Tke Ruffed G

rouse

Marine Biological Laboratory

Accession Nn 60794

Given Rv The Macmillan Co.

i%'' York City

Place ^

Tke Ruffed G

roti5e

^^

.L lie JLVulIed vx

rouse

ITS LIFE STORY, ECOLOGY AND MANAGEMENT

By frank G. EDMINSTER

THE MACMILLAN COMPANY • NEW YORK • 1947

Copyright, 1947, by
THE MACMILLAN COMPANY

All rights reserved — no part of this book may
be reproduced in any form without permission
in writing from the publisher, except by a re-
viewer who wishes to quote brief passages in
connection with a review written for inclusion
in magazine or newspaper.

First Printing

PRINTED IN THH UNITED STATES OF AMERICA

To my sons
DAVE and STEVE

to whom I
consider my greatest
duty is to help them
become good sportsmen

OGICyd^

Acknowledgments

I wish to express my great indebtedness to the New York State
Conservation Department, and particularly to (former) Commis-
sioner Lithgow Osborne and Director of Fish and Game, WilHam
C. Adams, for their sympathetic encouragement in this work. Full
credit for data used from the New York studies rests with the Con-
ser\^ation Department. All New York records cited are derived from
the Conservation Department's Annual Reports, unless otherwise
noted.

To my former colleagues I express my unbounded gratitude for
many years of stimulating and productive collaboration in "roughing
it with the ruffed grouse." Those whose association with me has
extended over most of the past decade or more and to whom I am
most grateful are Gardiner Bump, Robert Darrow, Walter Crissey,
Victor Skiff, Ben Bradley, Robert Cameron, Arthur Holweg and Asa
Smith.

For writing the greater part of the chapter on grouse management
that deals with the propagation of woody plants, I am indebted to
Robert Thornton, formerly of the U. S. Soil Conservation Service.

Credit is due the U. S. Soil Conservation Service, Dr. A. A. Allen,
the Pennsylvania Game Commission, and others for the use of nu-
merous photographs and illustrations.

For critical reading of portions of the manuscript and invaluable
suggestions for its improvement, I am grateful to Dr. Paul L. Erring-
ton of Iowa State College; Dr. William J. Hamilton, Jr., Prof. Ralph
S. Hosmer, Dr. Philip Levine, and Dr. George M. Sutton, all of
Cornell University; Professor Ralph T. King of Syracuse University;
Arthur C. Mclntyre and Dr. William R. Van Dersal of the U. S. Soil
Conservation Service; Dr. Neil Hosley of University of Connecticut,
College of Agriculture; Richard Gerstell and James N. Morton of
the Pennsylvania Game Commission; Dr. James L. Peters of the
Museum of Comparative Zoology, Cambridge, Mass.; and John E.
Trainer of Muhlenberg College.

vil

viii Acknowledgments

The pen-and-ink drawings at the chapter endings are by William
Montagna of Cornell University.

To W. L. McAtee of the U. S. Fish and Wildlife Service, I am most
grateful for careful editing of the whole work.

I have left for the last my deepest obligation. I want to thank
Dr. Arthur A. Allen of Cornell University for his wise and kindly
guidance in my chosen field for many, many years, and for his help-,
ful suggestions on the organization and writing of this book.

Introduction

Three times since the turn of the century the ruffed grouse has
suffered severe and quite sudden reduction in numbers in the north-
eastern states. Similar declines occurred in the lake states and
Canada. Each time it has appeared to be threatened with extinction.
Each time the sporting and nature-loving public has become aroused.
And following each of these occasions, there have been investiga-
tions of the causes of the catastrophe. Woodruff in 1908 reported
promptl}' on the severe but rather short-lived decline of 1907 in
New York. Stoddart likewise pursued his questionnaire study of the
1917 scarcity promptly and published the results the next year.
When the great decline of 1927 occurred (possibly the most severe
of all ) , studies were already under way by Allen and Gross in New
York and New England respectively. By the autumn of 1929, with
two more hunting seasons passed and no apparent improvement in
the numbers of grouse in New York, sportsmen urged tliat the state
again undertake to determine the cause of, and cure for, the periodic
decimation of the ruffed grouse. As a result the Conservation De-
partment undertook a complete, scientifically conducted study of
this species. I worked on this study for over seven years, mostly at
the Connecticut Hill Game Refuge, near Ithaca, N. Y. These studies
are the nucleus of experience about which this book is written, but
insofar as possible its application has been broadened to cover the
whole northeastern region. Eight years' additional work on wild life
throughout the northeastern states with the Soil Conservation Serv-
ice, U. S. Department of Agriculture, has aided in providing the
breadth of field observation that makes this possible.

Many other studies of the ruffed grouse have been carried on in
recent years. I have cited freely from the reports of these in order to
provide a comprehensive picture of this most fascinating bird. State
game agencies and colleges in Maine, New Hampshire, Massachu-
setts, Comiecticut, Pennsylvania, Virginia, Ohio, Michigan, Wis-

iz

X Introduction

consin and Minnesota are among those reporting on various aspects
of the subject.

It is not intended that this book be an exhaustive treatise on all
phases of study of this species. Rather it is hoped to cover the high
spots of the life story, the ecology, and management of the ruffed
grouse in a manner interesting to the sportsman, the nature lover and
the lay reader, and at the same time be accurate and adequately
documented for the technical man in the field of wild life manage-
ment.

I want to emphasize that the responsibility for this writing is
entiiely mine and it is not sponsored by any public agency. In par-
ticular, it should be understood that this is not a report of the New
York State ruffed grouse investigation. Throughout, where the edi-
torial we is used, it refers only to the author.

Preface

The ruffed grouse is a creature that has evolved over milHons of
years along with thousands of contemporaries in the animal and
plant world of today. The fact that the species exists at all is evi-
dence of its ability to compete successfully in a world of constant
strife. Like all other creatures it developed certain innate reproduc-
tive abilities, together with skills of escape and concealment that
enabled it to increase and maintain its numbers. Also, other condi-
tions developed to offset these increases. These opposing forces that
play on each species tend to preserve the status of each. In discussing
this mechanics of ecology for game species these forces have come to
be called the factors of abundance.

These elements, or factors, that control the numbers of a species
are essentially the same for all. But the different factors assume
vastly different degrees of importance with different species and
under varying circumstances with each species. Under any set of
conditions, some one of the factors will keep grouse from increasing
beyond the level it actually does; or, stated another way, will reduce
grouse to the minimum numbers they decline to prior to the breed-
ing season. This limiting factor does not cause all the mortality;
merely the ultimate part. But if the limiting factor is reduced or
eliminated, some other will take its place at a somewhat improved
level of population. There will always be some factor to set the
actual limit of survival for each species. And other factors, no matter
what their importance or how man may affect them, cannot change
the survival level of our game species. The limiting factor deter-
mines the greatest number that will survive on a particular area.

The eight major factors that control grouse populations may be
divided into two types, as shown graphically in Fig. 1, page 3. The
ones that compose the environment are in one group, while those
that are inherently a part of the species are in the other. These we
will describe briefly.

xi

xii Preface

SHELTER

Fundamental to the environment that supports grouse is the
vegetation that provides its home. Shelter includes the trees, shrubs,
and herbs that compose the crown cover, imdergrowth, and ground
cover as well as the surface litter on the ground, and occasionally
boulders, ground holes, and other protective materials. For the
grouse, the evergreen conifers and broad-leaved trees and shrubs
are important shelter, especially in the winter. Brush piles, fallen
tree tops, and deep snow also furnish important winter protection.
In the summer the leaf litter and vine and brier tangles are important
shelter. Shelter must furnish not only protection from the elements
and from mortal enemies, but the medium for all of life's functions
including nesting, coiui;ing, sunning and dusting.

FOOD

Probably the most obvious factor affecting abundance is the food
supply— an ample supply for good health at all seasons of the year.
Water and grit requirements are a part of this factor. With the
grouse, food as an independent factor is not often as limiting in
effect as it is with many animals. Cover which furnishes adequate
shelter usually possesses considerable numbers of grouse food plants.
But the distribution and quality of grouse foods has a vital bearing
on the distribution of the birds, even though grouse rarely succumb
directly from diet deficiencies.

The cover which furnishes both shelter and food for grouse is
termed the habitat, and is the foundation of the environment.

PREDATORS AND OTHER ANIMALS

Those creatures other than man that utilize the grouse for
food are its predators. Primarily mammals and birds, they are the
most common immediate cause of death for adult grouse. However,
in considering the importance of the predators and other animals in
determining grouse abundance, the correlation with other factors
must also be considered. Most of the other factors of abundance are
closely associated with predation. The snow that makes the roosting
grouse vulnerable for attack, the parasite that weakens its flight,

Preface xiii

the innate urge that forces a grouse to expose itself in order to
perform for the benefit of other grouse— these and many other
conditions predispose the predation which actually disposes of tlie
grouse. The lack of adequate shelter in the cover, an inconvenient
distribution of staple food supplies, man's sudden changes in the
habitat, are examples of environmental deficiencies that may induce
grouse mortality from predator attack. The density of the grouse
population itself affects the degree of predation.

Even under the best of physical circumstances, predators will
affect grouse population to some degree, and over most of the north-
eastern range they constitute one of the major factors. Many other
mammals and birds that are associated with grouse habitat tend to
serve as buffers for grouse by furnishing the predators with the bulk
of their food.

DISEASES AND PARASITES

Organisms that derive both food and shelter from the grouse are
parasitic upon it. Those that live upon its skin are ectoparasites and
belong to the insects and other arthropoda. Endoparasites, those that
live inside the body, are generally roundworms, flatworms, or free-
floating protozoa or bacteria. Some of the disease-causing organisms
are classed as viruses, while one is a fungus. Many are capable of
causing death, either alone or in combinations.

Most animals have a rather large population of parasitic organisms,
even when in excellent health. This is true of grouse. Most specimens
of grouse when thoroughly examined will be found to have parasites
of various types. Ordinarily they do the bird no harm. When certain
destructive kinds gain access, the bird may be seriously affected.
Other kinds of parasites, when present in large numbers, may have
serious effects.

Generally a grouse, so ill from the effects of parasitic organisms
that its functions are impaired, is picked off by some enemy before it
can die from its affliction. Some do actually die without violence,
at times in such numbers that the presence of an epidemic is in-
dicated.

When disease is rampant it is a grave matter. Fortunately this is
not commonly the case, and the direct losses from this source are
generally occasional and scattered

XXV

Preface

WEATHER CONDITIONS

Violent changes in the physical conditions surrounding grouse
are exceedingly important in affecting abundance. Weather condi-
tions and some of man's activities come in this category.

As with ourselves, weather conditions affect the grouse's activities.
The wind, rain, snow, sleet, heat, cold, and sunshine all play a part in
determining where a grouse will be and what it will be doing. For
the young ones, the exposure to the elements may cause death,
quickly and surely. To the grownups it often makes them more
vulnerable to their enemies than they might be under more favorable
weather. For tlie grouse, as for most living things, the weather is of
the utmost importance in their lives.

man's activities

Man may be either beneficial or destructive to grouse. As a hunter
he is a killing agent, yet the removal of the surplus population may
actually be beneficial to the species he kills. In other ways he affects
the physical conditions of the environment. As a farmer he both
improves and destroys habitats, through plowing, cutting, and
fencing; he constantly arrests and alters the natural plant succession.
His use of fire is often detrimental to wild life, yet sometimes help-
ful. His livestock and pets affect both plants and animals of the wild.

Man as a lumberman creates desirable openings and bushy growth
by breaking up the forest cover. But by making forests into open
land, he has often completely destroyed the range for grouse. As a
conservationist he recreates habitat, offers protection in refuges, and
makes laws to improve wild-lffe conditions. With some definite
limitations, it may be said that the future of the ruffed grouse, as
with all wild Iffe, rests with him.

HABITS OF THE SPECIES

Even as with you and me, the grouse has habits imbued in him
that he could do better without, at least in some respects. Many
inherent urges affect survival. With us it may be to drive automobiles
at breakneck speed, to jaywalk in tra£Bc, or to eat or drink exces-
sively. With the grouse it is to drum on an exposed log, to strut

Preface

XV

around in the open to please a lady, or to fly at breakneck speed
where obstacles are everywhere. But even though it is courting
death to indulge in these whims, the grouse will continue to do it.

REPRODUCTIVE CAPACITY

The second of the major survival and increase factors that is
inherently a part of each species is its innate ability to reproduce.
It is elemental that, other things being equal, a bird that habitually
lays eleven eggs ( like the grouse ) has a higher reproductive capacity
than one that lays, let us say, four eggs (like the woodcock). Like-
wise, a species that is polygamous is better fitted to assure complete
fertility than is one that is monogamous; a creature that matures to
breed in a single year has a higher reproductive capacity in that
respect than one that cannot breed until it is two or three years old.
So in these and in other reproductive constituents, the survival and
population of the grouse is vitally affected.

When we combine all the forces involved in these eight major
factors of abundance, we have the whole ecology of the grouse and
its associates with all the tragedy, pathos, drama, and comedy of
their dynamic existences.

Table of Contents

PAGE
INTRODUCnON ix

PREFACE Xi

Part 1— Its Life Story

Chapter 1.

the bird itself

1

Classification, Nomenclature

1

Description

Pterylography

References and Citation Sources

2

9

17

Chapter 2.

chronicle of the ruffed grouse in east-

ern UNl'i'ED states

19

References and Citation Sources

26

Chapter 3.

biography

27

References and Citation Sources

58

Part 2— Ecology of the Rxjffed Grouse

Chapter 4. shelter 60

Types of Grouse Range 60
Cover Types— Characteristics and Composi-
tion 63
Interspersion of Cover Types 70
Plant Succession Changes Cover Types 71
Relation of Shelter to the Grouse Nesting 73
Relation of Shelter to Grouse Broods 77
Changes in Cover Types Utilized at Different
Times of Day 79
xvii

60794

XVlll

Table of Contents

Chapter 5.

PAGE

The Effect of Slope on Selection of Cover

Type 80

Cover Selection in Relation to Openings 80

Cover Types Chosen as Affected by Weather

Conditions 80

Relation of Shelter to Adult Crouse 81

The Effect of Openings on Cover Selection 91
Variation in Cover Type Preference at Dff-

ferent Times of Day 92

The Effect of Slope on Selection of Cover by

Adult Grouse 93

Cover Types Chosen as Affected by Weather

Conditions

94

References and Citation Sources

99

)OD AND WATER

101

Winter Foods

105

Spring Foods

105

Summer Foods

107

Fall Foods

110

Summary of Foods Utilized

111

The 25 Primary Grouse Foods in the North-

east

113

Secondary Grouse Foods

138

Quality of Different Grouse Foods

171

Variation in Summer Food Habits of Yoimg

and Adults

172

Relation of Food Supply to Cover Types

173

Distribution of Birds as Affected by Food

Supply

174

Effects of Cultural Operations on the Food

Supply

174

Food as a Limiting Factor

176

Economic Aspects of Grouse Food Habits

179

Use of Grit

180

Water Requirements of Grouse

182

References and Citation Sources

182

Table of Contents

XIX

PAGE

Chapter 6. weather conditions in relation to grouse 184
Effects of Weather on the Habits and Num-
bers of Grouse 184
Population Fluctuations Affected by Weather

Conditions 190

References and Citation Sources 194

Chapter 7. interrelationships of ruffed grouse to

mammals and to other birds 196

Species that Prey Upon Grouse

Relation of Predation to Cover Types

Predators as an Agent of Sanitation

The Effect of Predator Control on Ruffed

Grouse Populations
The Abundance of "Buffer" Species as It Af-
fects Predator Food Habits and Grouse
Mortality
Other Animals that Affect the Grouse Wel-
fare
Predation as a Limiting Factor
References and Citation Sources

196
209
210

211

215

219
219
221

Chapter 8. the diseases and parasites of wild ruffed
grouse

Characteristics and Occurrence of Disease

Agents
Total Parasite Infestation
Disease as a Limiting Factor
References and Citation Sources

223

225
249
250
252

Chapter 9. man's relation to the grouse

As a Hunter and Trapper
As a Farmer and Lumberman
As a Conservationist
Man as a Limiting Factor
References and Citation Sources

254

255
267
271

275
275

XX Table of Contents

PAGE

Chapter 10. productivity and populations 277

Potential and Actual Productivity 277

Annual Fluctuations in Grouse Numbers 278

Life Equation 284

Variation in Different Habits 286
Characteristics Inherent in the Species that

Condition Productivity 286

Populations in Relation to Productivity 292

Carrying Capacity and Saturation Point 298

Environment Affects Productivity 300

Summary of Analysis of Periodic Fluctuations 322

References and Citation Sources 323

Part 3— Management of the Ruffed Grouse

Chapter 11. management of the ruffed grouse 326

Extensive vs. Intensive Management 327
Development of Habitat 329
Woodland Protection 329
Improvement of Woodland Cover 331
Interplanting of Conifers 336
Improvement of Shelter by Cuttings 337
Improvement of Food by Cuttings 339
Establishing Woodland Shrub Borders 340
Use of Poison in Establishing ShiTib Borders 341
Shrubs Along the Woods Road 342
Clearings for Food and Cover Improvement 343
Selective Cutting to Improve Food Condi-
tions in the Whole Woods 345
Provision of Drumming and Dusting Places 347
Plantings and Seedings 348
Propagation of Woody Plants for Plantations 356
Control of the Harvest 364
References and Citation Sources 372

index 375

List of Illustrations

Figure Page

1. The twelve grouse shown represent parents and a brood of
grouse 3

2. Feather tracts and spaces of ruffed grouse 13

3. Type of cover preferred by grouse. Taken from Analysis of Cer-
tain Cover Requirements of the Ruffed Grouse in New York
State. G. Bump, 1938. 82

4. Intensity of use of cover types by adult grouse by seasons 84

5. Cover types used by adult grouse at different seasons 87

6. Seasonal utilization of food by adult ruffed grouse according to
type of food (Northeastern States) 101

7. Utilization of food by young ruffed grouse according to types

of food ( Northeastern States ) 102

8. Twenty-five most important food plants used by the ruffed
grouse in the northeast 112

9. New York State Game Kill Records 1930-1938 217

10. Generalized Population Trends of Ruffed Grouse, certain Preda-
tors, and certain Rodents, New York State 1930-1939 (Based

on State Game Kill Records and field data) 218

11. Reported take of ruffed grouse by hunters 256

12. Correlation of grouse recovery rate with breeding density, Con-
necticut Hill, N. Y., 1930-1937 283

13. Life equation of grouse for an increasing population 285

14. Life equation of grouse for a stable population 285

15. Normal grouse infant mortality curve June to September 304

16. Hypothetical imit of grouse cover 333

17. Plan for cover type changes in a woodland 335

Plate

1. (Upper). When the white man came, farms were hewn out of
the wilderness. (Lower). Where the land clearance work was
confined to small scattered units, the range was improved for
grouse.

2. (Upper left). The drumming of the cock grouse is one of the
wonders of the bird world. (Upper right). Strutting before the
female bird. (Lower). Young grouse in a brood dust bath.

xxii List of Illustrations

Plate

3. ( Upper left) . The nest was built at the base of a big tree, on the
sunny side. (Upper right). Eleven eggs composed the clutch,
an average number. (Lower left). The hen grouse watches
carefully for enemies before stepping onto the nest. (Lower
right). The chicks remain in the nest only a few hours after
hatching.

4. (Upper left). Grouse chick one day old. (Upper right). At five
days of age. (Lower left). Seventeen days have passed, (Lower
right). At five weeks of age.

5. (Upper left). Grouse going to roost in a tree. (Upper right).
Grouse just after landing in the snow. (Lower left). Pheasant
and grouse eggs in one nest. ( Lower right ) . A snow roost after
being vacated.

6. (Upper). Disconnected cover, woodlands separated by open
fields. (Lower). Continuous cover, extensive forests found in
the several mountainous areas from the Appalachians to the
Adirondacks and Maine.

7. An aerial view of a portion of the Connecticut Hill sub-marginal
farming area.

8. (Upper left). Open land, overgrown land and hardwood woods
in close juxtaposition. (Upper right). Shrub cover next to conif-
erous woods. (Lower left). Woodlands lacking a shrubby edge
lose valuable interspersion. ( Lower right) . Pure alder most often
occurs on moist soils, and is much used by broods.

9. ( Upper left) . Subtype of mixed hardwoods composition. ( Upper
right). Mixed conifer-hardwood subtype. (Lower left). Mixed
oak composition. (Lower right). Pure stand of popple along
woods border.

10. (Upper). Newly cut subtype. (Lower). Old cuttings.

11. (Upper). Pure hardwoods are deficient in shelter. A scattering
of conifers improves this type for grouse. (Lower). Broadleaved
evergreens serve somewhat the same purpose as conifers.

12. (Upper). Small openings made by removing a large tree, or
two or three in hardwood stands is the best of the many sub-
types. (Lower). Tlie mixed oak association is quite unproduc-
tive of grouse where it occurs in large, unbroken areas.

13. (Upper). Hardwood overstory of near-mature trees with much
hemlock in lower growth. (Lower). Young stand of hemlock
and hardwoods.

14. (Upper). Overstory of mature white pine with completely
hardwood understory. (Lower). Hard pines (Virginia pine)
and oaks compose a mixed woodland cover type.

15. (Upper). Hemlock on left has sheltering branches close to
ground, white pine on right has shelter only in crown. (Lower).

List of Illustrations xxiii

Plate

Close-up of low-hanging hemlock boughs showing their provi-
sion of winter protection.

16. (Upper left). Some of the broad-leaved evergreens may sub-
stitute for the conifers in furnishing winter shelter close to the
ground. (Upper right). The hard pines are the predominant
conifers in much of the Appalachian range. (Lower). Spruces
furnish most of the winter shelter in the northernmost range.

17. (Upper). An old white pine has established the makings of a
stand of sohd pine around it. (Lower left). A view of the nortli
end of the Connecticut Hill area, showing how natural plant
succession of hardwoods with scattered conifers is gradually
changing the open fields, first to brush overgrown land, ulti-
mately to forest. ( Lower right ) . Among the factors that influence
plant succession is the nearness to an existing woodland or
hedgerow.

18. (Upper left). Hen grouse on nest at base of a large beech.
(Upper right). A successful nest showing the characteristic
appearance of hatched eggs. (Lower left). Grouse incubating
on nest at base of stump. (Lower right). Mother grouse brood-
ing her chicks.

19. (Upper). A single grouse meal. Crop contents of a female
grouse from Sulhvan County, Pennsylvania, December 13, 1940.
(Lower). A grouse with a full crop, skinned out to show the
immense capacity of this organ when fully distended.

20. (Upper left). Ferns, especially the woods fern, the fronds of
which are eaten mostly from autumn to mid-spring. (Upper
right). Sedges (Carex) are used from spring to fall. (Lower
left). Important among winter foods of grouse are tree buds.
(Lower right). Sheep sorrel (upper plant), a small field weed.
The Canada mayflower (lower plant), a plant of the woodland
floor.

21. (Upper left). Brambles. (Upper right). Blueberries. (Lower).
The partridgeberry provides food all year round.

22. (Upper). Acorns. (Lower left). Cherries. (Lower right). The
dogwoods.

23. (Upper left). The mapleleaf viburnum. (Upper right). The
sawbrier. (Lower left). Flowering dogwood. (Lower right).
Red osier.

24. (Upper left). Old apple orchards furnish excellent feeding
cover all year. (Upper right). Wild apples near pine clumps are
a perfect combination in autumn. (Lower left). The beech
furnishes a preferred food when it has a nut crop. (Lower
right). The sumacs are an important source of winter food.
(R. typhina shown in center).

xxiv List of Illustrations

Plate

25. (Upper left). Mountain laurel, whose leaves are a staple food.
(Upper right). Highbush cranberry, one of the viburnums.
( Lower left ) . Wild rose, whose hips are eaten when accessible.
(Lower right). Teaberry, or wintergreen, is a small plant of the
woodland floor.

26. (Upper). Wild black cherry. (Lower). Pin cherry; and choke-
cherry.

27. (Upper left). Nannyberry. (Upper right). Hobblebush. (Lower
left). Blackhaw. (Lower right). Arrowwood.

28. (Upper). Prolific growth of wild grape {Vitis labrusca) along
a woodland edge. (Lower). Close-up of fruit.

29. (Upper left). Black chokeberry, a small bush. (Upper right).
Bittersweet, a cHmbing vine. (Lower). Winterberry, one of the
hollies.

30. (Upper left). Virginia creeper, vine. (Upper right). Bayberry,
most prevalent near the coast. (Lower left). Elder, a common
fencerow shrub. (Lower right). Mountain ash, a small wood-
land tree. ( Center ) . Serviceberry, another woodland tree whose
fruits ripen in early summer.

3L (Upper). Farming activities greatly affect the grouse food sup-
ply. (Lower). When deep snows blanket the ground, grouse
resort to buds for most of their food.

32. ( Upper ) . Feathers and tracks in the snow reveal fox evidence.
(Lower). Horned owl work.

33. (Upper). Grouse nest destroyed by a horned owl killing setting
hen, after which a fox ate the eggs. (Lower left). The common
skunk loves eggs, and occasionally stumbles onto a grouse nest.
(Lower right). Great homed owl, one of the most efficient of
all predators.

34. (Upper left). The cottontail, found mainly in the disconnected
cover areas of grouse range. (Upper right). The snowshoe hare,
found mainly in the extensive northern forests. (Insert). The
squirrels affect the grouse in two ways, as a buffer between the
grouse and its enemies, and as a food competitor. (Lower). The
empty shells of a walnut planting after being dug up and eaten
by squirrels.

35. (Upper). Diagram of alimentary canal of the ruffed grouse
showing location of some parasite infestations. (Lower left).
Grouse stomachs, normal and parasitized with Dispharynx
spiralis. (Lower right). Stomach worms (Dispharynx spiralis)
and intestinal worm (Ascaridia lineata).

36. (Upper left). The modem hunter pursues grouse as a sport.
(Upper right). A split-second shot brings a clean kill. (Lower).
Man's use of well-trained pointing dogs has done as much as
anything to make hunting a fine sport.

List of Illustrations xxv

Plate

37. Ploughing and cultivation of crop fields prevent woody cover
from expanding, and at the same time maintain the valuable
woodland edges next to the fields.

38. ( Upper) . Man's interest in managing the land to produce grouse
is often directly affected by what he does or does not do to
control other animals, such as deer. (Lower). Damage to white
pine seedling from deer browsing.

39. (Upper). Overcutting destroys shelter, eliminates much of the
interspersion of types, and prevents high quahty regeneration
by eroding the forest floor. (Lower). Woods roads break up the
cover, make valuable edges, feeding lanes, and, in summer, dust-
ing and sunning spots.

40. (Upper). Seed stock refuges as a medium of hunting control
are of httle value in grouse management except in the most in-
tensely shot areas. ( Lower ) . The woodcock, shown on its nest,
is one of the more important game birds commonly sharing
range with the ruffed grouse.

41. (Upper). Protection from grazing is one of the first essentials
of woodland management, whether for game or timber. ( Lower
left). Overgrazing in a woodland not only prevents plant re-
production, but has a detrimental effect on the soil itself.
(Lower right). Fire in woodlands may destroy grouse cover
almost completely for a period.

42. (Upper). Farm woodlands that have been heavily grazed by
cattle can often be interplanted successfully. ( Lower left ) . Nor-
way spruce interplanted in a woodland glade. (Lower right).
An old field partially taken over by naturally-seeded white pine.
Interplanting of other species could well be done here.

43. (Upper). Brush piles made from the waste tops and branches
of cuttings serve well as temporary winter shelter. (Lower). The
margin of the woodland lane is a good place to improve food
conditions.

44. (Upper). Interplanting of white pine in an area of cut-over
hardwoods improves the shelter value. (Lower left). Shrub
border of silky dogwood planted along a woodland edge. (Lower
right). Shrub borders along the woodland edge may be devel-
oped by cutting out tree species and favoring the shrubs in the
woods margin.

45. (Upper). A quarter-acre clearing just after cutting shows
stumps, and sunshine reaching the ground. (Lower). Same after
two years, showing thicket development of shrubs, and tree
sprouts and seedfings.

46. (Upper). Aerial view of a Connecticut Hill woodland showing
both unit and lane cuttings. (Lower left). Close-up of a one-
acre clear-cutting grown to briers, elderberry and other shrubs

xxvi List of Illustrations

PUae

and trees. (Lower right). A lane slashing, thirty feet wide, pro-
viding needed brush cover and valuable edges.

47. (Upper). Clear-cut areas provide edges, so needed by game.
(Lower left). An experimental slashing unit created by poison-
ing the trees, leaving the dead trees standing, (Lower right).
The edge of a unit slashing showing the change in the character
of the cover.

48. (Upper). Contour-furrowed field being planted. (Lower). The
same plantation five years later.

49. ( Upper ) . In a ten-year-old plantation, the arborvitae, while sur-
viving, has grown but a very few inches due to an unsatisfactory
site. (Lower). The second row in the shrub border planting is a
failure because of the use of a species unsuited to the soil.

50. (Upper left). An aerial view of two fields on Connecticut Hill
showing pattern of conifer and hardwood plantings. (Upper
right and lower left). Same two fields from the ground. (Lower
right). Field planting showing alternate bands of conifers and
hardwoods.

51. (Upper). For the first few years young trees affect the old
field complex very little. (Center). From about five to fifteen
years of age the stand is mixed with natural reproduction, usu-
ally hardwoods, and serves as overgrown land. (Lower). After
the crown closes, often between fifteen and twenty-five years of
age, the competing natural vegetation is rapidly driven out.

52. (Upper left). The trees gradually prune themselves and the
tightly closed canopy prevents any germination of ground cover.
(Upper right). At around the forty-to-fifty -year age period, the
crown begins to open and the return of herbs and hardwoods
begins on the woodland floor. (Lower). At maturity we find a
top crown of pine, partially broken, with a complete hardwood-
shrub-herb stand underneath.

53. (Upper). Mixed planting of conifers and hardwoods to provide
both food and cover. ( Lower ) . Border of shrubs provides a de-
sirable thicket type next to a pine planting.

54. A ten-year-old planting of northern red oak.

55. Nursery beds of seedling bayberry, one of the shrubs recom-
mended for planting.

56. (Upper). Game refuges are generally associated with pub-
he shooting ground areas when established on public lands.
(Lower). Trapping fur-bearers such as the red fox for their
pelts on a sustained-yield basis is the only predator control
recommended on grouse range.

Tke RuffeJ G

rou5e

1

The Bird Itself

CLASSIFICATION, NOMENCLATURE

Bonasa umhellus (Linnaeus)/ the ruffed grouse, belongs to the
great, almost cosmopolitan order Galliformes. Four families of this
order are found in North America, the Cracidae, Tetraonidae,
Phasianidae, and Meleagrididae— respectively the curassows, guans,
and chachalacas; the grouse and ptarmigan; the pheasants and New
World quail; and the turkeys. The family to which the ruffed grouse
belongs, the Tetraonidae, is represented by seven genera and eleven
species on the North American continent.

The generic name Bonasa, derived from the Greek Bovayn^ and
Latin Bonasus (classical nouns meaning "a bison") embodies the
idea of a bison's bellowing, hence that of a ruffed grouse's drum-
ming. The zoologist Stevens, who named the genus in Shaw's Gen-
eral Zoology in 1810, may never have seen a living ruffed grouse, but
he obviously had heard of its courtship performance. The specific
name, umhellus, is a Latin word meaning an umbel or umbrella.
It is descriptive of the bird's striking ruffs. The genus Bonasa is
found only in North America (in the United States and Canada)
and has but one species.

The accepted common name is ruffed grouse. However, the bird is
known colloquially by many other names, some of which occasion
considerable confusion with other species. The various Indian tribes
called it by many names, including: Wen'-gi-da-bi-ne' or Pinai
(Chippewa); paupock ( Narragansett ) ; papahcogh (Mohican); cut-
quass ( Pequot ) ; ohquase ( Oneida ) ; pabhackoo or mimituiis ( Dela-
ware); Mutch-i-es'— "Bad Bird" because in old times he was mede-
oulin, had magic power, and plotted against Gluskap, according to
old stories— (Malecite, in New Brunswick); Pul-o-wetch (Micmac);
Ajack, Puskee, or Pupuskee (Algonquian around Hudson Bay); Kh-

^ A.O.U. Check List No. 300.

2 The Ruffed Grouse

tuk ( Esquimo ) .' Probably the first name given it by the early white
men in New England was "wood hen." Today it is commonly called
"partridge," "patridge," or "birch partridge" in New England, al-
though it is not actually a partridge at all. In the southern Appa-
lachians it is referred to as the "pheasant," but it is not a true pheas-
ant either." Other names include "mountain pheasant," "fool hen,"
"drumming pheasant," "wood-pilequarker," "drumming grouse,"
"drumming partridge," "drummer," "shoulder-knot grouse," "ruffed
heath-cock," "moor fowl," "tippet grouse," "carpenter-bird," "white-
flesher," "hazel hen," "red-tail," "gray-tail," "silver-tail," and just plain
"grouse."

DESCRIPTION

Adults. The grouse is a symphony of rich browns, sometimes with
a grayish tendency, sometimes rufous, with plump form, medium-
proportioned beak, legs and tail, and a neck ruff of varying promi-
nence. Length varies from fifteen to nineteen inches, wing spread
twenty-two to twenty-five, tail four and one-haff to seven and one-
haff, and weight from sixteen to twenty-eight ounces.

The head is topped with erectile feathers that are raised into a
crest when the bird is excited. The crown is crossed with numerous
lines or dabs of black and buff that gives the brown ground color a
mottled appearance. There is a quite prominent buff eye-line from
the base of the beak to the back of the head and a buff chin patch.
The ruff feathers, which form a somewhat triangular patch on each
side of the neck, are generally black with a green and purple irides-
cence. Occasionally on a very reddish bird the ruff is a rich reddish
brown. The ruff is carried obscurely beneath the brownish neck
feathers until erected into prominence by some excitement.

The browns of the neck, back, upper wings, and rump have vary-
ing patterns composed by the different characters of black, buff, and
white. The neck is rather prominently marked with whitish, the back
and rump mottled with long, black-edged spots along the feather

1 1 am indebted to W. L. McAtee of the U. S. Fish and Wildlife Service for most
of the Indian names.

2 The "true partridges" and the pheasants belong to the family Phasianidae, natives
of the Old World. The European ("Hungarian") Partridge (Perdix perdix) and the
ring-necked pheasant (Phasianus colchicus torquatus) are well established in northern
U. S. and parts of Canada.

The Bird Itself 3

shafts; the rump is somewhat more rufous than the back; the wings
usually show less rufous in the browns than the other upper parts,
the primaries are quite dark with buff to whitish markings along the

Fig. 1. The twelve grouse shown represent parents and a brood of grouse. We
assume that the parents survived the vraiter on an area that has a carrying ca-
pacity of but two grouse. Therefore ten of this group must die by next spring if
they remain in the area vmder the same conditions. The birds are illustrated as
meeting the elements that may reduce their numbers. The higher the element,
the greater the toll it may possibly take. The highest element, here shown as
shelter, is the hmiting factor. Only two grouse, in this instance one youngster
and one parent, are able to surmount it. The reduction or eUmination of any of
the other seven lower elements would not result in a greater survival but merely
a delayed mortaHty, since the shelter will permit the maintenance of only two
grouse. Under difFerent circumstances the eight factors might assume any order
of importance, and any one might be the limiting factor.

narrow outer vanes; and the secondaries and coverts are bordered
with buff or whitish with occasional black spotting.

The ground color of the eighteen^ tail feathers (rectrices) is a

^A few specimens having twenty rectrices have been examined, usually males,

and a few with only sixteen, usually females. ''^l C/ /

4 The Ruffed Grouse

clear rufous brown transversed by a series of from six to eleven thin
black bars. The tip of the tail is gray, but there is a broad black
subterminal band that shows clearly in flight. The band is often
broken widi gray on the center feathers, especially in the females.

The lower parts vary from whitish to gi'ayish or bulfy white, tend-
ing mostly toward buff on the lower neck and upper breast. Numer-
ous short bars of brown or blackish, daikest along the sides and fad-
ing toward the center, break up the gray background. The under
wings and under tail coverts are predominantly an even gray or
buff.

The beak is dark brown, the iris hazel, and the feet dark horn
color. The tarsi are spaisely feathered above, naked below. Dirring
the winter months lateral pectinations appear on the toes, commonly
and aptly known as "snowshoes." When spring comes these ap-
pendages are shed.

Plumage Differences Between the Sexes. The full-grown male is
consistently larger and heavier than tlie full-grown female, although
large females may match small males in these respects. The rectrices
of the male are from sLx to seven and one-haff inches long, while
those of the female are from four and one-half to sLx and one-quarter,
thus giving tlie male a distinctly longer tail. The ruff is relatively
prominent on the male as compared with the female, always con-
tinuous across the breast where that of the female fades. The broad,
black subterminal tail band is always broken with gray in the fe-
male, but is usuallv continuous in the male. The colors of the female
are generally more subdued than those of the male and there is
usually a slight pinkish wash to the brown of the forebreast. During
the breeding season the male has a bright reddish naked patch over
the eye that is not developed in the female.

With a bird in the hand, the sex is readily perceptible in almost
all cases because of one or more of these characters, even though
they are not each always constant. In the field the lengdi of tail is
a safe indicator most of the time, while niff and tail band will often
aid in making the sex identification. Sex identity in the wild may also
be aided by actions characteristic of males or females, such as broodi-
ness, drumming or strutting.

Color Phases. For most parts of its range, the grouse exhibits wide
variation in basic ground color. Those birds with a rufous cast are

The Bird Itself 5

called the "red" phase, while giayish birds are "gray" phase. Every
gradation between the extremes of dichromatism may be found.

Subspecies. Six subspecies are recognized by the 1931 American
Ornithologists' Union Check List. The originally named form B.
umbellus umbellus, has been described above. B. umbellus togata
(Linn.), the Canada ruffed gi'ouse, or northern ruffed grouse
(A.O.U. No. 300a), is essentially like B. u. mnhelhis, but seems to
average a little larger and heavier. A very large male will sometimes
scale over thiity ounces. While this subspecies has a red and gray
phase, it never attains the fullness of rufous or cinnamon color of
the more southern subspecies. In B. u. togata the tail is generally
giayer, the upper parts are a darker and more chestnut brown, and
the under parts are more heavily barred. The feathers on the tarsus
are somewhat longer than in B. u. umbelltis.

B. umbelltis thayeri (Bangs), the Nova Scotia ruffed grouse
(A.O.U. No. 300d), is similar to togata in being two-phased but is
darker gray, dusky or sooty, on the upper parts. Banding of under
parts is regular and prominently dusky and more sharply contrasting
against the background color. The bill is somewhat longer than in
B. u. umbelltis. The color phases are both very dark and not markedly
drfferent.

B. timhelltis umbeUoides (Douglas), the gray ruffed grouse
(A.O.U. No. 300b), is usually a gray-to-slatish colored bird with
very little brown on the upper parts, although red ruffs and brown
tails occur occasionally. From mid-back to the top of the tail it is
veiy slate gray, the arrow-head spots on the back feathers are
gray with black edging, and the tail delicately vemiiculated with
iDlack. Beneath it is whitish with a tawny caste. There are several
brownish crossbars on each feather, largest and most sharply de-
fined on the sides. The flank and vent feathers are nearly clear
white and the foreneck and scapulars are a varied mixture of grays,
browns, whites and rufous shades. The wing covert feathers gener-
ally have white shaft lines. The female is inclined to be more rufous
than the male. The feathers on the tarsus are long, sometimes extend-
ing to the base of the toes.

B. umbellus sabini (Douglas), the Oregon (or red) ruffed grouse
(A.O.U. No. 300c), is a large dark race. There is more blackish to
the brown and tlie bruwiis often are a marked chestnut shade. Ac-

6 The Ruffed Grouse

cording to Coues ( 1903 ) , this subspecies was discovered by Lewis
and Clark in 1805-06 and first named Tetrao fusca by Ord ( Guthrie's
Geography, 2nd Am. ed. ii, 1815, pp. 317). "In strictness . . . this
bird should be called B. u. fusca Coues," but he waives the point in
favor of B. u. sahini as redescribed by Douglas.

B. umbellus yukonensis (Grinnell), the Yukon ruffed grouse
(A.O.U. No. 300e), is the largest and grayest of the subspecies, simi-
lar to B. u. umhelloides but the light colors are more ashy and the
dark markings finer.

While the six subspecies just described are all that were recognized
ofiicially in the 1931 edition of the A.O.U. Check List of North Ameri-
can Birds, several others have been described since this check list
was printed.^ One, from Vancouver Island, named B. u. hrunnescens
by Conover (1935), has since been accepted for the A.O.U. Check
List (Auk Vol. 61, No. 3, July 1944). It is described as being most
closely related to sabini but with a browner (less reddish) upper
surface except for the tail. The tail is either a dull ochraceous umber
instead of ferruginous in the red phase, or a clear giay without red-
dish cast and without double crossbaning in the gray phase. It is
much darker brown on the upper parts than either umhelloides or yu-
konensis, and is more buffy below with more brown barring. It is
thought to be restricted to Vancouver Island and islands adjacent to
British Columbia and the mainland from Vancouver to Malaspina
Inlet.

Three additional subspecies are described by Todd (1940) as fol-
lows: B. u. monticola, Appalachian ruffed grouse, "similar to B. u. um-
bellus but general coloration darker; the under parts more regularly
and more heavily barred and more strongly suffused with buff."
This subspecies is ascribed to the Appalachian mountain region from
West Virginia southward. According to Wetmore (1941), birds
from this area are indistinguishable from B. u. togata.

The second new subspecies described by Todd is B. u. medianus,
Minnesota ruffed grouse, "Similar to B. u. umhelloides, but the upper
parts are less grayish, more rufescent, and the under parts are more
albescent and less heavily barred." Its range is given as the "Transi-
tion Zone from Alberta to southeastern Minnesota (and probably
farther east)."

Third of Todd's new subspecies is B. u. canescens, northern ruffed

1 A subspecies, called E. u. Johdi, was described in 1871 by Jaycox from birds
taken near Itliaca, N. Y. It was not accepted for the check list, however.

The Bird Itself 7

grouse. "Similar to B. u. togata, but the upper parts in general are
giayer, less brownish, and thus lighter in tone, particularly on the
secondaries, scapulars, rump, and upper tail-coverts. The under
parts are also lighter colored on the average, and the dark barring is
less intense. Similar to B. u. medianus, but the upper parts are gen-
erally darker, more grayish and less rufescent, and the tail is obvi-
ously darker gray; the buff of the under parts as a rule is not so
rich and deep. The general coloration is lighter than B. u. umbel-
loides." Its range is described as "from Labrador west to James Bay
and thence to eastern Manitoba."

While Todd is proposing the addition of three new subspecies
of Bonasa umbellus he suggests that the subspecies B. u. thayeri, ac-
cepted by the A.O.U. Check List as authentic, is actually identical
with B. u. togata. He also indicates his belief that B. u. brunnescens,
proposed by Conover, is adequately covered by B. u. sabini. In sum-
mary, Todd would recognize eight subspecies, although he says
frankly, "A number of problems concerning distribution and sys-
tematics are necessarily left unsettled."

This is apparently the case, for only a year later Bailey ( 1941 ) de-
scribes another subspecies from Long Island, N. Y. Proposed as B. m.
helmet, it is described: "A small ruffed grouse . . . very light buffy
throat and a light-colored upper mandible . . . more whitish belly
than any of the other races, heavily baiTcd below with blackish;
dark-breasted, but with more buffy breast than togata; neck ruffs
and subteiTninal tail band black . . . gray-tailed type with few ex-
ceptions."

Recently, a complete taxonomic revision of the species was pro-
posed by Aldrich and Friedmann (1943). They did not simplffy
the matter, but did provide sound ecological basis for their oganiza-
tion. Twelve subspecies are recognized, eight previously described
and four new ones. Three of those discussed above are discarded.
Those substantiated are: umbellus, togata, umbelloides, sabini, iju-
konensis, brunnescens, monficoh, medianus. Discarded are: thayeri,
recognized in the A.O.U. Check List but considered by these authors
to be the same as togata; canescens, described by Todd but here in-
cluded with umbelloides; and helmei, which is covered by the type
form B. u. umbellus. The four new subspecies described are cas-
taneus, affinis, phaios, and incanus. These are distinguished as
follows :

The Olympic ruffed grouse, B. u. castaneus, is the "darkest and

8 The Ruffed Grouse

most richly colored of all the predominantly brown races of the
species . . . darker than sabini and more reddish than brunnesceiis;
l^rown of upper parts deep chestnut to dark auburn with no grayish
mixture . . ." No gray phase is known. Its range is western Wash-
ington from Puget Sound to Oregon, south of the range of brun-
ncscens and north and west of sabini.

The Columbian ruffed grouse, B. u. affinis, is described in both
the gray and brown phases, the gray being the more prevalent. The
gray birds are in ". . . general appearance intermediate between
the gray phase of umbellus and that of sabini . . " Brown-phase
birds are "closest to the brown phase of togata, but have the black
markings less extensive." The range extends from east-central Ore-
gon and southwestern Idaho northward east of the Cascades through
most of eastern Washington and the interior of British Columbia,
to Hazleton.

The Idaho ruffed grouse, B. u. phaios, occurs in both color phases.
In both it is most similar to B. u. mnbelltis. The gray birds are
grayer, much darker, less brownish than the like phase of umbellus,
while brown-phase birds of pJmios are ". . . much darker, less
rufescent, more brownish (more like corresponding phase of brun-
nescens, but with more grayish or dusky ) . . ." than their umbellus
counterpart. It is found on the west slopes of the Rocky Mountains
in Idaho, and in portions of northeastern Oregon, southeastern
Washington, and northeastern Washington.

The hoary ruffed grouse, B. u. incanus, in its brown phase, is
"... a very ashy bird, similar not to brown but to gray phase of
B. u. umbellus, but paler and, except for tail, less brownish, more
like that of B. u. umbelloides but paler and less brownish on inter-
scapulars, back, and upper surface of wings . . ." The gray phase
is ". . . similar to brown phase but with tail feathers smoke gray
with no buffy tone; ventral barrings duskier— light brownish olive
darkening to sepia on the sides and flanks." It is found ". . . from
west-central and central-northern Utah, southeastern Idaho and
central- western Wyoming, northeastward across Wyoming and the
Dakotas to northeastern North Dakota . . ."

It is well to note that Aldrich and Friedmann indicate in many in-
stances that the subspecific characters gi-ade from one to another
along the boundaries of their ranges. Thus the delineation of the
ranges of subspecies is somewhat arbitrary and the number recog-

The Bird Itself 9

nized is dependent upon the degree of variation one requires for
such distinction.

I lay no claim to being an authority on the taxonomy of the ruffed
grouse. Even so, having examined many hundreds of specimens of
all ages and at all seasons from the northeastern states, I am thor-
oughly impressed with the imending variety in pattern and colora-
tion of this remarkable bird. I have also seen birds of different sub-
specific pattern in single broods. How we view the question of
subspecies depends upon whether we subscribe to the beliefs of the
"lumpers" or the "splitters." I go along with the lumpers, for the
sake of simplicity. My inclination is to go along with Todd in aban-
doning B. u. thayeri and B. u. hrunnescens. But I am equally un-
S}Tnpathetic to the eight new subspecies suggested by Todd, Bailey
and Aldrich and Friedmann. If the authorities reconsider the sub-
species of Bonasa umhellus for the next edition of the Check List,
I would cast my vote for simplification to four— corresponding to
the present subspecies umhellus, togata, sabini, and umbelloides.
However, for the sake of those who find merit in taxonomic detail, I
commend the revision of Aldrich and Friedmann.

Immature. The juvenile plumage is similar to that of the adult but
with less of contrasting blacks and whites, more of a pattern of
somber browns. The ruffs are inconspicuous and the sexes are identi-
cal. The shape of the primaries and secondaries is less attenuate
than in the adult, due to the greater width of the outer vane and
the more blunt tips. The upper throat and chin are whitish.

The downy young are brownish buff on the upper parts mottled
with pale buff, and yellowish buff below. There is a prominent black
line extending behind the eye, occasionally broken.

PTERYLOGRAPHY

The fact that the feathers on a bird do not grow out of the skin
evenly over the body, as does hair on a mammal, comes as a surprise
to the average person. As every bird student soon learns, there are
bare areas on the bird from which no feathers arise, and the areas
from which the feathers do grow are called feather tracts. The ex-
tent and arrangement of these tracts (which vary greatly among
])irds) is called pterylosis and their study is called pterylography—
the geography of feathers.

10 The Ruffed Grouse

Ptilosis— Types of feathers and their characteristics/ Before dis-
cussing the feather tracts and spaces, let us consider the character
and texture of the feathers themselves. This is known as ptilosis, or
plumage:

Neossoptiles make up the natal down, distinguishable from simi-
lar feathers in the adult by the presence of pigment. They are the
tips of the barbs of the succeeding juvenile feathers, sometimes
called mesoptiles. Their length varies from six millimeters in the
capital and dorsal cervical tracts to thirteen millimeters in the sternal
and femoral.

Filoplumes are degenerate feathers, appearing almost like hairs.
They arise from the skin elevations from which come also the teleop-
tiles and semiplumes. From zero to four may arise from these skin
elevations. They range from half to three-quarters the length of the
larger feathers.

Plumulae are adult down feathers, similar to neossoptiles but
without pigment. They differ from semiplumes in the absence of a
rachis. Strictly speaking there are no plumulae in the grouse, but
those feathers resembling plumulae but having slight pigmentation
have been called by this name.

Semiplumes are between plumulae and teleoptiles in character.
They are always covered with other feathers, are downy, but possess
a rachis. Large semiplumes, for all purposes merely downy teleop-
tiles, are found on the sides of the abdomen. Smaller ones occur in
the posterior part of the spinal tract, in the cervical spaces, beside
the primaries and many of the secondaries, and just above and below
the bases of the inner rectrices.

Teleoptiles are the typical feathers of the adult— rectrices, remiges,
coverts, contour feathers, and others. They vary widely according to
modifications for various functions from the highly developed,
almost completely pennaceous (i.e. not downy) rectrices and
remiges to others difficult to distinguish from semiplumes. Some ex-
tremely modified teleoptiles are those of the eyelid, the oil gland,
the ear, and the anus. Some, as those of the eyelid and oil gland, are
probably functional and are small and modified of necessity; the
others are likely only vestigial. In these modified feathers the barbi-
cels are usually absent, the rachis ( oil gland ) is sometimes missing;

^ For good photographs of several types of bird feathers, see Life magazine for
July 17, 1944, pp. 8-11.

The Bird ItseK 11

the barbs and barbules are few and stiff in the auriculars to allow
free passage of sound; and in the anal feathers and those of the oil
gland the aftershaft is practically gone.

The aftershaft is well developed in the ruffed grouse in all teleop-
tiles except the rectrices and functional remiges; in the anal and oil
gland feathers, as noted above, it is often not distinguishable; it is
rather poorly developed on the alula quills, axillaries, and the
auriculars. It is always plumulaceous (i.e., downy or fuzzy) and pig-
mented only at the tips of the longest barbs.

Sexual Variations in Feathers. Variations in the pterylosis between
the sexes are minor. The number of feathers modified to compose the
ruff is greater in the male. There is a tendency for the male to have
more feathers than the female, possibly due to his larger size. The
size of some of the feathers, as the rectrices, is larger in the male.

Snowshoes. These are cuticular outgrowths along the edges of the
toes that grow in the fall and shed in the spring. Their purpose is to
assist the biid in walking on snow by widening the toe area— hence
the name snowshoes. Normal length is about two millimeters but in
the northern subspecies tend to grow a little longer.

Moult. The ruffed grouse has a single moult each year. With the
immature birds this takes place in August and September when the
juvenile plumage is replaced by the first adult feathers. There is
some evidence that the outer primary in this first adult plumage is
shorter than in subsequent years.

The adults begin their annual moult in late July and continue into
September. There is a period when many of the flight feathers have
been dropped and the new ones are not well developed that the
birds have some diflBculty with flight. At this time it is not uncom-
mon to see a "tailless" grouse.

It has been believed by some that a restricted pre-nuptial moult
takes place in the ruffed grouse. This was originally based on two
specimens of B. u. sahini described by Dwight (1900), which were
collected in British Columbia in 1889, one on May 20 and the other
on June 2. These specimens have been re-examined by Trainer who
likewise found a few growing feathers on the head and throat
regions (letter, January, 1942). He says, however, that the evidence
indicates that these two birds are unusual and that the normal thing

12 The Ruffed Grouse

in the ruffed grouse is a complete absence of moulting in the spring
of the year. None of the thirty birds Trainer examined in the Cornell
University collection that might have shown this moult did so.

Pterylosis— Feather tracts and spaces. The area of skin on the ruffed
grouse may be divided into ten feather tracts which are separated
by twelve bare areas, or spaces ( see Fig. 2 ) . These may be described
as follows ( Trainer, 1938 ) : ^

The Capital Tract includes all feathers on the head except those
on the under center portion. Several isolated spaces, noted later, are
found here. The feathers in this tract are all small, all point back-
ward. Longest are the erectile crest feathers. Some of the feathers
around the eyes are among the smallest on the body, being about
two millimeters long. The feathers in this tract are divided into ten
regions, named according to the part of the head: frontal, coronal,
occipital, superciliary, loral, rictal, ocular, malar, auricular, and
post-auricular.

The Spinal Tract covers the upper surface of the body proper,
from the capital tract to the caudal tract. It is divided into three
regions: cervical, interscapular, and posterior.

The Oil Gland Tract is at the posterior end of the spinal tract and
consists of eight little (three and four-tenths millimeters) feathers
arranged in a circle on the tip of the gland.

The Caudal Tract consists of the rectrices (tail feathers) witli
their upper and under coverts and the feathers around the anus.
There are normally eighteen rectrices, although occasionally birds
are found with twenty ( usually males ) or sixteen ( usually females ) .

The Ventral Tract includes the whole under side of the body from
the mental angle to the anal region. The foremedian portion is di-
vided into the interramal region from the chin to below the eyes, the
submalar region and the cervical region; the posterior median por-
tion is called the abdominal region, and the lateral portions are
divided into the sternal and axillar regions on each side.

Most notable feathers in the cervical region are the five rows
which compose the ruff. They are situated near the middle of the
length of the neck. The length of the male ruff feathers increases in
size from sixteen millimeters to about seventv millimeters and then

■^ The descriptions of feather tracts and spaces are derived entirely from Trainer's
work.

i3
a

>

>

o
Q

X

>

a

3
O
1-1

O
i±i

■XI

a.

-d
c

ci
ID

6

13

14 The Ruffed Grouse

decreases to fifty millimeters at the back row; the female ruff feathers
average shorter. All of the ruff feathers have truncated ends made by
the exceptionally long proximal barbs.

The Humeral Tract bears the scapular feathers, which consist of a
band of large feathers extending obliquely across the humerus. Be-
giiming at the lower end of the neck they extend to the back edge
of the upper ann.

The Alar Tract consists of all the wing feathers other than those
of the humeral tract. Many of these wing feathers are identifiable as
individuals and hence are significant taxonomically. The ti-act is
composed of primaries, greater primary coverts, middle primary
coverts, secondaries, gi-eater secondary coverts, middle secondary
coverts, carpal remex, carpal remex coverts, lesser secondary coverts,
marginal coverts, alula, alula coverts, carpo-metacai-pal coverts,
under gi'cater primary coverts, under middle primary coverts, under
lesser primary coverts, under greater secondary coverts, under mid-
dle secondary coverts, under lesser secondaiy coverts, and under
carpal covert.

The Femoral Tract covers most of the sides of the thighs with the
rows radiating posteriorly from an apex at the anterior end, and the
more posterior row curving somewhat dorsally.

The Crural Tract is divided into two regions, internal and exter-
nal, covering the lower leg. The feathers are rather sparse, soft and
hairlike, with a plumulaceous aftershaft extending about half their
length.

The Pedal Tract is a continuation of the crural tract onto the tar-
sus. The feathers are also similar to the crural feathers, and are even
more modified in the same direction. The rachis and calamus are
almost entirely gone, and the plumulaceous aftershaft is short.

The spaces, or bare areas, separating these tracts are described as
follows:

Capital Spaces: The temporal space, in front of and above the ear,
is covered by the feathers of the ocular and upper auricular regions
of the capital tract. The superciliary space above tlie eye is covered
by the feathers above it, while the rictal space posterior to the nos-
tril is covered by feathers from the rictal region above it. Over the
ramus of the lower mandible is the submalar space, covered by the
adjacent feathers of the submalar and interramal regions, while be-
hind the culmen the bare frontal space is located.

The Bird Itself 15

The Lateral Cei-vical Space extends along each side of the neck
from a point in front of the middle of the neck back to a point oppo-
site the hmiieral tract. Occupying about one-sixth of the circum-
ference on each side, this space lies predominantly above the median
line.

The Lateral Trunk Space includes all the several spaces around
the v^ings and legs. At the front it is continuous with the lateral
cervical space above the wing; farther back it separates the femoral
tract and the posterior part of the spinal tract, and then curves
medially in front of the upper tail coverts to meet its other lateral
half at the oil gland. It is covered by the feathers of the back part
of the spinal tract and on the sides by some of the femoral tract. It
is also connected with the inferior space by narrow bands extending
between the stenial-axillar and abdominal regions on the ventral
side.

The Spinal Space is found along the median line of the spinal
tract from the middle of the interscapular region to a point opposite
the middle of the femoral tract. It is covered by the adjacent feath-
ers of the interscapular region.

The Inferior Space lies between the two halves of the ventral tract
along the medio-ventral line, beginning on the neck just in front of
the ruff and extending back to within about fifteen millimeters of
the end of the sternum. It is covered by feathers of the lateral cervi-
cal, sternal and abdominal regions.

The Upper Wing Space is the area distal to the humeral tract,
bounded by the marginal feathers of the shoulder, the secondary
coverts and the feathers on the posterior edge of the upper arm. It
is very irregular in shape and is covered by the marginal feathers of
the shoulder, the proximal lesser secondary coverts, and the distal
scapulars.

The Lower Wing Space is continuous with the lateral space of the
trmik and extends to the underwing surface in-egularly to the base
of the first primary. It is mainly covered vdth the feathers anterior
to it but next to the body the adjacent body feathers cover it.

The Crural Space separates the femoral tract from the lower leg
plumage. It is a very small naked ring on the outer surface of the
upper tibial region, continuous with the lateral trunk space.

These spaces vary from being entirely bare to having a sparse
covering of plumulae. Their function is to facilitate movement or

16 The Ruffed Grouse

position, as flying or walking, or fitting one part of the body into
another, as in roosting, and probably economy of weight.

The Brood Spot: The female bird plucks the feathers from her
abdominal region during incubation in order to make proper body
contact with tlie large clutch of eggs. This bared area is called the
brood spot. Normally all of the feathers on this spot are plucked
except the filoplumes. Most of the abdominal region is plucked, as
well as adjacent portions of the sternal and axillar regions and the
femoral tract.

The Numbers of Feathers on a bird may vary considerably. Some
indication of the number of feathers (teleoptiles and semiplumes)
in the various tracts may be gained from the figures of Trainer ob-
tained by plucking a female weighing five hundred thirty-nine and
seven-tenths grams, collected April 27, 1937. He points out that the
distinction between certain tracts is indefinite owing to the indefi-
niteness of some of the tract boimdaries.

Number of Feathers

Tract

(both sides)

Capital
Spinal
Anterior

776
197

Posterior

360

Oil gland
Caudal

8
164

Ventral

Cervical, submalar.

interramal 241

Sternal, axillar

158

Abdominal

196

Humeral

144

Alar

1104

Femoral

284

Crural and Pedal

710

Total 4342

Range. The ruffed grouse in its six subspecies is found over most of
the wooded areas of the United States and Canada. B. u. umbellus
ranges from the Appalachian Mountains in northern Georgia,
northern Alabama, and the Garolinas, through western Virginia and
Maryland, Tennessee and Kentucky, southern Ohio, Pennsylvania
and northern New Jersey, and northward approximately to a lati-
tudinal line through the center of Minnesota, Wisconsin, Michigan,
New York and northern Massachusetts. Here it blends into B. u.

The Bird Itself 17

togata. Scattered outposts remain in eastern Kansas, Missouri (?),
Iowa, Illinois, and Indiana.

B. u. togata ranges northward from the Minnesota-Massachusetts
line referred to above through Ontario, Quebec, and New Brunswick
provinces, practically to the limit of trees in northern Ontario and
Quebec/

B. u. umbelloides is found from Alberta and west-central Macken-
zie south to northern Utah, northern Colorado, and western South
Dakota, including the whole of British Columbia east of the Coast
and Cascade ranges (A.O.U. Check List, 1931).

B. u. sahini ranges from Vancouver Island and the adjacent main-
land coast of British Columbia south to Humboldt County, Calif,
(op. cit. ).

B.u. thayeri is restricted to Nova Scotia and probably eastern New
Brunswick (op. cit.).

B. u. yukonensis is found in the interior of Yukon Territory and
Alaska (op. cit.).

Eggs. The eggs of the ruffed grouse are oval to short ovate in shape,
broad and blunt at one end and more pointed at the other. The color
is grayish-white to buffy, and only occasionally speckled. After a
period of incubation and exposure they often become quite stained.
The size averages about thirty-eight and five-tenths by thirty milli-
meters. Extremes of size of the eggs examined by Smythe (1925)
were: largest, forty by thirty-two millimeters; smallest, thirty-three
by twenty-five millimeters.

REFERENCES AND CITATION SOURCES FOR DESCRIPTION OF
THE RUFFED GROUSE

Aldrich, J. W., and Friedmann, H. A Revision of the Ruffed Grouse, Condor,

Vol. 45, No. 3, May-June, 1943.
A.O.U. Check List of North American Birds, 4th Ed., 1931.
Bailey, H. H. An Undescribed Race of Eastern Ruffed Grouse, Bull. No. 14,

Bailey Mus. & Lib. of Nat. Hist. Jan. 5, 1941.
Bent, A. C. Life Histories of North American Gallinaceous Birds, U. S. Nat.

Mus. Bull. 162, 1927.
Conover, H. B. A New Race of Ruffed Grouse from Vancouver Island, Condor,

Vol. XXXVII, July, 1935.

■^ Wetmore ( 1941 ) identifies birds from the mountains sonth from West Virginia
as B. u. toaata.

18

The Ruffed Grouse

Coues, E. Key to North American Birds, 5th Ed., Vol. II, 1903.

Jaycox, T. W. [Bonasa Jobsii, n. sp.]-The Cornell Era, Vol. 4, p. 182, Dec. 8,
1871.

Smythe, T. Studies in the Life History of the Ruffed Grouse, unpublished
Ph.D. thesis, Cornell University, 1925.

Todd, W. E. C. Eastern Races of the Ruffed Grouse, Auk, Vol. 57, No. 3,
July, 1940.

Trainer, J. E. The pterylography of the Ruffed Grouse, unpublished M.S. the-
sis, Cornell University, 1938.

Uttal, L. J. Tarsal Feathering of RuflFed Grouse, Auk, Vol. 58, No. I, Jan., 1941.

Wetmore, A. Notes on North Carolina Birds, Proc. U. S. Nat. Mus., Vol. 90,
No. 3117, Wash., 1941.

19th Supplement to the A.O.U. Check List of N. A. Birds, Auk, Vol. 63, No. 3,
July, 1944.

Chronicle of the Ruffed Grouse in Eastern United States

During Indian times the papahcogh was an important item in the
economic life of the Amerindians of the Northeast. How long this
relationship had existed before the advent of the white man is not
recorded, but it probably went back to an era when the aborigines
were far more primitive than when first observed by the Pilgrims in
1620. It may have begun when the people first moved in from the
West to settle what is now the northeastern United States. The
grouse, or at least a bird very closely related to our modem species,
has been recorded in remains from the Pleistocene period (circa
25,000 years ago ) from areas in both the eastern and western parts
of the country.

It is highly probable that the native birds and mammals were
fairly stable elements in the environment of those times. At least we
find no evidence that the Indians either exterminated or seriously
threatened any species. Even though the giouse was a staple item
of food, the proportion of game to the sparse population of humans
was such that hunting was not a serious threat to it. The grouse and
its natural enemies, including man, lived in a rather harmonious
balance. There is evidence, however, that there were years when
game was scarce, and presumably grouse were included. Indian lore
tells of winters of famine, and Longfellow refers to them in his "Hia-
watha." It should be stated again, though, that such temporary
periods of scarcity were not likely due to overhunting.

The White Men Come. When the Pilgrims came to Massachusetts
they found a land abounding in game. Of course there were not
many of them at that time, but still the hunting was wonderful for
the few that there were. Many of the old prints of the Pilgrim days
show the grouse, along with the deer and the turkey being brought

19

20 The Ruffed Grouse

home, as the prize of the hunt, to the log cabin as the fitting larder
for a pioneer family.

It is to be expected that such a creature would quickly find its
way into the writings of the time. Probably the first printed record
is that of Morton in 1632, just twelve years after the initial white
settlement. He was greatly impressed with the abundance of the
bird, having seen as many as forty in a single tree at one time. Baron
de Lahontan, in 1703, wTote of the great abundance and "comical"
stupidity of the "wood hen," as he termed it. This "fool hen" charac-
teristic did not detract from the high regard in which the bird was
held as a table delicacy. Audubon, writing in 1856, considered it
second only to the wild turkey among the upland game birds as a
delectable food.

A Change of Character and Economics. The journals of the early
American ornithologists contain many notes on the distribution,
abundance, and habits of the species. From Wilson ( 1812) in Penn-
sylvania and Nuttall ( 1832 ) in New England to Bartram ( 1751 ) in
Georgia, and Swainson and Richardson (1831) in Saskatchewan,
the story developed until by 1849 Audubon determined that its
range was the whole breadth of the continent. As the settlers ad-
vanced into the wilderness, the grouse continued to play an impor-
tant part in the home diet.

With the forests being cleared to make way for agricultural
lands, grouse range was vastly restricted. In the process, the range
was temporarily improved by the vast development of edges ^
through the extensive woods, but ultimately the clearing reached
such proportions that the species was eliminated from large areas
and confined to a small fraction of the former range in others.
Throughout the eighteenth and nineteenth centuries this process
continued. Concurrently, as farming mtensified and communications
developed, cities sprang up all over the nation and the grouse ceased
to be of great importance as a staple food. An era of commercializa-
tion developed, with professional hunters and trappers replacing
the scattered pioneers. Along with the passenger pigeon and other
game birds, the grouse were mostly shipped to the big city markets.
The birds became scarce in the vicinity of big cities as this traffic

"^ "Edge," as used in wild-life management, refers to the boundaries between
dissimilar types of cover, as for example hay field and woodland. It is so significant
in animal ecology that game is sometimes said to be a product of edges.

Chronicle of the Ruffed Grouse in Eastern United States 21

developed. As early as 1812, Wilson records the retreat to the in-
terior of the species away from the vicinity of Philadelphia.

The abundance apparently fluctuated considerably in different
parts of the Nortlieast and the market price varied accordingly.
From twenty-five cents a brace (Boston 1831, Cincinnati 1820) in
the first third of the 1800's prices ranged to as high as two dollars a
bird in New York later in the century.

Even when civilization pushed the species back from the cities,
hope was held out for its recovery, as witness the item in the U. S.
Gazette of November 18, 1832: "We are glad to perceive that the
citizens of Bucks County [near Philadelphia] are giving heed to the
preservation of game. . . . Few of that fine species of bird [the
grouse] are to be seen. . . . Let us encourage their growth and we
shall be repaid by purchasing them a few seasons hence at fifty cents
per dozen."

During the last two decades before 1900, the importance of the
grouse in commercial channels waned with the diminishing supply,
and at the same time interest in hunting as a sport was growing
rapidly. Demands for the legal elimination of commercial hunting
became more and more vociferous until one after another the states
enacted the needed restrictive legislation early in the present cen-
tury. New York's law prohibiting sale of grouse was passed in 1903,
Pennsylvania's was enacted in 1897.

The emphasis on grouse hunting as a sport and form of recreation
has increased by leaps and bounds since the passing of the era of
exploitation. Today he is called "the king of game birds" by most
hunters familiar with him. In Pennsylvania it is the oflBcial state bird.
That the grouse should attain such importance in the sporting field
is a tribute not only to the wisdom of creating and enforcing ade-
quate conservation laws but even more to the ability of the species
to adjust itself to changing environment.

Reference has already been made to the "fool hen" characteristic
of the bird that so amused de Lahontan. One early author told of
hunting grouse with dogs— the more mongrel and yappy the better-
wherein the trick was to first flush a covey. They would alight in a
near-by tree and watch the dogs as they gathered at the base of the
tree and ran around in circles, all the while making a great noise.
The hunter then picked off the birds with his rifle. So long as he was
careful to pick the lowest bird each time, so that its falling did not

22 The Ruffed Grouse

disturb the others, he would get them all without any flying. While
I have never witnessed a phenomenon like this, I have shot gi-ouse
in the deep Adirondacks with a high-powered rifle by taking their
heads off, and within the past ten years. However, this is a far cry
from the sporty bird of most of our Northeastern coverts today. Even
those in remote mountainous areas are seldom the "fool hen" type.
The abflity of the grouse to make this change is truly remarkable.
Instead of a dumb biddy sitting on a limb awaiting its execution,
today the normal grouse flushes so fast that a hunter hardly has time
to get his gun braced to his shoulder in time to shoot, let alone to
take aim. How amazing is this transformation of character may be
judged by comparing it with the record of its near relative, the
Canada spruce grouse ( Canachites canadensis canace ) . This grouse
was unable to meet the adjustment necessary for survival. It re-
mained a fool hen, and today is exterminated from all of its former
range except where an area of wilderness has remained beyond the
hand of man.

Cycles and Surveys. When the transition from market produce to
game bird extraordinary was completed, the grouse assumed a status
of far greater importance to man. It is natural then that the sporting
fraternity and nature lovers as well would become greatly concerned
when sudden scarcity seemed to threaten the species with extinction.
Then, as each crisis passed and was followed by a resurging abun-
dance, the alternating periods of scarcity and plenitude appeared to
possess an inexorable rhythm. Thus came the concept of the popula-
tion cycle. The baffling nature of the cause of this cyclic phenomenon
led many observers to the conclusion that some inscrutable, all-per-
vading force must be guiding it. This led naturally to the suggestion
that the cause must be ethereal, probably emanating from the sun.
Concurrent investigations indicating cycles in solar conditions abet-
ted this theory. This being the case, these cycles must have been
going on from time immemorial. And so the search for records be-
gan; records that would substantiate the existence of periods of
scarcity prior to those that had been fully documented. They were
not hard to find.

The Indian lore telling of winters of game scarcity are probably
the earliest indication of the then unrecognized cycle. It did not take
the white man long, however, to register the occurrence of grouse

chronicle of the Ruffed Grouse in Eastern United States 23

depletion. Nuttall, in 1832, in New Hampshire, an article in the U. S.
Gazette for November 18, 1832, and an anonymous editor in New
York the same year, called attention to the disappearance of the
birds in the winter of 1830-31 where they had been abundant the
year before. Simple explanations were offered: migrated southward
and hard winter respectively. They had not learned enough of the
problem to make it complicated.

Written records reveal a few more so-called disappearances in the
nineteenth centuiy. The most definite records of scarcity are for a
few years after the Civil War in eastern New York, for 1866 in Ver-
mont (but abundant in Massachusetts that year), 1877 in New
York, 1883 in Ontario, and 1896-97 in Pennsylvania. After the turn
of the 1900's, when the interest in sport had greatly increased, the
story is far more complete.

An exceedingly hard winter in 1903-04 seemed to have resulted in
serious grouse losses in the Northeast. Following this, Forbush un-
dertook the first of several questionnaire studies by sending out an
inquiry to several hundred naturalists and sportsmen in Massachu-
setts. When the very severe scarcity of 1906-07 came it was quite
generally recognized throughout the Northeast. It was feared that
the end of the grouse was at hand. So alarmed was the Forest, Fish
and Game Commission of New York that an investigation was
ordered. The questionnaire survey which resulted ( Woodruff, 1907 ) ,
in addition to the four standard explanations— disease, bad weather,
predators, hunting— added five more to the growing list of possibili-
ties. He concluded that it probably was caused by an unhappy com-
bination of the first three of these four.

While the meeting of the minds brought about by the 1907 survey
resulted in the concept that a combination of factors may have im-
pelled this gi'ouse decline, the survey covering the next great decline
disclosed just how complicated the problem can become when more
attention is given to it. Following the 1914-16 period of decline, the
New York State Conservation Department, this time in cooperation
with the American Game Protective Association, again sponsored a
questionnaire survey (Stoddart, 1918) in an attempt to ferret out
the cause of the decline. No less than forty-five contributing causes
are listed, the most prominent being foxes, bad nesting seasons, and
hawks. Great emphasis is laid on predation; there are eighteen
factors within this classification. Among the also-rans are: driven out

24 The RufiFed Grouse

by pheasants, shooting from autos, pasturing of cattle in woods, and
summer boarders and foreigners without hcenses. It is interesting
to note that disease was given relatively Httle importance in this
inquiry, whereas it had been considered veiy important in the 1907
study. The note of new stock needed is the first mention we find of
the inbreeding factor, destined to be highly ballyhooed a few years
hence.

In 1924 an out-of-step decline seemed of sufficient severity to
institute another investigation. At the American Game Conference
that year Dr. A. A. Allen, who had been investigating the artificial
propagation of grouse since 1919, reported the discovery of the
stomach worm (Disphanjnx spiralis) of such severity in New York
grouse as to assume epidemic proportions. He suggested that it
might be the cause of grouse cycles. At the instigation of Senator
Walcott a national grouse investigation was created, with the late
John B. Burnham as chairman of the committee. The work of this
committee, directed by Dr. A. A. Allen, was concentrated largely on
a study of grouse diseases. Specimens were received from aU parts
of the Northeast. Allen meanwhile continued his studies on methods
of raising grouse in captivity. The next year the New England por-
tion of the work was continued independently by Dr. A. O. Gross
under the New England Ruffed Grouse Investigation sponsored by
the Massachusetts Fish and Game Association, and Allen continued
his work in New York, examining birds from Minnesota, Michigan,
Pennsylvania, and New Jersey, as well as New York.

The 1924 decline was short-lived in most parts of New York and
the birds were again plentiful in 1925. But the gains were not held
for long. Indications of decline were noted in 1926 and by 1927 the
species had declined to the lowest levels known. Hunting seasons
were completely closed in New York and most other near-by states
for a year or more. Both Allen and Gross examined hundreds of
specimens sent in by interested sportsmen and scientists from all
over the Northeast. In 1929 Gross reported: "Work on all phases of
the life history of the ruffed grouse will be continued, but this fall
we will give special emphasis to an intensive study of Dispharynx
which has been found to be the most important parasite affecting
the grouse. . . ." Altogether, seven progress reports of these investi-
gations were made, two by the committee of the American Game
Conference and five by the New England Investigation.

Chronicle of tlie Ruffed Grouse in Eastern United States 25

Chafing under the repression of two successive closed seasons on
grouse in New York, leading sportsmen of the state met to consider
what might be done to remedy the alanning situation. They re-
quested Dr. Allen and Herbert Stoddard to draw up a compre-
hensive plan for a thorough investigation of the grouse in New York
State, the proposed study to be financed by the New York State
Conservation Department. As a result of their recommendations the
Conservation Department in 1930 entered upon a thorough scientific
study of the whole problem.

From 1930 to 1932 the grouse in the Northeast recovered from the
depression of 1927-29. From 1932 to 1944, the population had nu-
merous moderate ups and downs from a general plateau of moderate
to great abundance, none of which can be compared in intensity
with the 1927 crash or probably with some of the high populations
of previous periods of abundance. The declines that have been ob-
served in the East during the 1930's have been very spotty geo-
graphically, and usually quite local. In 1945 the number of grouse
in the Northeast again declined to a low level. Owing to war, no
studies were in process at the time.

Concurrently with the modem New York Investigation, a studv
of the ruffed grouse was being made in Minnesota under the leader-
ship of Dr. Ralph T, King. Begun in 1929, these studies reached a
climax in 1934 when the observers witnessed a very severe grouse
decline. As a result of his studies. Dr. King came to the conclusion
that the cyclic decline was precipitated by an upset in the reproduc-
tive functions of the female followed by an almost complete loss of
young birds of the year. The imderlying force causing this condition
appeared to be beyond the control of man.

In the 1930's there were a number of smaller studies of the grouse,
some confined to limited phases of the broad problem. C. H. D.
Clarke in Ontario (1932-36), L. W. Fisher, F. W. Baumgartner
and others in Michigan ( 1932-39), studied many of the factors gov-
erning populations. Other studies were made in Pennsylvania, Con-
necticut, Maine, Ohio, Virginia, New Hampshire, Wisconsin, and
Massachusetts.

And so, with scientists of all classes sitting on the sidelines waiting
to examine every aspect of the long overdue cyclic decline— theoreti-
cally to have occurred in 1937 in the Northeast— we find that appar-
ently the phenomenon wasn't what it was thought to be after all.

26

The Riiffed Grouse

REFERENCES AND CITATION SOURCES FOR THE CHRONICLE OF
THE RUFFED GROUSE

Audubon, J. J. The Birds of America, 1856.

Audubon, J. J. Ornithological Biography, Vol. 5, 1849.

Bartram, J. Observations in His Travels from Pennsylvania to Onondaga, Os-
wego and Lake Ontario, 1751.

Forbush, E. H. Birds of Massachusetts and Other New England States, 1929.

Gross, A. O. Annual Report of New England Ruffed Grouse Investigation,
1929.

Lahontan, Louis Armond deLorn d'Arce, Baron de. New Voyages to North

America, 1703.

Morton, Thomas. New English Ganaan, containing an Abstract of New Eng-
land,' 1632.

Nuttall, Thomas. A Manual of the Ornithology of the United States and
Canada, 1832.

Pahner, T. S. Glironology and Index of the More Important Events in Ameri-
can Game Protection, 1776-1911, U.S.D.A. Biol. Surv. Bull. #41, 1912.

Stoddart, A. M. Ruffed Grouse in New York State, 1918.

Swainson, Wm., and Richardson, John. Fauna Boreali; Americana, Vol. II,
1831.

Wilson, A. The Natural History of the Birds of the United States, Vol. VI, 1812.

Woodruff, E. S. The Ruffed Groiise, a Study of the Causes of Its Scarcity in
1907, 1908.

3

Biography

April Showers and May Flowers. Each dawn arrived a Uttle sooner
than the day before and each dusk delayed a Uttle longer. The sun,
when it shone, became wanner day by day. The winter snow had
disappeared except for a few residual patches on the cool north
slopes. These changes brought new experiences and sensations to
the grouse that had been bom and raised on the other side of the
woodlot the summer before. They had not witnessed such changes
and sensations before. It was thrilling.

All winter long they had lived along the hemlock ravine. They had
several other companions off and on since autumn but they had not
seen any of them for several weeks. One day the female felt more
and more the need of solitude. The deep shelter of the ravine did not
seem to fit the new feeling in the air. And so she wandered off— over
near the little slash where first she had seen the light of day. There
she encountered another grouse and, still wanting a place all her
own, she decided to press on farther.

Just beyond the slashing was a woodland of young maples and
beech. A logging road traversed this woods from the slashing, and
wound its way down over the hill to the farmstead in the valley be-
low. As the young hen bii^d leisurely picked at fresh spring gi-eens
and jumped for the first insects of the season, the sun came up higher
through the lower tree branches and the old trail with its grassy
edges seemed very homy. She was nervous. Something within tugged
at her emotions. There was an increasing urge to respond to an un-
known need. And it was not long in coming. A strange sound rent
the air. Hardly had the first few beats of its thumping passed before
she was tingling with anxiety to the tip of every nerve. Surely this
was what she had been seeking, all unknowingly.

Slowly she made her way back towards the edge of the slashing—
the direction from which this strange sound had come. Stealthily
she chose her footsteps so as not to expose her presence. In just a

28 The RuflFed Grouse

few minutes the tlu-ihiug beats started again— slowly at first, then at
an increasing tempo, until at the end they blended into an indefinite
whir— as though the speed had so increased as to have passed the
field of audible vibrations. The hen bird's heart was in her throat as
she came closer. Then suddenly, as she peered around the end of an
old windfall, she beheld the object of her search. It was the grandest
grouse she had ever seen. She froze in her tracks.

The Courtship. Apparently the old cock bird had spied her the in-
stant she saw him. Facing her with his head turned slightly sideways
he stood erect on a huge old fallen log. His tail was held high and
spread into a huge fan; his wings drooped deliberately at his side,
while the head was drawn stiffly back into an encircling black ruff,
raised high ( see Plate 2B ) . His sharp eyes held her attention stead-
fastly, as though he were attempting hypnosis. Slowly and carefully
he took a stiff step forward— then another, and another. For several
feet along the old log he thus slowly strutted towards her. Then he
broke his gait and quickly jumped down from the log, and just as
quickly regained the strutting pose again. After a moment he reached
quickly down with his head and pretended to peck at each of a
couple of leaves in front of him. In an almost continuous gesture, he
began shaking his head forward and somewhat downward and side-
ways, first on one side and then on the other, and with each shake
emitting a most peculiar sound. With each headshake he made a
double hiss— something like "shh— ushh," the first half with an exhale
and the second with an inhale. After a few preliminary, evenly
spaced hisses, the headshakes gradually became more rapid and the
hisses speeded up with an increasing crescendo, finally reaching the
point where it became physically impossible to shake the head any
faster. A final, long, letting-oS^-steam hiss following the finish of the
headshaking was accompanied by a quick little forward run toward
the hen bird during which every muscle in his body relaxed in a
convulsive quiver.

By this time the female was so overcome with emotion at this
devastating exhibition that she squatted, limp, on the giound. But
when the male suddenly set upon her and struck her vicious blows
about the head, she quickly regained her senses and beat a hurried
retreat.

This performance Allen ( 1934 ) has called the "intimidation dis-

Biography 29

play" and apparently, insofar as the male is concerned, has little
relation to mating. If the intruding bird had been anotlier male, the
same events would have taken place except that a fight might v^ell
have ensued. In this case the winner would remain in the territory
and the loser would seek another location at some distance.

Both the drumming and intimidation display performances are
acquired by instinct. The drumming takes a little time and practice
to attain proficiency— the wings make only a swisliing noise until the
bird learns to make the drum sound, but it needs no guidance to
know what to do. Allen ( 1934) had a captivity raised male learn to
drum, never having seen or heard another grouse.

The hissing performance is a fascinating display. The whole action
takes almost fifty seconds, the first thirty seconds being devoted to
the slow, evenly timed double-hisses, about a second or slightly less
apart; then about the same number of hisses again, thirty to forty,
in fifteen seconds at a rapidly increasing tempo, and finally about
twenty or more hisses at top speed for about five seconds culminating
in the last hiss and run act. In all, the bird makes about ninety
double-hisses in each display, with the beak held wider and wider
open as it progresses, and the final long hiss is emitted with the beak
nearly closed.

The Mating. Each day thereafter, the cock bird could be heard
drumming on his log near the edge of the little slashing. From before
dawn till after sun-up he was most ambitious, dmmming every few
minutes. Then he would slacken ojff and little would be heard from
him until along toward nightfall. In other directions, but farther
away, the hen bird could hear other grouse drumming too— inviting
any and all grouse to combat, as she learned to her sorrow. She
wanted to seek them out too, but her first experience had taught her
the wisdom of caution. After a few days had passed, she could no
longer control her emotions; so off she went to meet her adversary
once more. Again, as he discovered her presence, he ceased his
drumming, paused for a brief moment, then slowly descended from
his log. This time his tail was held low and compact, all the feathers
were depressed, and the look in his eye was one of appealing gentle-
ness, not of fiery defiance. He walked toward her slowly, in a some-
what circuitous route, shaking his head a little, pecking at a leaf now
and then.

30 The Ruffed Grouse

To the hen this new advance was even more convincing than the
first. His appeal was simply irresistible. He pecked gently at the base
of her bill, then, placing first one foot and then the other upon her
back he mounted for coition. She responded with slightly raised tail,
spread wings, and body flat to the ground.

Probably the most peculiar aspect of the sexual relations in the
grouse ( and possibly in most birds ) is the apparent lack of recog-
nition of sex, as such. Differences between birds seem to be exempli-
fied as relative strength or weakness, audacity or timidity. Not only
will a strong male attempt coition with any subjugated bird, whether
male or female, but timid males may actually accept such advances.
In captivity, and possibly in the wild, this exhibiting of physical
superiority is not confined to sexual activity. Once a bird is decisively
beaten, regardless of sex or by what sex, it attains an inferiority com-
plex, and from then on is easily drubbed by any other bird in the
pen. If not removed, it will be killed but, if placed in isolation for a
time, it often will regain its self-confidence.

Sex Rhythm. Why should the male bird have challenged his visitor
to a duel upon the first meeting, and received her as a mate upon the
second? The explanation lies in the phenomenon called by Allen
(op. cit. ) "sex rhythm." Both sexes have a short, restricted mating
period, which may recur several times during the spring season.
Only when both birds are in the mating phase of their physiological
cycles will mating take place. At other times, without this syn-
chronization, and regardless of whether either bird alone is in oes-
trus, mating will not take place. In these periods fighting— the
working of the law that only the fittest can survive— is the order of
the day.

Displays that appear superficially to be courtship or mating per-
formances may take place at any time of year, by birds of either sex
or any age. These are primarily the manifestation of the continuous
urge to show superiority. Coition is often attempted, by either sex
with either sex or by chicks as young as a single week old, but these
exhibitions of male behavior are in reality an attempted demonstra-
tion of individual physical superiority.

Drumming. The truest exhibit of the male character is in his drum-
ming. At once a challenge to other males or an invitation to receptive

Biography 31

females, it is the one display that sets him definitely apart from his
sisters. Grouse have been known to drum during every month of the
year and during every hour of the day and night. The period of
really intensive drumming, however, is the early spring during late
March and April. The amount of drumming begins to increase in
early March and tapers off in early May. There is also a short period
in midautumn when this activity increases for a few weeks. Apart
from these periods drumming is only occasional and spasmodic.

The act is usually performed on a log, but in the absence of appro-
priate, prostrate logs, mossy mounds or boulders, stone walls, rail
fences, or similar objects are sometimes used for the stage. The log,
or logs, selected are usually of considerable diameter, averaging
around twenty inches. They may vary greatly in length but usually
are not less than ten feet, and are almost invariably long-fallen and
moss-covered. Drumming always takes place at one spot. Since the
same log is often used many years, this spot becomes well worn,
e\'en below the moss covering and the bark. The bird roosts at vari-
ous places on the log most of the nights of the spring mating season
and hence a well-used log will have many droppings piled on it.
Considerable drumming-log roosting is also done through the fall
and winter months, and the identity of a drumming log is most easily
made by the presence of piles of droppings and the telltale worn
spot.

Anyone who lives in grouse country can hardly be considered an
experienced woodland naturalist if he has not witnessed a wild
drumming grouse. It is one of those supreme thrills that makes life
so surelv worth living in a world full of sadness and strife. A careful
stalker may approach a bird guided by the sound of the drumming
if there are enough low-hanging evergreens or other cover to obscure
the approach. However, the surest way is to locate a log in regular
use, then sleep near it overnight. By building a screen of evergreen
boughs in front of the blanket roll, about twenty or thirty feet from
the log, the thrilling sight may be obser^^ed with comparative ease.
Any movement must be made with the gi'eatest of caution, for the
slightest disturbance and the grouse will be gone in a flash.

Before a cold April dawn in the spring of 1931 I doubled my
"frozen" feet under me in the sleeping bag with slow, soundless
movement so as not to disturb the cock grouse sleeping on his drum-

32 The Ruffed Grouse

ming log only about fifteen feet away. I had hoped to photograph
this bird and had rigged a litde hemlock blind in which the camera
was ready for action. This particular log was a very old, well-rotted,
mossy granddad of a log, and had been used for drumming for many
years, judging by the deeply worn drumming spot. The drumming
spot was quite near one end of the log, where it hung suspended a
few inches off the ground. And it so happened that at this particular
moment the log broke off, right at the drumming spot, and the free
end fell to the ground. Of course the cock left the place in a hurry.
I figured that my chance of getting a photograph, or even of seeing
the grouse again at tliat place, was remote, although at the moment
I was not sure just what had happened.

Before completing this episode, I must note that there was a small
mossy mound right next to the foot of my sleeping bag, and I went
back to sleep. The next I knew I awoke suddenly just as dawn was
breaking to hear the bird drumming so close it seemed to be almost
on top of me. And truly, he almost was, for I could feel the wing
beats against my feet at the side of the bag— he had selected the little
mossy momid to drum on after having his regular log broken. To say
that I thrilled to the tips of my hair is putting it mildly.

In dmmming, the bird stands crosswise of the log, braces himself
back on his tail, and brings the wings forward and upward with
quick strokes like the beats of a well-trained crew ( see Plate 2A ) .
Slowly at first, then wdth increasing speed, the beats roll on until
finally ending in a rapid whir. The quality of the drmus may best be
called a dull, hollow thumping. They possess a peculiar ventriloqual
quality, often deceiving one as to the direction from which they
come. They are also deceiving as to distance, drumming a quarter
of a mile away sounding practically as loud as if only a hundred
yards off.

The mechanics of the sound was long in dispute. Some held that
the wings struck together behind the back, others tliat they came
together in front; it was believed by still others that the wings struck
the breast, or the log. It remained for Allen to prove by slow-motion
movies that it actually results from the cupped wings striking the
air. Mathews ( 1904 ) places the pitch of the sound at from A flat to
B flat. Smythe (1925) savs: "... the first tones are staccato and
widely separated but the last are run together in a rapid roll . . ."
(see musical expression on next page).

Biography 33

The drumming characteristic has been used in bygone days by
hunters to lure tlie birds to their destruction. By beating on drums
made of an inflated bladder, market hmiters were able to shoot or
net them. In more recent times numerous observers have noted that
grouse are attracted by the sound of wood chopping.

Accel, et cres.

presto

dim.

— -^ dodo

Boom - Boom Boom Boom Boo Boo Bur -r-r-r-r-r-r-r-r-r-r/ "

Some male grouse apparently do not drum at all, nor do these
birds establish drumming logs. Possibly there are one of two reasons
for this apparently abnormal behavior. Most likely is the condition,
demonstrated in captive birds, of having the need for drumming
short-circuited by the ready presence of one or more female birds.
The second condition might be an abnormal physiological deficiency.
Many males use more than one log, but in these cases there is usually
•one primary log, and one or more secondary logs which are used
more or less irregularly. Some birds use the same log year after year.
On an average, there are about one and one-half active drumming
logs per male grouse.

Location of Drumming Log. As has already been noted, the log se-
lected for drumming must have certain characteristics of size and
condition. The log is selected to a considerable extent on the basis
of its location and smroundings. It is almost invariably inside the
woods, not in the open nor on the very edge. It is also almost always
very near to an edge of some type of opening, usually within two
hundred feet. Very rarely is a log used that is deep in a woodland,
far from a clearing. Preference seems to be for a young, second
growth woodland, predominantly hardwood but with a fair scatter-
ing of conifers, especially young conifers. Mature woodlands, solid
coniferous stands, open slashings, and brushland seem to be avoided.
The immediate location preferred is one well exposed to the sun
on one side, preferably the side with a warm, morning sun. At the
same time, one end on one side at least should have good escape
shelter immediately at hand. This shelter is usually a clump of low
hanging conifers, most often hemlock in the transition zone, spruce
or balsam in the Canadian zone.

34 The Ruffed Grouse

Nesting. When last we left our young hen grouse she had just re-
ceived the male in coition. Following her first mating experience she
made her way back to the sunny woods road in the near-by hard-
woods. As she walked easily along, no doubt daydreaming of the
wonderfid new world mifolding to her, she would pick up dead
leaves and toss tliem over her shoulder in careless abandon. She defi-
nitely was in a homemaking mood. The sunlight shining along the
north side of the logging trail made it a delightful spot in the cool
morning hours. There were several trees and old stumps close to the
trail that offered protection for a home and she determined that she
would build somewhere in this area. After looking over the various
possibilities she chose an old yellow birch tree about twelve inches
through. On the south side, next to the trail, a few sprigs of dead
hemlock brush hung about a foot off the gi'ound, the butt end
braced on the gi-ound near by. She would build her nest up against
the trunk beneath this extra little sprig of camouflage, and so provide
additional protection for her coming family.

Scratching in the leaves, twigs and humus she dug a little hollow
about eight inches in diameter and this she lined with dead leaves.
Simple as it was, it would serve as home for about five or six weeks
to come ( see Plate 3 ) .

After completing her handiwork she went about the ordinary
business of living— eating, sunning, resting, roosting— for several
days. She never strayed far from her new headquarters, except for
the occasion when she returned the few hundred yards to the edge
of the slashing where the old cock grouse lived. As the days wore on
it was apparent that a great change was taking place in her body.
The region back of the legs became enlarged, and after about a
week following nest making, she stayed close by the homesite,
finally laying her first egg. After the egg was deposited she carefully
covered it with dead leaves. Rarely did she stray from the near
vicinity. Successive eggs were laid at intervals of about a day and a
half. On one occasion, feeling in an adventurous mood, she barkened
to the drum of another cock grouse, and set off to visit him about a
quarter mile away near the opposite end of her own woods. This visit
added new zest to her exciting new experiences. In a little over two
weeks, eleven eggs were laid and the urge subsided. No longer did
she have the desire to consort with the cock grouse, no longer did
she want to add to the potential family she had already created. Now

Biography 35

she wanted to take care of her eggs, to mother them on to hatching.
She was broody.

Incubation. Although the duty of setting on her clutch for twenty-
three or twenty-four long days was bound to be somewhat of an
ordeal, her devotion to the job was great enough to overcome any
notion of giving up. As the days wore on, her determination to com-
plete the incubation, regardless of any dangers that might arise,
increased with each slowly passing hour.

She had hardly begmi setting when it was apparent that she could
not get all eleven eggs snugly close to her body for even warmth. So
she plucked the feathers from a large part of her lower abdomen,
using the feathers to make a softer lining for the nest, forming a
large brood spot that enabled her to cover all the eggs evenly.

Early each morning she would leave the nest soon after dawn,
being careful to cover the eggs with dead leaves, to get food and
exercise. Habitually she went around behind the birch tree and off
down the hill. Just as regularly she returned by way of a little draw
on the opposite side of the woods trail. Being anxious not to allow
the eggs to chill she would take only twenty or thirty minutes for
her meal and toilet. And before settling on the eggs once more she
would carefully turn each one over.

Late each afternoon the young hen again took time oflF from her
duties to eat and relax. But apart from these two periods each day,
her life was an miending and almost motionless vigil. Life continued
as usual all aromid her. The ovenbird and the hermit thrush sang to
their mates, who were also incubating nests of eggs among the dead
leaves on the ground near by. In the trees above, the black- throated
green warblers and the red-eyed vireos sang the whole day long.
Squirrels and chipmunks scampered about, scolding at any other
creature that came their way. Mice and shrews ran cautiously
through the ground litter. Life was good. A feeling of freshness and
new vitality was in the air, and it was manifest in the actions of all
the woodland folk. When dusk fell, the activity of the day went too.
But new creatures were abroad, many of whom she knew instinc-
tively were not her friends. Now and then a waddling old skunk
would amble by, seemingly not yet fully awake from his long win-
ter sleep. And though he occupied himself in digging for grubs and
small rodents in tlie humus of the forest floor, she would automati-

36 The Ruffed Grouse

cally draw herself in rigidly if he happened to wander near to her
abode. In the distance a homed owl hooted, and occasionally she
would spy a fox sneaking stealthily along the lumber road. But she
knew she was fairly safe if she remained motionless.

She had not seen another grouse since she last visited a drum-
ming log, and in recent days tlie reassuring drumming of the cock
birds had almost ceased. While feeling quite alone, she became more
and more anxious for the fateful time when she could leave the woods
road with her owti precious little family. As the days became weeks,
the dangers seemed to mount. She would often see sharp-shinned
hawks darting through the trees, and soaring red-tails high in the
sky renting the soft air with their screams. Then, what should come
up over the hill along the road but a team of horses with a wagon.
A man, queer creature she had never seen before, was on the wagon
and a dog ran along beside. As they approached, her nervousness
increased and she tightened her feathers against her body. Closer
and closer they approached; the dog headed straight for the place
where she lay. When it seemed certain that the dog would step on
her the next instant, she rose from the nest with a terrific whir of
wings and flurry of leaves. Luckily— or by design— the rush of wind
behind her drew a flood of leaves which settled over the nest, obscur-
ing the eggs from view. The dog was startled, and after turning to
chase her a moment, wagged his tail in delight at the excitement and
followed his master down the road.

The experience was terrifying to the grouse, but she was im-
mensely relieved upon returning a half hour later to find her nest all
safe. Life became routine again until suddenly, just a bit over three
weeks from the time she started incubating, she felt the first little
pecking within the eggs. This signal that hatching was only two days
off made her all the more anxious, and her mother love for the little
chicks that she could not yet see surged to new heights. But her
troubles were not yet over for, just as the dawTi of her twenty-third
day of setting broke and she was about to go off to forage for a little
food, she heard soft footsteps approaching. When it came into view
it proved to be a fox. She had seen several foxes before and so she
sat tight, as she always had, waiting for it to pass on. But as luck
would have it, his course of travel came straight toward her as he
wandered off the trail to look for mice, or a rabbit— or a grouse. At
the last moment Ijefore the animal would surely grab her she de-

Biography 37

cided to make her escape. But reahzing that other hves than her own
were now at stake she had to be more resourceful than to merely
flush quickly to her own safety. She must also draw the intruder
away from her pipping eggs.

Jumping off the nest and behind the birch, she ran, lamelike and
with wings dragging as though mortally wounded, all the while
squealing as if in terrible pain. The fox lunged at her but was just
too short, and she kept on with her broken-wing performance. Again
and again he sprang upon her and each time she would just barely
elude him until finally, having drawn him well away from the nest,
she flew away to safety. She remained away for a quarter of an hour
—a lifetime it seemed— and then returned home with the greatest of
apprehension. But the eggs were there untouched, and they now be-
gan to show the chipping of the egg teeth of the chicks. She was
lucky— lucky that the particular fox she encountered was not wise
to the ways of hen grouse and the nests full of delicious food they
sometimes leave behind them.

A Family Is Born. The next day dawned bright and warm. This was
fortunate, for many early June days are rainy and rain is apt to be
lethal to tender young grouselets. All night long the nest had been a
iDeehive of activity. The new mother tolerantly and tenderly shifted
her weight, first one way and then another, to make room for the
hatching chicks. One after another, the ring around the large end
of each egg was chipped complete and the dowmy, wet little creature
within pushed the cap back to enable it to shove its way free of its
prenatal tomb. There was a continual, pathetic sort of peeping— in
fact there had been peeping for a couple of days if one were to place
his ear close to the eggs to hear it— as the whole family extricated
themselves from the shells. Some of the shells got pushed clear out
of the nest to make room for the expansion. The mother hovered
them carefully for several hours until about noon they had become
well dried.

She raised herself clear of the nest for the first time since return-
ing from the encounter with the fox, the sun shone in on her precious
brood, and they first saw the light of day. Stepping just a few inches
away, she clucked softly for them to follow. In this, their first lan-
guage lesson, the chicks learned quickly for they were aided by a
nivsterious and imcrring instinct to react to such sounds. With a

38 The Ruffed Grouse

little necessary physical encouragement to a few, all were finally
gathered about two feet out of their one and only home. As this was
theii- first physical effort, they naturally tired easily; so the mother
gathered them into a compact group beneath her sheltering body
and hovered them for their first rest. Thus they left their home only
a few short hours after birth, never to return. For such is the way of
precocial birds.

The First Few Weeks Are the Hardest. Birds cannot count very
well, and it's probably fortunate they can't. This grouse was no ex-
ception to the rule, for ff she had been able to take stock of numbers,
she might well have been concerned over the egg that still remained
whole in the nest. For some reason, possibly because it had been
out at the edge of the clutch for much of the time and was a bit re-
tarded from too much chilling, this chick was about a day behind
the others in its development. Another dawn and it would have
emerged from its shell, possibly as healthy as the rest. But no, it was
left to die; deliberately abandoned by a mother who was following
blindly those inexorable laws of nature that only the fittest sumve;
and the species is what counts, not the individual. Thus, even before
it embarked upon its journey into the world, this family had suffered
its first casualty. This was only the beginning of unending trials and
tribulations destined to reduce the number in the family again and
again.

With a half hour's brooding, the hen again led her young hopefuls
on, away from the homesite that held memories of two narrow es-
capes. They went off toward the edge of the little slashing, for there
the insects would be especially plentiful and escape cover always
close by. Progress was slow and the mother spent anxious and busy
minutes keeping her charges from straying too far off, teaching them
to obey instructions, and showing them how to catch insects. Every
little while she would gather them together for brooding, but each
time the space between broodings increased. By dusk they had
reached the slashing, having covered the three hundred yards more
or less in seven hours. Here they hovered together under their
mother's protecting coat of warm feathers for their first night away
from home, this time beside a tuft of grasses beneath a low hanging
hemlock branch.

They were up and away wnth the dawn, searching industriously

Biography 39

for insects— almost any kind of crawling or flying insects tliat they
could catch; scratching a little in the leaf litter on the ground, pick-
ing them off the lowgrowing vegetation, or jumping for them in the
air. Eating insects was mainly a question of taking those kinds that
were most available and easiest to catch— flies, ants, beetles and
many other insects and related forms. Only a few species, such as
some lightning bugs ( Lampijridae ) and lady bird beetles ( Coccinel-
lidae), are definitely avoided by some birds, apparently due to their
offensive taste. While insects and other arthropods constitute almost
the entire food of the chicks for a period after hatching, they soon
learn to add a little fruit to their diet. A few strawberries and similar
early season, soft-bodied fruits are eaten almost from the start.

After the morning meal was over the brood hovered for a short
time before setting forth again. Straying chicks would call a plaintive
"tsee-tsee-tsee-eee" in a rising pitch (Sawyer, 1923). More practice
in discipline was then in order, particularly in responding to warning
calls of danger, for all giouse soon have to learn ever to be on the
alert for the many forms of sudden death that may strike at any time.
They quickly learn to scatter and "freeze" among the dead leaves
at the first warning signal.

It was not long before they had their first opportunity to test their
skill in dead earnest. The alert mother spied a shadow darting
through the trees in their direction. A quiet, stern "freeze" signal, a
sort of low "pe-e-e-e-e-u-u-r-r-r" (Sawyer, 1923) was quickly given
and the whole family squatted motionless instantly. It was too late.
The female sharp-shin had spotted some movement and she plied
her speed and dove earthward in what seemed to be a suicide plunge.
As soon as the grouse mother realized that the attack was on she
gave the violent scatter call, and herself flushed in a broken-wing
act in an attempt to draw the accipiter from her young ones. But the
hawk had concentrated its whole attention on the one chick whose
movement it had first seen and the kill was made with little trouble.
During the excitement the others had made good their escape and
were perfectly concealed in the ground litter in a space about thirty
feet across while the mother hung near by continuing her keep
quiet instructions wdth mewing squeals and sharp "quit-quit" clucks.
By this time the hawk had carried its prey to its habitual butchering
log where it picked the little grouse before taking it to its young in
their nest high in a near-by hemlock.

40 The Ruffed Grouse

When the danger appeared to be past the grouse hen returned to
their midst and gave the all-clear signal, a low "puk-puk-puk." In-
stantly, as if by magic, nine little fluffy brown chicks arose from their
shelter and, peeping joyously, quickly gathered beside their mother.
Off they went at a quickened pace, with no thought on the part of
the parent to check to see ff all were present. She had her family
safe through the first skirmish and she was both thankful and anxious
to put distance between them and the source of their trouble.

That day and the next they traveled across their home woodland
to the edge along the fields that sloped down to the farm home in
the valley below. Lots of shrubs and briars grew along this margin,
and it provided both plenty of food, many tangles and small hemlock
clumps for escape shelter. On the way over, during one of their for-
aging periods when the chicks had spread quite widely, one little
fellow strayed beyond the mother s call. When it realized its predica-
ment it cheeped wildly and pathetically, running first this way and
then that, in an effort to locate the brood. That night it had no warm
mother's breast to keep it warm, and nowhere near enough feathers
of its own yet to do the job. So passed the third member of the origi-
nal eleven. In but a few hours its tiny carcass disappeared into the
ground as the scavenger beetles dug beneath to lower it to their ad-
vantage.

A week passed. The weather had been very kind, there having
been only one light shower one afternoon. The chicks easily kept dry
during that time beneath their mother's wings and breast. Several
times predators had been sighted but attack had been avoided by
freezing. The wing-flight feathers were developing rapidly and the
chicks were enjoying a new half -running, half-flying type of exercise.
By the middle of June, at twelve days of age, they could actually fly
twenty or thirty feet if an occasion warranted real exertion.

Life Without Father. The wind began to whine through the tops
of the near-by hemlocks. It was still several hours before dawn, and
the brood were hovered under the edge of a briar and grape thicket
along the woodland border. In the distance flashes of lightning were
visible and the far-off rumble of thunder gave warning of the ap-
proach of a storm. It was not long in arriving. The wind whipped the
trees savagely as the rain came down in sheets. The temperature had
dropped to the vicinity of the lower fifties Fahrenheit. Such a turn

Biography 41

of the weather is very dangerous to young grouse during the early
weeks of their Hves. A single wet chilling, if not followed soon by a
warm sunning, is apt to be fatal. In violent storms like this some
chicks may be seriously soaked and chilled in spite of the hen's best
efforts. Any abnormal exposure would lead to almost certain trouble.
It was calamitous that upon such an occasion a water-soaked old
skunk, caught away from his den by the storm, blundered through
this particular thicket on his way home and scattered the grouse
family. As quickly as possible the mother recalled her chicks and
hovered them again, but the damage was done. Although the day
came along bright and cheerful, three of the family failed to respond
to the breakfast call. If they had been picked up by a man and taken
to a game pathologist for autopsy, his report would probably read
"cause of death unknown."

It was soon after the encounter with the storm that proved so dis-
astrous that the family met another grouse, an old cock bird. This
was the first of their kind other than their mother that the chicks had
ever seen. The hen had not encountered another grouse since her
last visit to a drumming log before starting incubation. The chicks
stayed close to the mother's side, for they were not quite certain
whether the newcomer was friend or foe. The cock bird, for his part,
was completely nonchalant, paying scant attention. Soon the family
were on their way, the male bird following a short distance behind
for a few minutes, and then disappearing. He might have been their
father ( or one of their fathers ) but if so he showed no paternal in-
terest or sense of duty. While famed for his magnificent courtship,
the male grouse is not a good parent. Among hundreds of grouse
broods observed, only about one in fifteen had the benefit of a
father's presence near by, and it is doubtful if many of these asso-
ciations were anything but accidental. Further, it is probable that
most of the relationships of male to the brood did not involve the
actual father anyway. There is little inclination on the part of the
male bird to associate with the brood at any time, but what solici-
tude does exist is stronger during the early weeks than during the
last half of the summer. About one in ten broods up to six weeks of
age are likely to be found vdth a male adult near by (within two
hundred feet), whereas with older broods the ratio is only about five
per cent.

Each day that the ground was dry considerable time was spent in

42 The Ruffed Grouse

taking dust baths. Following their mother's example, each chick
would wiggle and scratch a little hollow in the dry dust of an open
spot or rotted wood of an old log, whichever was most convenient
at the time ( see Plate 2C ) . They would lie in their bath, first on the
belly, then one side, then the other, all the while scratching and rub-
bing dust with the feathers. Among them all, they kicked up quite
a smoke screen.

Dusting was a regular and necessary event. When handy they
would use the same baths over again. It serves primarily to stimu-
late proper growth of the feathers, although it may help to get rid of
external parasites too.

The little grouse grew rapidly. At three weeks they had become
proficient flyers; at four weeks the feather coat was well enough de-
veloped to make hovering by the mother no longer necessary under
average conditions. By six weeks the full juvenile plumage was at-
tained, and the young grouse looked very much like miniature repli-
cas of the adult. By this time the family was spending its nights roost-
ing in trees, usually hemlocks or other evergreens. It had experienced
many critical moments when natural enemies or the elements had
threatened, but no additional casualties had resulted. Every one of
the five remaining youngsters was now very proficient in making its
escape from running or flying predators, and had grown hardy to
the vicissitudes of the weather. But they still depended very much
on the mother for leadership and warnings of danger.

Revolt in the Ranks. Almost as soon as the youngsters had left the
nest, family squabbles began. The assertion of physical superiority,
or at least equality, is a prime essential for successful survival in a
grouse family or group. Often the affirmation of strength led to
fights, but more often a chick would satisfy his ego by strutting to
amuse or antagonize his brethren. Always the mother bird main-
tained control over these trivialities, though this control became
more difficult as the chicks grew older. As the summer progressed
to the dog days of late July and August, a surging of insubordination
was noticeable on the part of the chicks. They were now half grown
and were beginning to feel a need for freedom, and resented parental
control. They would stray quite far apart in foraging for raspber-
ries and the other summer fruits which now furnished a good pro-
portion of their diet. They would fly off, chasing each other as their

Biography 43

rivalries became increasingly belligerent. In spite of the sometimes
frantic orders from the mother, they would not heed her warning of
impending dangers. So it was natural that the probability of addi-
tional mortality in the family would rise at this time.

One day a Cooper's hawk sat waiting on its perch in a dead chest-
nut tree as two chicks raced recklessly up the near-by draw. In a
flash the hawk plummeted earthward as they approached and with
a quick sideward surge grasped the second of the chicks. Thus the
family was reduced to four youngsters— only thirty-six per cent of
the number in the little nest at the base of the birch tree two months
before.

The Family Grows Up with a Change of Clothes. The youngsters
soon learned the dangers of independence and became more alert
to observe dangers for themselves. Mother was more cautious than
ever now because she was losing feathers and flight was not as pow-
erful or accurate as usual. In the course of a few weeks she com-
pleted her moult and had a full new set of feathers. No longer were
her tail and wing feathers ragged at the tips; every one was as fresh
as a new leaf in the spring. Before she was through with her moult,
the chicks began losing their juvenile plumage. By mid-September
their first adult plumage was almost complete and they could hardly
be distinguished from the mother. Two of them had very rufous tails,
one was mottled rufous and gray like the mother, the fourth proved
to be a gray-phase bird. The family allegiance became very loose,
and the mother no longer concerned herself with her children's
safety. It was every one for himself now, and they were found by
themselves as often as they were together.

Crazy Flight and the Fall Shuffle. As the calendar turned to Octo-
ber, there was a new tang in the air. Many new fruits were ripen-
ing so that the grouse were to be found along woodland edges, in
brush lots and even out along hedge-rows where the hawthorns,
dogwoods, viburnums and other preferred fall foods were to be
found. The youngsters attained their sexual maturity with an ac-
companying resurgence of quarreling, more vicious than ever be-
fore. Now when one of the brood was chased by another the victim
was not likely to return. There was a definite antagonism on the
part of most of the birds for all other giouse. It was not exactly an
exhibition of territorial behavior as in the spring, but operated simi-

44 The Ruffed Grouse

larly and arose from similar physiological causes. This dispersal of
the family groups is called the fall shuffle.

When a bird is driven from its family group it seeks its lebensraum
(living room) elsewhere. However, it may soon encounter another
grouse or brood of grouse, and is then more than ever likely to be
driven out again. With each failure to find a fall territory it can de-
fend, the bird becomes more nervous, and more desperate as a re-
sult of its growing inferiority complex. It is alone in a strange and
unfriendly world for the first time. Although it was brought up in
an area of only forty or fifty acres and had never traveled over a half
mile before, it may now find itself many miles from its summer home.
Under such circumstances, a grouse often will fly far out of its normal
coverts and many times come to a sudden end by flying into obstacles
such as buildings.^ This phenomenon has been termed crazy flight.
Various explanations have been given for these actions, such as a
nervousness brought on by the leaves falling, or irritation caused by
internal parasites. While the falling of the leaves may well add to
the nervousness of the birds, the crazy flight is merely an aberration
of the normal fall shuffle— a social phenomenon that likely occurs
with most sedentary species of birds and is well known in the bob-
white quail.

A-Hunting We Must Go. If we may assume that the young grouse
in our biography did not reside on a survey area of some ruffed
grouse study, it is quite probable that they have never encountered
that most formidable of adversaries, man, up to this time— mid-
October. Then comes a day, according to the calendar and the laws
of the state, when the status of the grouse changes from a protected
bird to fair game: the hunters, who have been chafing in their anxiety
for weeks, swarm over fields and woods as the hand of the clock
passes the fateful point.

Since our grouse famfly has now dispersed into four different
coverts, with only two of them remaining together, we find it con-
venient to continue the life story through the experiences of only

^ These escapades often result in human interest stories; for example, the March
27, 1942, issue of The Boston Herald headlined: "Shipyard Worker Keeps Bird Fly-
ing," then recounted: "The supper which William Catterall, Fall River shipyard
worker, was enjoying . . . last night, was interrupted when a grouse entered the
kitchen amid a shower of broken glass. Catterall, unperturbed, patched the bird's
cuts . . . and released it. . . ."

Biography 45

a part of them. Let us take the two that have remained together
—the mother and one young one.

Not far down the hill from their home woodland toward the farm
in the \ alley below, was an old apple orchard. Between it and the
woods ran an overgrown fence-row, with gray dogwood, wild apple,
thomapple, black cherry and wild grape growing in tangled pro-
fusion. This was just the place for the grouse to be feeding this crisp,
sunny October morning— either along the woods edge, in the or-
chard, or in the hedge-row between. An experienced grouse hunter
would recognize this almost as well as a grouse. So it was not sur-
prising when two hunters, well bedecked in their red plaid hunting
garments, came up the hill from the farm, straight toward the or-
chard. Three beats across the orchard convinced them that no grouse
were there. One of the hunters was particularly slow and deliberate,
examining carefully each likely bit of cover for fresh sign. They
turned up the hill again, one on each side of the hedge-row, working
carefully towards the woods.

Just about fifty feet along the fence-row was a clump of thorn-
apple loaded with fruit, and it was here that the two grouse had been
feeding. They saw the two tall creatures coming and, true to their in-
stincts and experience, crouched motionless on the ground until
they should go away. But these were not ordinary adversaries. Just
as they were about to pass the birds, one of the men gave a kick at
the low-hanging thorn branches. With a terrific whir of wings, the
startled birds burst into the air. Swinging out of the obstructing
hedge-row on the side opposite the hunter who had flushed them,
they made a splendid shot for his companion. The gun blazed twice,
and one of the birds crumpled to the ground. The other, having ex-
perienced one hunting season before, had cut sharply back to the
top of the hedge and on into the woods just at the critical moment,
causing the shot to go wild. She flew far out of sight into the woods,
coming to rest high in a clump of protecting hemlocks. From past
experience, this was the safest place to avoid a repetition of the
death-dealing flight of lead.

With each escape from the hunters' bullets the grouse becomes
more wary and more able to make a successful getaway. This ability
to dodge shot and baffle hunters has been developed in a relatively
short time. Wilderness grouse even today sometimes exhibit the
fool-hen characteristics that most grouse possessed when the white

46 The Ruffed Grouse

men first introduced them to guns. But in coverts frequented by
man, the grouse is well able to hold its own, given reasonable laws,
good cover, and not too great a hunting pressure. Its ability to
change direction in flight, to put trees between itself and the hunter,
to outwit the hunter by flushing behind him are some of the char-
acteristics that endear the grouse to millions of sportsmen and earn
it the title "king of the game birds."

The four remaining grouse of our original brood, each in its own
covert, met and evaded many hunters during the remainder of the
open season. As the hunters' guns became a memory the season rap-
idly changed, and with it the coverts changed too. The leaves fell
from all but the evergreens, making the hardwood areas quite bare
of cover. The berries on most of the fruit-bearing trees and bushes
were dropping rapidly, and although much fruit remained avaflable,
the birds' diet included more and more of tender buds; insects were
practically gone now.

Winter Comes. As the wind swung around to the northwest and the
skies assumed a leaden color, the temperature dropped below freez-
ing and snow began to fill the air. There had been a few snow flur-
ries earlier in tlie fall but this tiine it looked Iflce more serious busi-
ness. Winter had come. To many grouse this meant a change in their
habitual coverts, a move to the shelter of the coniferous woodland.
The hen bird returned to the hemlock ravine where she had spent
the previous winter. Several other grouse had settled there too, and
they banded together much of the time to form a covey. Gone was
the antagonism of the early fall; no longer did each bird look upon
the others as competitors. They settled down for the winter in an
area particularly adapted for winter use; and since such areas were
not over plentiful or too large, it was somewhat of a necessity for
them to get along together if many were to survive.

Snow-Roosting. The birds roosted mainly in the thick hemlocks
but sometimes, when the nights were bitter cold and the wind
high, they would roost in the deep snow. If it were snowing at the
time, a bird might simply squat deeply into the snow and then al-
low the weather to cover it up. On other occasions they would dive
into a deep snowbank from the air. If the weather stayed bad, they
might remain in a snow roost for several days. Occasionally it is
reported that grouse are trapped in snow roosts by a quick freezing

Biography 47

crust but this I have never observed. Knowing the abihty of the
bird to go without food for days on end, it is doubtful if this is ever
a very serious matter, although it is possible that it may actually
happen.

Snow roosts do make the birds quite vulnerable to attack, espe-
cially by foxes. There is always a little breather hole from the birds'
heads to the outside of the snow that is often easy to spot. Grouse
tracks that end in a slight mound of snow are a sure indication of
the birds' exact position. Even a man, if he is skillful, can catch the
birds by hand under these conditions. There is good evidence that
some foxes learn to catch on to these signs too.

The proportion of winter roosting beneath the snow varies in dif-
ferent years and also with the period of the winter. Winters with
little snow will prevent the use of snow roosts to a considerable ex-
tent, but other than this limitation of opportunity the degree of
snow-roosting depends upon the prevalence of abnormally severe
weather, low temperatures and high winds. The first severe storms
of the season always induce more snow-roosting than later ones.
This appears to be due to the relatively greater change from late
fall weather, whereas in the later storms the change in conditions
is not as marked and the birds have then become accustomed to
the hibernal extremes.

Gregariousness. The ruffed grouse is not a gregarious bird in the
full sense of the word: living in flocks. However, it does exhibit vary-
ing degrees of cohabitation with those of its kind. It has already
been noted that in the early spring there is a shuffle from the winter
groupings to the male and female breeding territories. Thus, in the
spring there is an almost complete loss of the gregarious tendency.
We have noted two females nesting concurrently within fifty feet
of each other on occasion. Two females occasionally lay in the same
nest although, so far as we have been able to ascertain, only one
bird does the incubating. Aside from these definitely abnormal as-
sociations, we may say that the species is solitary throughout the
spring period.

During the summer two males are quite often found together, or
a male and female that has no brood. It has already been noted
that a male is occasionally found associating with a brood, although
this is hardly an example of flocking. Several cases were observed of

48 The Ruffed Grouse

two grouse broods consociating but most of these were not regular
associations. In one case two broods stayed togetlier throughout
the summer. This seems to have been a true example of brood
communism.

It is interesting to recall the clear distinctness of the brood identi-
ties in these associations. Upon one occasion of making a brood
count, the observers were confused both by a poor count and an ap-
parent extra female. One of the observers climbed a tree to await
the regathering of the family (the chicks could not fly yet) while the
other observers proceeded on out of sight. Soon one female reap-
peared and, finding the danger past, called her brood from the sur-
rounding vegetation. Out they trooped in perfect order, assembled
their skirmish line, and off they went. A few moments later the
second hen returned and she too called out her brood. From the same
terrain as the first appeared another whole family with just as un-
erring discipline, and they trooped off in another direction. It was
clear that, even though only a week old, these chicks knew the call
of their own mother as compared to other grouse and responded only
to her. On some occasions, usually with birds just after hatching,
there does occur some confusion in brood reassembling.

The groups of grouse that are occasionally found together during
the fall are usually the remnants of a family and hence are not gre-
garious. But the regrouping of birds in the late fall and winter is the
one strong period of communal activity they have. The usual groups
run from three to six birds, the commonest numbers being in order
from the lowest. Groups of from seven up to twelve or fourteen or
even more are occasionally encountered, but not commonly. The
period of greatest group activity is late fall to February. In late Feb-
ruary and March the groups begin to break up to seek out their
spring territories.

The Annual Wheel of Fortune Is Completed. When last we left the
grouse family, one of the young ones had fallen before a hunter's
gun. There were left only four, the mother and three offspring, each
in a different area of coverts. If we may add an adult male to make
the full complement of our original reproductive unit, we have now
carried five of the original thirteen into the winter. We have shovm
some of the problems faced by them as a result of the change of
seasons. Now we may see what happens to these five birds as the

Biography 49

winter progresses and the story returns to the time of beginning,
early spring.

With each change of season the cover types used have shifted,
and likewise the food habits are altered. As fall gave way to winter,
the diet gradually changed from a predominance of fruits to a pre-
dominance of buds. Throughout the winter some fruit is eaten, an
occasional insect and a little green leaf material. But the fare is
largely buds, mostly those of trees found in association with the
conifers or near by— birches, hophornbeam, soft maple, popple,
cherry, and others. Traveling through the woods in winter one can
readily understand why the grouse resort to budding, for food in
other forms is mighty scarce indeed. Yet they are very skillful in
locating available fruit. Even with a deep blanket of snow on the
ground, grouse will find a few cherries, dogwood fruits, and similar
fare when ordinary man is entirely unable to find them.

Food is not usually an important factor in limiting or reducing
winter grouse numbers in coverts that are otherwise satisfactory.
Shelter is very often a delimiting factor, according to the extent,
type, and distribution of coniferous types and dense slashings. The
winter season does bring on problems of survival in relation to the
elements and to predators. While it is rare for ruffed grouse to be
killed directly by freezing or exposure, the rigors of the season do
increase their vulnerability to predation. Snow-roosting, the lack of
protecting leaves on deciduous plants, telltale tracks in the snow,
the bare snow ground cover, all contribute to this problem. Toward
the end of the winter the arrival of the courtship activities greatly
increases the danger of predation.

So it was that in February one of our five grouse was picked out of
a snow roost by a fox and early in March another was taken from
its tree perch in the dead of night by a great horned owl. The three
remaining birds, one the old male, the other two a young male and
a young female, left the shelter of their hemlock winter homes to
locate new homes for the spring. The old male reestablished his ter-
ritory of the previous year and once more boomed his defiance of
the rest of the grouse world from his favorite drumming log. The
young female set forth to find a suitable nesting territory from which
she could seek out desirable males for mating. Possibly she would
return to the part of the woodland where she was bom, now left
vacant by her mother's death.

50 The Ruffed Grouse

Her brother ( or half brother ) also looked for an opportune drum-
ming territory. Several times he was challenged by other males
whose established territory he invaded and each time moved on
until he finally located a desirable log that no other grouse immedi-
ately claimed. This he successfully defended against later comers
and displayed his superiority with all the vigor and arrogance of
his father before him. One dawai while he drummed his challenge
to the world and paraded his wares on his mossy stage, a little thin,
mottled brown and white, hunchbacked animal crept slowly toward
him through the near-by hemlocks. The weasel was unappreciative
of the fine points of the grouse's art; to it this creature was just an-
other opportunity for a full belly. With a final spring from behind
the log the weasel sank its teeth into the grouse's neck. A hectic
stmggle among flying feathers lasted but a few minutes.

As we complete the wheel of fortune of our grouse for the year,
we note that of thirteen grouse early in June— a father, mother, and
eleven youngsters— only two remain, a male and a female. Thus, al-
though eighty-five per cent of the birds have succumbed, we have
as many as we started with. The species carries on as plentiful as
before. What more can we ask?

In recounting the biography of a grouse family around the seasons,
I have attempted to cover the life history and habits of the species.
Inevitably there would be some items that would not fit easily into
this story. These are given below in several subheadings.

Territory. The term territory has been used several times, and in
two different senses. In connection with the breeding season the
male bird establishes an area centering around his favorite drum-
ming log, from which he will drive out all other grouse. Thus he
considers this his own backyard and, except for the reception of
females when he is in the mating cycle, will permit no competition
therein from his ov^ai species. Nice (1941) calls this the "mating
station only" type of territory, one of six types recognized. The size
of this territory is difficult to ascertain and undoubtedly varies
greatly, depending upon the population density and the character-
istics of the habitat. The maximum average size may be derived
by dividing the available covert area by the number of male terri-
tories. This is no doubt too large for a true average, since much of
the area, and notably the nesting localities, are often not frequented

Biography 51

by the male birds. Observations show these maximum averages in
various coverts to run from about ten to over a hundred acres. Cer-
tainly no male bird ever tried to defend a hundred-acre territory,
but what the limits of a bird's ambition would be when given full
freedom to take as much territory as he wished is a matter of con-
jecture. Also, some males have gotten along with considerably less
than ten acres. It is my judgment that when the spacing of male
territories is such that less than twenty acres are available to each
bird, the pressure between them becomes continuously active.

The second concept of territory is that area that an individual,
or a brood, or a group, requires and uses in its year-round or seasonal
activity. There is less connotation of conflict or of exclusive rights
in this type of territory, although at times conflict does occur. The
fall shuffle is the main occasion for this type of territorial activity.
Evidence is that an average of four acres per bird is the minimum
requirement. It will be noted that this figure corresponds to maxi-
mum cariying capacity ( saturation point ) and roughly corresponds
to the minimum average figure for male spring territories ( ten acres
—or five per giouse if sexes were equally divided). This type of
territory establishment does not mean a separation of area for each
bird or group; rather it is the area utilized by them as a unit and
only during the fall shuffle may it result in the exclusion of other
birds from overlapping. An average of eight acres per bird for a
group of six would indicate a forty-eight-acre territory for that
season.

Territory on the part of the brood is simply the area they cover
during the summer. No interbrood conflict is involved and terri-
tories often overlap— even occasionally are practically concurrent.
The area required to raise a brood also varies— generally twenty to
forty acres is utilized. Cases have been observed of broods living the
entire summer in an area of about ten acres while other cases have
involved in excess of a hundred acres. The latter cases are usually
the result of abnormal movements resulting from disturbance.

Daily and Seasonal Range. The spring range of the male corre-
sponds to his territory, while the sum of the seasonal ranges of all
grouse add up to their yearly territory. However, seasonal ranges
vary considerably. The spring range of the female, during egg lay-
ing and incubation is very restricted, often as little as three or four

52 The RufiFed Grouse

acres, with travels of only one or two hundred yards, except for
occasional longer trips to a drumming log. During the summer the
range of the female covers the same territory as the brood, often
extending half a mile or more, while males and broodless females are
more restricted, often remaining within ten acres or less and travel-
ing only two or three hundred yards in distance.

During the fall the greatest distances are traveled. Extremes are
almost always birds of the year seeking new territories. Distances ex-
ceeding two miles are not uncommon, and fairly authentic records
have been made of birds traveling up to twenty-two miles. Terri-
tories are large and daily travels are greater than at any other time.
When winter arrives the territory is more restricted. Except for
occasional warm, sunny days when they may wander afield in search
of berries, they remain close to the winter shelter areas. Two or three
hundred yards would be a normal range on an active day and it
would not greatly exceed this for the season.

Daily range also varies widely, with the season, the weather, and
the type of activity. Normally, the day's travels consume only one
to three hundred yards, but if flushed and reflushed the bird may
extend to a half mile or more. Occasionally a single flight wfll carry
them a quarter of a mile. Females nesting far from a drumming male
travel up to half a mile on their visits for mating.

Year around range normally adds up to about three-quarters of a
mile, outside measurements.

Characteristics of Flight. Having short, rounded wings and power-
ful muscles for rapid wing movement, the grouse is a powerful and
accurate flyer. Its ability to dodge obstacles and change its course
at full speed are famed. However, its accuracy in flight is sometimes
exceeded by its recklessness. A grouse flushed at night will often fly
off as quickly as if it were daytime, unmindful of the branches hit
along the way. Occasionally one hits a tree, fence or other obstacle
and closes its career with a broken neck.

Probably the most marked characteristic of the grouse's flight in
the mind of the average observer is the whirring, sometimes terrify-
ing noise that accompanies it. Springing from the ground near by,
it puts all its power into a quick getaway with the resulting loud
whir. Even the most experienced woodsman wfll on occasion be so
taken unawares by a grouse as to be scared nearly out of his wits.

Biography 53

Few people have the opportunity of observing an unflushed grouse
taking flight at its leisure. When it takes the time to do so it can fly
as silently as an owl.

Duration of flight varies but never very greatly. Assuming that
it may make its own choice of satisfactory cover to alight in, the av-
erage flight from flushing is between three hundred and six hun-
dred feet. Flights made for its own purposes usually are shorter.
If the same bird is flushed about three or four times in quick succes-
sion it can be picked up by hand, exhausted. About half a mile is the
limit of power flight (not counting long coasting) without recupera-
tion of the muscles.

Speed of flight varies with the cover. In woodland with an average
number of trees to dodge, it will be from thirty to forty-five miles
per hour according to the strength of the bird, its anxiety to escape,
and the amount of dodging required. In the open it can fly con-
siderably faster. I timed one bird across an open field, using a stop
watch, at fifty-one miles per hour, measured from the time it left
the ground on one side of the field to the time it entered the woods

o

on the other.

Relation of Nest Location to Drumming Logs. The positional re-
lation of a female nesting site to the nearest active drumming log
is normally negative. If the female has a definite thought in select-
ing her nest site with regard to the location of the male it is to get
the nest as far away from him as practical. I have observed one
drumming log fifteen feet from a nest but I was unable to check
whether or not the log was used after the nest was made. Allen
( 1934 ) says that the distance between log and nest is sometimes as
low as fifty feet and sometimes as far as half to three-quarters of a
mile. The closest I have observed both nest and log in use at once
was seventy-five feet. In this case these two birds habitually roosted
in a hemlock tree over the dmmming log prior to the incubation
period. This is not usual, however.

In numerous cases nests have been observed half a mile from
the nearest known drumming log in areas where we believed we
knew the location of all active logs. The average distance is approxi-
mately one-quarter mile.

Nest Desertion and Renesting. Desertion of the nest is rare in the
ruffed grouse. The few cases observed that appeared to be deser-

54 The Ruffed Grouse

tion (as contrasted to the cases where the hen is killed away from the
nest) have all been in the early stages of incubation or before egg-
laying was completed. It is rarely possible conclusively to prove
willful desertion since it is usually conceivable that the hen may
have been killed and her remains not located. It is true, however,
that the will to stay with a nest is weakest at the beginning and
strengthens rapidly. It never is absolutely complete. If the setting
bird is disturbed enough she may desert at any time. In oiu experi-
ence in trapping and marking setting females about a week before
hatching time about one bud in twenty would not return to the nest
after release.

Renesting is the building of a new nest and laying of a second
clutch of eggs after the first nest has been destroyed. If the nest is
broken up before two weeks of incubation the probability of renest-
ing is fairly good. If the nest is broken up after two weeks of incu-
bation, and most destroyed nests are, there is practically no possi-
bility of a second nest. This is due to the loss of the urge to breed
so late, by both sexes. Explamed in terms of the sex rhythm, the
probability of both hen and cock reaching the mating stage of the
sexual cycle together after about the third week of May is almost
none. Second nests that are made following early season nest losses
are always in a new location, have fewer eggs than first nests and a
higher proportion of infertility. In most cases the later the second
nesting, the smaller will be the clutch, and the higher the number
of infertile eggs.

Dust Bathing. Dust bathing by the brood has already been dis-
cussed. This practice is just as necessary for the adults as it is for the
chicks, for it seems to be a requisite for proper maintenance of the
feather coat as well as for normal feather growth. Grouse grown or
held in captivity without benefit of dusting facilities usually develop
a ragged coat. Dusting is done mostly from spring to fall with the
greatest activity in the summer. Little dusting is done in the winter
for lack of opportunity, but if the ground allows them to do so they
will dust all year round. The increase in dust bathing in the summer
lends weight to the supplementary theory of need for elimination of
ectoparasites, although whether this is a primary reason for dusting
is doubtful. The same bath is used repeatedly although one bird may
have several in different places.

Biography 55

Relations of Grouse to Other Animals. Except for those animals
that are recognized as enemies, grouse pay little attention to their
woodland cousins. Other birds, as well as the small rodents and in-
sectivores are taken for granted most of the time. Occasionally if a
squirrel or chipmunk comes too close to a brood of young chicks the
mother will drive off the intruder with a whirlwind attack. Some-
times jays will scold at grouse, apparently for theii- own amusement.
Ring-necked pheasants will occasionally parasitize a grouse nest
(see Plate 5C).

Grouse select their nest sites without regard for nests of other
species. They occasionally will flaunt danger and locate the nest in
the vicinity of an enemy. We watched one hen bring off a clutch
successfully within fifty feet of an active red fox den. The fox pups
had played within ten feet of the nest as evidenced by the "toys"
(uneaten animal remains such as a bone or a piece of fur) left there.
Another bud nested successfully almost beneath a sharp-shinned
hawk's nest. Generally, however, they pay healthy respect to those
creatures that they instinctively know to be dangerous.

Adaptability to Changing Environment. That the grouse is to be
found within a few miles of New York City and is scattered in fair
abundance throughout most of the highly populated northeastern
states is a tribute to its ability to adapt itself to changing conditions.
There is little question but tliat certain environmental conditions
combine to make optimum survival of the species. But there is
hardly a single one, or gioup, of tliese conditions that the bird cannot
circumvent to maintain itself in reasonable numbers. In coverts
throughout its range, it is constantly facing changes: lumbering,
forest fires, gi-azing, changes in populations of predators and buffer
species, in unending variety and combination. Grouse populations
may go down as a result of changes in environment, in some ex-
tremes they are even extirpated, but as time wears on the species
adapts itself and continues with surprising versatility.

Adaptability to Artificial Propagation. The story of man's attempts
to raise ruffed grouse in captivity is even older than the profession
of game breeding itself in America. As early as 1750 Bartram (In
PhU. Trans. Roy. Soc. London, 1754) wrote a friend in England to
the effect that the ruffed grouse could not be tamed or raised under
hens. For over 150 years every other attempt met with the same

56 The Ruffed Grouse

quick failure. The first faint hope of ultimate success came in 1903
when Professor Hodge of Massachusetts's Clark University suc-
ceeded in maturing several grouse from wild eggs. Two years later
he passed another milestone with hatching of the first grouse from
captivity-raised parents. From that time until 1916 Merrill and Tor-
rey successively carried on the grouse raising experiments for the
state of Massachusetts that Hodge had begun as a hobby. They
proved conclusively the futility of trying to use wild-caught breed-
ing stock; they succeeded in raising the first second-generation
grouse, but finally became discouraged and released the seventeen
birds remaining in the spring of 1916.

In 1912 the American Game Protective Association became inter-
ested in the problem and engaged Torrey to do their experiments.
After he moved to the Massachusetts state game farm in 1914, the as-
sociation encouraged other experiments. These, however, added lit-
tle to the knowledge of the subject. In 1919 they persuaded Allen
( 1929 ) of Cornell to try his hand at the problem. Thus began the era
of scientific fact finding in game breeding. In twelve years of alter-
nating hope and despair, Allen proved that grouse could be raised in
captivity with some success in small numbers. It was he who took
the birds off the ground, placing them in wire-bottomed pens, thus
partially solving the disease problem.^ Allen uncovered the primary
essentials of a breeding technique with the captive hens and cocks,
so essential in obtaining a high fertility ratio and in preventing ex-
cessive mortality. And in many other aspects of grouse propagation,
Allen developed the first satisfactory techniques.

In 1931, Bump took up the problem in connection with the New
York Grouse Investigation and for several years Allen and Bump
worked cooperatively, with Allen pursuing further definite research
problems and Bump attempting to develop methods for practical
large-scale propagation. In the ten years of work with grouse propa-
gation. Bump raised nearly 2,000 grouse, including birds of the
tenth generation. However, in spite of continually improving meth-
ods and a growing knowledge of the requirements of the species,
grouse propagation is still not economical, dependable, or practical
for restocking purposes.

The vital problems that still must be solved before the raising of

^ Other breeders, with other species, developed the wire floor technique inde-
pendently at about the same time.

Biography 57

grouse becomes a routine success are: increase in number of eggs
laid by breeders ( 10.5 to 16.9 average per hen ) ; ' reduction of inf er-
tihty in eggs (27.8 to 61.9 per cent a year); ^ increase in hatchability
of fertile eggs ( 58.3 to 87.5 per cent ) ; ^ increase in survival of
hatched chicks (16.2 to 37.7 per cent).' In the latter case, the high
mortality occurs mainly during the first two weeks after hatching.

There is little question that these problems will be solved in due
time, but it will likely take many more years. The difficulties in pro-
ducing grouse in confinement are to a considerable extent due to
the relative unadaptability of the species to artificial propagation.
The exacting conditions of the oestrus cycle, the vicious nature of
the bird when not in the proper physiological condition for mating,
its susceptibility to diseases and parasites of many kinds, and the
natural fitness of the digestive system for insects almost exclusively
in the early weeks of life, all contribute to make the grouse a difficult
species to rear. On the other hand, the bird is very tractable in cap-
tivity, even more tame than the pheasant or bobwhite. It is poten-
tially able to lay many eggs; one female examined during the
egg-laying period proved to have 177 eggs visible to the naked eye,
from pinhead size to one ready for laying.

In spite of its tameness in captivity, man-reared grouse readily
revert to natural wildness when released in good coverts. The ma-
jority of game farm grouse restocked in New York became normally
wild within a week after release. Some individuals do not lose their
affinity for man so readily, however. I recall one bird in particular
that ran up to greet a man as if he were a long lost brother, this in a
woodland fairly remote from human habitation and about a month
after the bird had been released.

Farm-reared birds appear to have little trouble in shifting from
captivity food over to their natural diet but do experience consider-
able difficulty in avoiding predator attack. Probably this is as much
due to unfamiliarity with the habitat as with a lack of wariness or
ability.

1 These figures are taken from N. Y. S. Cons. Dept. Ann. Reports for the five-year
period prior to 1941.

58 The Ruffed Grouse

REFERENCES AND CITATION SOURCES FOR THE LIFE STORY OF
THE RUFFED GROUSE

(including general references on the species)

Allen, A. A. Ten Years Experiments in the Rearing of the RuflFed Grouse in Cap-
tivity, Trans. 16th Am. Game Gonf., 1929.

. Sex Rhythm in the Ruffed Grouse and Other Birds, Auk, Vol. LI, No. 2,

April, 1934.

Breeding Season Behavior of the RuflFed Grouse, Trans. 20th Am. Game

Conf., 1934.
Allen, A. A., and Gross, A. O. Report of the RuflFed Grouse Investigation, Season

of 1925-1926, Am. Game, Oct., 1926.
Anonymous. U. S. Gazette of Nov. 18, 1832, as reprinted in Pennsylvania

Game News, Feb., 1933.
Bendire, C. Life Histories of North American Birds, Smithsonian Gontrib. to

Knowl. XXVIII, 1892.
Bent, A. C. Life Histories of North American Gallinaceous Birds, U. S. Nat.

Mus. Bull. 162, 1927.
Bump, Gardiner. RuflFed Grouse Studies, Trans. 17th Am. Game Gonf., 1930.

. Roughing It with the Ruffed Grouse, American Forests, July, 1931.

. The New York Ruffed Grouse Survey, Trans. 19th Am. Game Gonf.,

1932.

Ruffed Grouse in New York State During the Period of Maximum

Abundance, Trans. 21st Am. Game Gonf., 1935.

Clarke, C. H. D. Fluctuations in Numbers of Ruffed Grouse, Univ. of Toronto
Studies No. 41, Biol. Series, Univ. of Toronto, 1936.

Darling, F. F. Bird Flocks and the Breeding Cycle, 1938.

Dwight, J., Jr. The Moult of North American Tetraonidae, Auk, Vol. XVII,
April, 1900.

Eaton, E. H. The Birds of New York, 1910.

Edminster, F. C, Darrow, R., and Bump, G. The First Fifteen Months of the
New York State Ruffed Grouse Investigation, Trans. 18th Am. Game Conf.,
1931.

Fisher, L. W. Studies of the Eastern Ruffed Grouse in Michigan, Tech. Bul-
letin 106, Mich. State Coll., June, 1939.

Forbush, E. H. Game Birds, Wild Fowl and Shore Birds of Mass. and Adj.
States, 1916.

. Birds of Massachusetts and Other New England States, Vol. II, 1929.

Gross, A. O. Annual Report of New England Ruffed Grouse Investigation,
1928.

. Same for 1929, 1930, and 1931.

. The New England Ruffed Grouse and Wisconsin Prairie Chicken In-
vestigation, Trans. 17th Am. Game Conf., 1930.

. Ruffed Grouse and Prairie Chicken, Trans. 18th Am. Game Conf., 1931.

Biography

59

King, R. T. Ruffed Grouse Management, Joum. Forestry, Vol. XXXV, No. 6,
June, 1937.

Mathews, F. S. Field Book of Wild Birds and Their Music, 1904.

Nice, Margaret Morse. The Role of Territory in Bird Life, Am. Midland Natu-
ralist, Vol. 26, No. 3, Nov., 1941.

Reese, Staber W. Mr. Grouse Goes a Courtin', Wise. Cons. Bull., Vol. VII,
No. 7, July, 1942.

Sawyer, E. J. The Ruffed Grouse with Special Reference to Its Drumming,
Roosevelt Wildlife Bull., Vol. I, No. 3, 1923.

Smyth, Thomas. Studies in the Life History of the Ruffed Grouse, Ph.D. the-
sis Cornell University, 1925.

. Sexual Dimorphism in Bonasa timhclhis, Proc. Pa. Acad. Sci. 13:62-67,

1939.

Stoddart, A. M. Ruffed Grouse in New York State, 1918.

Studholme, A. H. Ruffed Grouse Environment in the Scrub Oak-Pitch Pine
Forest Type, M.S. thesis, Penna. State College, June, 1941.

Woodruff, E. S. The Ruffed Grouse, A Study of the Causes of Its Scarcity in
1907, 1908.

4

Shelter

TYPES OF GROUSE RANGE

Grouse range, in its broad sense, is the geographic area that the
species inhabits. The types of grouse range, as used here, are the
major subdivisions of the species range— divisions having common
physiographic, chmatic, ecologic, and temporal factors that affect
grouse populations in an extensive w^ay. Grouse range is a product
of climate, soil, and time, the same as plant growth regions (Van-
Dersal, 1938). On this basis, the grouse range in the northeastern
states may logically be divided into three types— the northern New
York-New England, northern Appalachian, and middle Appalachian.
All three areas are humid and microthermal (having mean annual
temperature between 14° and 0° centigiade ) . They differ primarily
in the dominant plant associations, largely as regards woody plants,
and character of land use as affecting the distribution of these asso-
ciations.

Northern New York-New England Grouse Range. This, the most
northern of the grouse range types in the eastern states, is naturally
the coldest. It has the greatest amount of extensive forest range and
is thus most nearly a wilderness habitat. Its topography varies from
rugged mountains to level, wooded plateaus, all generally well
watered. The winters are very severe, with low temperatures and
heavy snows. The area was almost entirely glaciated during the
last ice age, only the tops of the highest mountain peaks escaped the
ice sheet. Most of this range is relatively uninhabited by man, al-
though practically all the forest has been lumbered at least once.
The sparsity of the human population has enabled the grouse in
many of these coverts to remain today the fool-hen type of bird it
formerly was everywhere.

60

Shelter 61

The extensive areas of continuous forest in this northern habitat
do not produce as high densities of grouse on an average as do the
more southern ranges. The lack of cover type changes ("edges")
well distributed over the range is without doubt one of the major
factors that keeps down the grouse carrying capacity. The severe
climate and the vegetative characteristics contribute importantly
too.

The gross vegetative characteristics of a grouse range are closely
allied with forest types and may best be briefly described by the
tree species associations found therein. The most indicative forest
types in this northern forest range are the spruce-fir group: red
spruce-sugar maple-beech (17);^ red spmce (18); red spruce-bal-
sam fir ( 19 ) ; paper birch-red spruce-balsam fir ( 20 ) ; white spruce-
balsam fir-paper birch (21); balsam fir (22); black spruce (23).
The last is a wet soil type, the others are found on well-drained soils.
The second very important group, though not as indicative of the
range as the first, are the northern hardwoods types, the most preva-
lent of which are: sugar maple (14); yellow birch (15); yellow
birch-red spruce (16); sugar maple-beech-yellow birch (12). Other
common types are: aspen (4); paper birch (6); tamarack (25); and
northern white cedar (24). Associated with these key species are
many more tree species, many shrubs and herbs. Those of most im-
portance to giouse are given in the cover type descriptions later in
this chapter.

The prevalence of the above forest types in the northern range
varies from area to area. Of great significance has been the changes
in extent of types brought about by lumbering— mainly a change
from a predominance of coniferous to hardwood types. This often
resulted in the familiar pattern of hardwood ridges and coniferous
bottoms.

The northern range, except that part extending into Canada, con-
sists of the Adirondack region of New York, the Green Mountain
range in Vermont, the White Mountains in New Hampshire, and
all Maine but the most southern portion.

Northern Appalachian Grouse Range. This middle range extends
tlirough all New York and New England not covered by the northern

'•These are tlie ofBcial Forest Cover Type numbers (Forest Cover Types of
the United States; SAF Committee on Forest Types, Journal of Forestry, April, 1932).
This reference gives associated tree species and variants.

62 The Ruffed Grouse

range, except the coastal plain of Rhode Island and Connecticut, the
majority of Pennsylvania and northern New Jersey, and a few locali-
ties in West Virginia and western Maryland at higher altitudes. It
is faii-ly temperate in climate and has rather severe winters, with a
dependable winter snow cover. Of the three ranges it is the most
densely inhabited by man and, with some exceptions, the woodland
is well broken up with farmland. Much of it was covered by glaciers
during the last ice age. The best of the coverts in this range probably
produce somewhat higher grouse densities than either the more
northern or southern ranges. This is a result of good interspersion
and desirable cover types and moderate climate.

The major forest types found in the northern Appalachian range
include: hemlock (11); sugar maple-beech-yellow birch (12); a
mixture of (11) and (12), which is very common; white pine (9);
white pine-red oak- white ash (8); aspen (4); pin cherry (5); gray
birch-red maple (7), and a mixture of (9) and (12).

To a considerable extent, this range is found on the more rugged
lands, the poorly drained soils, or unproductive agricultural soils,
which have been allowed to remain in woodland or to revert to
woodland because of their impracticability for farming. In some
areas, as in parts of northern Pennsylvania, and the Catskills of
southeastern New York, this condition is so extensive as to allow
almost continuous grouse coverts over many miles, resembling in
this respect the more northern range.

Over much of this range, the change in forest composition as a
result of cuttings for lumber and other wood products has followed
a trend similar to that in the North-an extension of hardwood types
at the expense of the hemlock and white pine. Other changes, as the
growing predominance of beech in many woodlands, are also im-
portant as affecting the ruffed grouse.

Middle Appalachian Grouse Range. The southern range type of the
northeastern states' grouse range extends southward from southern
Pennsylvania and mid-New Jersey to include most of West Virginia,
western Maryland and Virginia. There are a few small areas of the
northern Appalachian type at higher altitudes in this zone and a
few areas of the southern type in central Pennsylvania, Long Island,
and southern New England.

For the most part this range type does not have a dependable

shelter 63

winter snow blanket, and consequently has a moderate climate
compared with the more northern ranges. It is fairly densely inhab-
ited by man and quite broken up with farm land, although less so
than the northern Appalachian range type. There are some areas of
rather extensive forest, as in western Virginia and eastern West Vir-
ginia. It was mostly untouched by the last glaciers. Its better gi'ouse
coverts are highly productive but, other factors being equal, prob-
ably are not quite as densely populated with grouse as are the
middle range. The carrying capacity is generally somewhat higher
than that of the most northern range.

The more important forest types included in the southern range
are: southern red oak-scarlet oak (34); bear oak (35); chestnut oak
( 36 ) ; pitch pine ( 37 ) ; shortleaf pine ( 38 ) ; shortleaf pine-southern
red oak-scarlet oak ( 40 ) ; shortleaf pine-Virginia pine ( 42 ) ; Virginia
pine-southern red oak (43); Virginia pine (44); eastern red cedar
(46); black locust (47); white oak-black oak-red oak (49); red oak-
basswood- white ash ( 51 ) ; red oak ( 52 ) ; yellow poplar- white oak-
red oak ( 55 ) . It is clear that the dominant group in this part of the
range are the oaks. These types, by themselves, are not veiy pro-
ductive of grouse.^ The conifers are primarily the hard pines. In
middle age to maturity the forest is predominantly hardwood, as
these pines are all short-lived, pioneer species.

The grouse range in the southern part of the Northeast, as with
the middle area, is found mainly on the areas not suited to farming.
To a gieater degree than in New York and Pennsylvania, the ruffed
grouse is found in the mountainous country which is often continu-
ous forest except for the valleys. Forest fires and livestock grazing
have been more prevalent in this region than farther north. Another
commonly detrimental factor to grouse here, along with other game
species, has been the prevalence of continuous hillbilly hunting.

COVER TYPES— CHARACTERISTICS AND COMPOSITION

There is abundant evidence that grouse react to differences be-
tween certain types of plant associations and between some age
classes of a plant association. These constitute the cover types of

^ T. E. Clark of Virginia told the author that on the George Washington National
Forest ninety per cent of the grouse are on ten per cent of the range, almost none
in the extensive oak types.

64 The Ruffed Grouse

grouse range, from the point of view of the grouse, insofar as we can
interpret their needs as expressed by their cover choices. The types
most prominent in the northeastern range may be organized as
follows:

Open land— predominantly herbaceous plants.

Overgrown fields— predominantly shrubs and saplings, succeeding
from abandoned open fields.

Slashings— clearcut woodland, prior to reaching the pole stage of
regeneration.

Hardwood woodlands— woods composed predominantly of hard-
wood species.

Mixed woodland— woods composed of a balance of hardwood and
conifer species.

Coniferous woodland— woods composed predominantly of conifer-
ous species.

Many subdivisions of these major types have some importance to
the grouse. These types with the more important subtypes may be
described as follows:

Open land: The great variety of open-field plant associations that
are so vital to such species as the bobwhite quail and ring-necked
pheasant make substantially one type for the grouse. They do not
furnish satisfactory shelter. Some of them do furnish considerable
food, particularly during the summer, but are not very important in
this respect since other cover types fiunish ample food of the same
type and adequate shelter as well.

The greatest value of the open-land types is that they enhance
the value of adjacent woody types by creating edges. The edges
improve the fruiting capacity of woody plants, and many herbaceous
plants from the fields will penetrate the woods edge to a short dis-
tance, thereby increasing the food supply. The abundance of shrubs
is usually greater along an open field edge of a woods.

The edges of open fields are used some by grouse for feeding,
sunning and dusting. Old fields of weeds and grasses are most desir-
able. Most crop fields are of practically no use to grouse. Buckwheat
is sometimes used a little in early fall for food. In forest areas, roads,
lakes, streams, marshes, and beaver meadows serve the functions of
the open-field tvpes.

Overgrown land: Fields abandoned from agricultural use will

Shelter 65

normally succeed to woody vegetation. When enough of the land is
covered with a sprinkling of bushes, saplings, briars and vines to
attract grouse to use it regularly, it becomes overgrown land. This
point of change between open land and brush is sharpest when
about forty to fifty per cent of the ground is covered with woody
plants of significant size, bush size or bigger. The land remains in
this cover type until it develops into a pole-stage woodland, or is set
back in its development by some external action.

The species that compose the overgrown land-plant association
depends upon many factors: the type of soil, fertility and moisture
characteristics of the site, the land use in recent years, presence of
seed-producing plants near by, as well as climate and geographical
location. Often a group of short-lived, light-demanding pioneers
take over the area first— raspberries, poplars, "old field" pine, pin
cheiTy, hardback, and gray birch are examples. These are displaced
in relatively few years by longer-lived shrubs and trees. On fields of
good fertility, a good seeding of subclimax species will often result
with the virtual elimination of the pioneer association stage. I recall
an instance of a complete establishment of red pine the year follow-
ing abandonment of a potato field. Many times one may observe
similar stands of maple, ash, and other long-lived species.

The type of overgrown land is of considerable importance to
grouse. For convenience it may be divided into three subtypes (see
Plate 9). The first varies immensely but in each case is made up
largely of a single species, usually a short-lived one. If the species
is quaking aspen it is a high quality grouse cover type; if pin cherry,
scmb oak, or alder it makes rather good cover too, but if it is gray
birch or hardback it is definitely inferior. The second type may be
described as a mixture of hardwood shrubs and tree saplings, with
few if any conifers. The last important subtype is the same as the
second except that it contains considerable conifers in the mixture,
one third or more of the total.

Among the more important plants found in the last two subtypes
are: thomapples, hazelnut, sumacs, rose, brambles, blueberries,
viburnums, grape, bayberry, huckleberry, and elder among the
shrubs. White pine, red pine, and hemlock are best among the coni-
fers while oaks, apple, cherry, and poplar are among the more useful
trees— all, except the tree oaks, being useful as saphngs. Among the
more useful herbaceous plants found in these types are sedges

66 The Ruffed Grouse

(Carex), strawberry, sheep sorrel, everlasting, asters, cinquefoil,
buttercups, clovers, violets, and barren strawberry.

Slashings : Woodlands that have been lumbered in recent years by
the clear-cutting method serve substantially the same purpose for
grouse as do the overgrown-land types. These differ from over-
gi-own land basically in origin, in that there is a regenerating woody
plant association rather than a new seeding of woody plants in an
herbaceous association. This results in a large proportion ( except in
coniferous stands ) of coppice growth and a relatively small propor-
tion of seedling plants. The species in a slashing association also
normally vary widely from the overgrown-land groups. Both the
woody species and herbaceous species in a slashing are to a consid-
erable degree those that were present in the woodland before cut-
ting. There are fewer shrubs and more tree saplings than in
overgrown land. The newly seeded species often are quite different
too. The use of fire in disposing of brush following the cutting fre-
quently brings in a thick stand of Rubus, pin cherry, or other species
that germinate well after burning. On the other hand, slashings have
fewer of pioneer woody plants, grasses and field herbs than does
overgrown land.

The prime characteristic of the slashing, therefore, is that it is a
woody plant association with few, if any, trees. A woodland set back
in its plant succession, the few trees that may remain are usually old
gnarled stubs that were not worth taking out for wood products.
Another characteristic of many slashings for a few years after cut-
ting, is the presence of dead brush— the tops of the cut trees. This
often results in a tangle of stumps, brush, and tree tops that furnishes
very desirable shelter.

Slashings seem to be divided into two types, from the point of
view of grouse use, primarily on the basis of age (see Plate 10). One
we may call the briar-stage slashing, the other the sapling-stage. In
the first instance the slashing operation removes the trees, releases
many shrubs and herbs (which, however, may suffer from this re-
lease), and induces the germination of new species of herbs and
shrubs. Various species of blackberry and raspberry are usually
prominent in this stage, hence the term briar stage. Depending on
several factors, mainly the composition of the original woodland,
soil type, degree of burning, and aspect, this stage lasts from three
to ten or more years. As hardwood saplings develop, either as cop-

Shelter 67

pice or seedlings, they gradually displace the briars and herbs and
finally become a thick stand of saplings. This is the second type. It
is generally somewhat less useful to grouse than the first, especially
for young birds. When the saplings grow into the pole stage, about
three or four inches diameter at breast height (d.b.h.) the stand
becomes a woodland again. This stage generally lasts from fifteen
to thirty years after cutting.

The species that are prominent in slashings have already been
indicated to some extent. Brambles, pin cherry, coppice of the
former woodland hardwood species, and popple are among the
commoner woody species. Grasses, ferns (especially bracken), fire-
weed, wild lettuce, and other rank-growing tall herbs may also be
prominent. The conifers are usually absent. Unless the original
stand had a coniferous mixture in the understory, the slashing is apt
to be all hardwoods. In many northeastern woodland areas the
native conifers do not seed in readily in clearcut areas. The defi-
ciency in conifers following lumbering is among the most serious
problems in both forestry and vdld-life management.

Hardwood woodlands: Woodlands predominating in hardwood
species constitute a high proportion of northeastern gi-ouse range,
particularly in the southern portion. To a degi'ce, grouse need this
t\'pe of cover in their year-round life, but as a general utility type it
is seriously deficient in protective shelter— that is in conifers. Hence
the vast extent of this type indicates one of the major faults with the
northeastern range.

The hardwood woodland type may be divided into two or more
major subtypes from the standpoint of grouse use; the main criterion
is the age class. The mature hardwoods, averaging about twelve
inches or more d.b.h., are distinguished also by the sparsity of the
undergrowth. Second-growth hardwoods, those from pole stage up
to the mature stands, constitute the second subtv'pe. Here the under-
story is usually well balanced. Small openings scattered through
make an improved subtype of either age class.

The density of hardwood types varies considerably. In the mature
type, the stem density is usually sparse but the crown density close,
unless selective cutting or windfalls have made some openings. The
younger stands are progressively more dense in stems but usually
more open in the crown. In either case, those with scattered glade-
like openings are best. Corollary to the crown density, the herbaceoj

^t'fLlBRARY

<^

68 The Rufted Grouse

ground cover is more luxuriant when the crown allows plenty of
light to penetrate to the ground.

There are many other variations of the hardwood type, some of
considerable importance to grouse. If the woods is pastured, the
woody understory is likely to be absent; hence this factor may be
important in grouse range. The species composition may be very
important depending mainly upon the inclusion of staple food pro-
ducers. Some of the more prevalent hardwood compositions in the
Northeast are: northern hardwoods (birches, maple, serviceberries,
poplar, black cherry, hophombeam); northern Appalachian hard-
woods (beech, birches, maples, cherries, poplars, serviceberries,
oaks, hophombeam, hornbeam); Appalachian hardwoods (oaks,
hickories, gums, serviceberry, hophombeam); mixed oaks. These
are the more prominent tree species in the hardwood types that
occur in plant growth zones from Canada to Virginia.

The shrubs found in the understory of these subtypes vary too. In
the North, witchhobble {V. alnifolium) , withe rod (V. cassinoides) ,
nannyberry (V. lentago), are prominent; in the middle region,
mapleleaf viburnum (V. acerifolitim) , poison ivy, blueberries, grape
are among the commoner species; while to the south flowering dog-
wood, mountain laurel, greenbrier, blueberries, honeysuckle, poison
ivy, and grapes are common. There are also minor differences in the
ground cover. Bunchberry [Corrms canadensis), ferns, Canada may-
flower, violet, wood sorrel, jewelweed, shinleaf, miterwort, false
miterwort, skunk cabbage, and hepatica are significant in the North;
while to the south the ferns, lespedezas, hepatica, alumroot, skunk
cabbage, wood sorrel, and false miterwort are among the more im-
portant plants.

Mixed woodland: The difference between the mixed woods and
the hardwood types lies primarily in the tree composition, and in
the proportion of conifers in the mixture. From the standpoint of
grouse needs, the hardwood types are deficient in conifers. Most of
the areas of the hardwood type have no conifers at all, or but a
scattered few. These areas are clearly conifer-deficient, but what
about stands that have five per cent, ten per cent, fifteen per cent and
so on, of conifers? Since we cannot interpret the reactions of grouse
to shelter with such fine accuracy, and since these reactions will vary
under different conditions, we have to be somewhat arbitrary in
drawing the line between the two type groups. However, twenty
per cent of conifers appears to be a proper proportion.

shelter 69

Of course, the value to grouse depends on the character of the
coniferous element, too. Whether the conifers exist primarily in the
understory as reproduction, primarily in the crown as mature trees,
or in all age classes is of great significance; the latter is most satis-
factory, the second class poorest ( see Plate 13 ) .

Just as we find it necessary to distinguish the line between hard-
woods and mixed woods, so we must also define the point of change
from mixed woods to coniferous woods. This criterion appears to be
about seventy per cent of conifers. The mixed woods is thus a bal-
anced stand containing hardwoods and from twenty per cent to
seventy per cent conifers, based on area of crown coverage. It pro-
vides grouse with food and shelter for all seasons, although the cover
in summer is imperfect.

The subtypes of mixed woodland follow the same general distinc-
tions as the hardwood types: differences in age class, in prevalence
of small openings, and in species composition. The mature and
second- giowth stands are distinguishable as in the hardwoods, the
critical point being in the neighborhood of an average twelve inches
d.b.h. The species associations follow the hardwood types with coni-
fers added: in the North, spruce, red pine, and balsam fir grow in
combination with the northern hardwoods; white pine, hemlock, and
sometimes Banks' or other pines in combination with the beech-
birch-maple group; the southern oak and hickory types contain
some white pine and hemlock at higher altitudes, but, toward the
south, include more of the hard pines, especially Virginia pine,
pitch pine and shortleaf pine.

The shrub species are substantially those in the hardwood stands.
In the ground cover we find some significant variations, notably the
addition of partridgeberry, wintergreen, and club mosses {Lyco-
podium).

Coniferous woodland: Woodland, predominately coniferous trees
(excepting tamarack), having less than about thirty per cent of its
crown taken up by hardwoods, constitutes the last of the important
northeastern type groups. In young stands, this type is usually very
deficient in food though well provided with shelter, but in mature
stands there is usually a hardwood understory of significance. Sub-
types, as with the last two type groups, are based on age class and
species composition ( see Plate 15 ) . In addition to the mature stands
and the second-growth stage, it is sometimes desirable to add a third
ageclass subtype, the pure, one-species reproduction ( or reforesta-

70 The Ruffed Grouse

tion) stands after the stage when briars and other hardwoods are
present and before it reaches the pole size. This subtype could be
considered a phase of the overgrown-land type but since its function
for grouse is more associated with the coniferous woodland group
it is included here.

The important species subtypes are the same as given for the coni-
fer element of the mixed woods subtypes, from the spiiices and fir
of the North to the hard pines of the South. The understory and
ground cover of the coniferous types are generally quite different,
however. Shrubs are usually sparse or absent, and limited in species
when present. In mature stands the understory of shrubs and hard-
wood tiees is more complete and also more varied in species. The
ground cover is quite scanty, too, and varies markedly from that
found in the hardwood types. Amongst a heavy ground litter of
needles and leaves, we find wintergreen, some ferns, partridgeberry,
some orchids, bunchberry, club mosses, blueberries, polygala, and a
few others.

INTERSPERSION OF COVER TYPES

We have discussed the character and quality of the major types of
grouse cover. But regardless of quality, an extensive area of any
cover type is too much. And so it is with grouse cover. Large units
of a single type are relatively unproductive of grouse, no matter
what the type. That fact brings us to the importance of aiTangement
of types, and the need for interspersion. Interspersion as applied to
cover types refers to the mixture of different types in small enough
units so that the game species may conveniently make use of them
day by day or season by season as needed. The need for two or more
types within a short range is brought about by two basic facts in
grouse ecology; the bird requires several different types for satisfac-
tory year-round range and it prefers edges between woods and
overgrown land or slashings, woods and open land, or between over-
grown land or slashings and open land, rather than the deep interior
of any cover types. These are basic requirements for most wild-lffe
species.

It is clear then, that the consideration of the character and carry-
ing capacity of any grouse range must give full weight to the arrange-
ment of the cover types and their degree of interspersion. Likewise

Shelter 71

management of grouse range must take cognizance of these prin-
ciples.

PLANT SUCCESSION CHANGES COVER TYPES

Grouse range is an unstable, ever-changing complex. The plants
change with the years, and as they mature, particularly the tree spe-
cies, the character of the cover types gradually changes. Not only do
the plants grow, and thereby alter the physical nature of the cover,
some of them die and are replaced by new plants, often different
species. Thus the composition of species in a cover type changes
too. This phenomenon of plant succession follows definite laws.
Grouse management must be based largely on these laws.

In order to understand the character of plant succession as it
affects grouse habitat, let us consider the history of an abandoned
farm field next to a woodland. To be specific, let it be a timothy
meadow which has about run out due to lack of fertilizer and loss
of topsoil by erosion. Many annual and perennial weeds have al-
ready replaced a part of the timothy at the time of abandonment.
Upon going wild these weeds gradually displace most of the timo-
thy and the field is taken over by goldenrod, daisies, deWl's paint-
brush, poverty grass, wild carrot, and associated plants. If the field
had been recently ploughed before abandoimient the weeds would
be primarily annuals for a time, such as ragweed, foxtail, and smart-
weed, which would then be succeeded by the perennials.

The decline in fertility may continue for a time after abandon-
ment and portions of the field may be so poor as to be barren of
vegetation, or merely support mosses. But sooner or later the first
woody pioneers begin to take hold— often dewberries and blackber-
ries at first, or sometimes sweet fern, or plants like quaking aspen or
fire cherry. Up to the time these woody plants attain some size and a
fair density, the cover is essentially of one type so far as the grouse
is concerned— open land. But as the brushy type takes over, the area
becomes overgrown land, with an entirely different prospect for
grouse.

The changes that take place during the overgrown stage depend
upon the species that are initially established and, as in all stages of
succession, upon the distribution of mature seeding plants in the
vicinity ( see Plate 17 ) . Generally the composition gradually changes

72 The RufiFed Grouse

to subclimax or climax ^ species while the pioneers are maturing and
dying out or are being crowded out. Sometimes a mixture of moder-
ately tolerant shrubs, like the dogwoods, viburnums, thornapples,
alders, and others takes over as intennediate between the earliest
woody plants and the later tree saplings. This stage is particularly
useful to grouse, especially if some conifers too are in the mixture.
Another alternative that occurs commonly in some portions of the
Northeast is a dense stand of pine. This usually follows the weed
stage, or sometimes tlie early shrubs.

As the long-lived tree species gain height and crown, the shrubs
and pioneer trees are replaced to form a subclimax pole-size wood-
land. This is somewhere between twenty and fort)^ years, or even
more, along the path of change since the field was abandoned by
man. This type is again quite different from the overgrown land as
grouse cover. It may be any one of the three woodland types, hard-
woods, mixed woods, or coniferous woods. The rate of change in
type now slows down greatly. A new stage, except for age class
change, may not develop for many decades. Oft times there is a
gradual change in species composition, such as the replacement of
hard pines with hardwoods, or the introduction of hemlock into a
stand. Thus the woodland reaches its climax type. Changes then are
brought about by the dying of large trees and the ensuing plant
succession from the briar stage in the small openings formed.

Actually this unhampered trend of plant laws seldom runs a full
course. Catastrophes occur that set back the successive stages and
prevent the climax being reached. Fire, either natural or man-made,
may sweep the woodland and bring the land cover back to an herb-
briar stage somewhat different from that described above, one domi-
nated by plants that germinate well following a burn. More com-
monly, man steps in with ax and saw, and reduces the woodland
to a slashing or to a spotty, small-clearing pattern if he pursues
selective cutting. In either case, the succession is set back many
decades, often a century or more. If he follows clear cutting with
stump pulling and ploughing, or if he grazes the cutover area with
livestock, the stage is placed still farther back. Then too, grazing,
even without lumbering, will materially alter the character of the

''■ Climax plant associations are those that complete the stages of plant succession
for an area and, unless disturbed, maintain themselves indefinitely. Subclimax species
are those in the stage preceding the climax.

Shelter 73

woodland, eventually reducing it to grass or weeds if continued
intensi\'ely beyond the lives of the crown trees.

This is just one story of plant succession, with some alternatives
discussed. Many other similar trends of plant changes occur under
other conditions. But whatever the conditions, the potentialities of
ruffed grouse management are bound up in these principles. Thus
sound management should be based upon the manipulation of plant
succession by means of man's tools— axes, saws, planting irons, fire,
fencing, livestock, and others.

RELATION OF SHELTER TO THE GROUSE NESTING

Cover Types Used for Nesting. Most hen grouse shift their scene
of activity from a predominance of coniferous types in the winter
and very early spring to a predominantly hardwood type in nesting
season. This change to hardwood types on the part of nesting females
is much more pronomiced than the same trend by other grouse. The
predominantly coniferous types— most important through the winter
—are practically abandoned as nesting territory. Fewer than one
grouse in twenty uses a conifer type for the nest location.

A medium-age stand of mixed hardwoods is the most commonly
used nesting type in New York, usually with a scattering of conifers.
Closely following is the medium-aged stand of mixed hardwoods
and conifers. These types were selected for nesting in two thirds of
the 1,270 instances observed (see Table 1) (Bump, 1938). When
the records are weighted ^ to account for the variation in extent of
the different cover types on tlie area, these types are still predomi-
nant but the slashing types also rise to prominence. From the ratios
of nests to acres of types, we conclude that the three types— predomi-
nantly hardwoods, mixed woods, and slashings— are the best types
for grouse nesting. The variation in acceptance between them is not
significant. Open land is of no value for nesting, while the over-
grown-land types and coniferous types are used only to a small
extent, far less than their proportional existence.

Among the desii-able hardwood t)^es and mixed hardwood-coni-
fer types, the middle-aged stands are superior as nesting cover over
the mature stands by a very significant margin.

''■ Number of records is divided by acres of cover type to provide comparable data.

74 The Ruffed Grouse

Table 1

Cover Types Selected by Grouse for Nesting (Based on
1,270 Nests in New York, 1930-36)

Percentage of Nests Ratio of Nesting Use
Found in Each to Cover Type

Cover Type Type Prevalence ^

Open land 1.4 .002

Overgrown (brush) land 9.8 .05

Hardwood woodland 45.2 .12

Hardwood-conifer mixed woods 21.5 .13

Coniferous woodland 4.1 .07

Slashings 9.9 .13

Cover type not ^ Norm, including open land .09

specified— 8.1 Norm, excluding open land .11

Ratios based on Connecticut Hill cover typo
proportions.

Specific Location of Grouse Nests. Two thirds of the nests I have
observed vi^ere located at the base of trees, with about two thirds of
these being hardwoods and almost all trees of considerable size ( see
Plate 18 ) . Of the other third, the majority were at the base of a tree
stump, with a few located beside or under logs, bushes, or brush
piles.

There appears to be no significant selection among tree species
for nest location. The species chosen are those predominating in the
woodland. These were mainly beech, maple, birch, hemlock, pine,
chestnut (dead), oaks, etc. A somewhat disproportionate number of
nests at the base of dead chestnuts seems to be due to the additional
sunlight reaching the ground with the absence of the tree canopy
rather than to the species of tree.

If we may presume to interpret the desires of a hen grouse for a
nesting location on the basis of having observed several hundred of
their choices, the selection appears to be a compromise between ( 1 )
a position protected from attack by enemies; (2) good visibility of
the immediate terrain; (3) protection from the elements; (4) desire
for direct sunlight; (5) proximity to edges; (6) lack of obstructions
in the way of quick escape. The result is a fairly exposed situation
with a solid backing, and near the edge of a prominent cover type.
A spot at the base of a tree, or near a stump or log in a relatively
open woodland, and near a road, field, or thicket seems to meet
requirements.

Shelter 75

The setting bird always faces outward with her tail raised against
the tree. From this attitude she can view her surroundings in all
directions except the rear, where the tree gives protection. Usually
she chooses a tree or stump that has few, if any, bushes or saplings
immediately in front for several feet— thus giving an open front yard
—but often a very small sprig of living or dead brush will protrude
a few inches above and in front of the nest to assist in obscuring her
position. The desire for sunlight results in many nests placed along
old woods roads, beside stumps where the canopy has been broken
by the removal of a large tree, at the base of dead trees (notably
chestnut), and in similar spots where the advantages of the wood-
land are combined with more than ordinary exposure to the sun.

Most grouse habitat is quite free from evidences of man's civiliza-
tion. However, if its cover requirements are met, the grouse pays
little attention to such intrusions. Sights, sounds, or movement of
mechanical objects, as long as they are regular, cause no fear. Sev-
eral grouse nests I have observed bear witness to this fact. One was
located on the bank of a railroad right of way, protected only by a
bracken fem, and within a few feet of the track. The nest was dis-
covered by a keen-eyed railroad man who spotted the bird on the
nest from the train. Another nest, in the oil well region of southwest-
em New York, was situated beneath the iron drive-rod that connects
the powerhouse with the well. This rod, supported every little ways
with tripods, was only about three inches above the bird's head.
About four hours each day, when the wells were being pumped, this
rod squeaked back and forth without causing the bird concern.
Nests are occasionally found on the borders of public highways.
The passing of speeding automobiles a few yards away does not
seem to worry a grouse hen.

In selecting a nest site, the bird usually chooses one with a very
open undergrowth, as has already been noted. The crown density
of the nest sites observed also appears to average a little on the open
side but generally about in proportion to what the woods contains.
A very large majority of the nests I have seen were within a hundred
feet of a coniferous type, or a clump of conifers, in spite of the rela-
tive unimportance of conifers to nesting. The presence of a significant
amount of evergreen cover in the vicinity seems desirable for pro-
tection from the weather even though coniferous cover is little used
for nesting and a thick growth of conifers around the nest is avoided.

76 The Rufied Grouse

Location of Nests with Respect to Openings. With the exception of
slashings, which have as high a degree of nesting use as any type,
the openings (cover types without crown cover) are httie used for
nest locations. However, they do determine the location of nests in
other types to a considerable degree. The need for "edge" places
most nests close to an opening. More than half the nests observed
were within fifty feet of such an opening, and almost three quarters
were witliin a hundred feet. Very few were located more than three
hundred feet away from an open field, brushy area, slashing, or a
significant road opening. Most of the nests are nearest to an open
field or slashing type of opening, and the proportion placed in or
near slashings is significant.

The occurrence of grouse nests close to roadway openings in the
woodland, usually wagon roads, is notable. Nearly one third of all
nests were within fifty feet of a road. Although it is not certain that
the road location affected the nest site selection in all cases, it was
the closest opening by a considerable distance for many of the nests.
The woods road seemed to provide an adequate opening and still
avoided the exposure of a wide open edge.

Effect of Slope on Nest Location. The only effect of degree of slope
on choice of nesting sites is a relative avoidance of the very steep
slopes (twenty per cent and over). Only about one half of the nor-
mal proportion of nests are found on steep slopes. The moderate
slopes have a proportionally higher number of the nests while the
flat land has its proportionate share.

The slight variations in proportions of nests on the various ex-
posures are not significant.

Does Cover Affect Nest Mortality from Predators? The percentage
of nests broken up varies slightly according to cover type, tending to
be higher in slashings and conifers. Since these conclusions are based
on relatively few observations, it is doubtful if tliey are dependable.
It is interesting to note, though, that the seemingly safer cover types
are generally the preferred types, and one type ( conffers ) with the
higher mortality rate is used to a lesser degree. It is at least possible
that the predation hazard may affect the choice of type of cover for
nesting.

There also is a possible correlation between midergrowth density
and nest loss, but here again the evidence is not abundant enough

shelter 77

to be fully reliable. The majority of nests are iii cover with open
iinderstory and the mortality in these is average. The mortality in
cases having a dense understory was very excessive. The possibility
exists, and is logical, that a dense imdergrowth tends to increase
nest mortality. Again, the habit of the giouse in avoiding nest sites
where the understory is dense may be conditioned by the predator
hazard.

RELATION OF SHELTER TO GROUSE BROODS

Cover Type Preferences of Grouse Broods. Families of young
grouse observed up to tlie end of September may be considered as
broods. Since the location of the family and its activities are deter-
mined by the needs of the young birds, the mother bird is considered
a part of the gioup. So many of the requirements of the young birds
differ from those of the adults that it would be likely that their
shelter requiiements would di£Fer; and since their food habits vary
markedly from the old birds, the cover types chosen also may well
vaiy considerably. What are tlie cover type preferences of the
chicks as they grow up, and how do they vary from the cover chosen
by adults by themselves in the summer? The records of 1,515 brood
observations in New York from 1930-1936 are given in Table 2
(Bump, 1938).

Table 2

Cover Types Selected by Grouse Broods in New York, 1930-1936

Flushes per Acre Reduced
Percentage of Total to Part of Most Used Type
Cover Type Flushes Taken as 1.00 "^

Chicks Adults

Overgrown land 44.7 1,00 .61

Slashings 12.4 .65 .72

Hardwood woods 25.5 .22 1.00

Coniferous woods 4.2 .22 .17

Mixed woods 10.4 .17 ,25

Open land 2.9 .06 !o3

It is clear that the tendencies in cover type use in summer by
young and old biids is similar-a definite preference for the brushy
types, primarily overgrown land, followed by slashings. This tend-

^ Figiires derived from Connecticut Hill data. See Bump, 1938.

78 The Ruffed Grouse

ency is very much more pronounced with the broods, however.
When the figures are corrected to a per acre basis, these two types
prove to be far more intensively used than all others. Correspond-
ingly, the use made of the woodland types is much lower than for
adults.

The cover used by broods may be divided into two types of major
significance, brushy cover and woodland. Differentiation in use of
the several woodland types is of little significance. The greater pref-
erence for overgrown land as compared with slashings is of consid-
erable significance. It is one of the major explanations of the greater
grouse-carrving capacity of separated coverts as compared with
continuous forest cover. The extensive forest has very little of this
overgrown border cover, and the only approach that the forest has
in this direction is their slashings and the occasional alder runs along
stream courses. The need of brushy cover for broods also poses one
of the biggest problems in grouse management,— and offers one of
the biggest opportunities for raising the carrying capacity in many
coverts.

A more detailed analysis of cover used by broods brings to hght
further interesting trends, all of them consistent with the principle
that hardwood bushy cover is best for broods. Of the overgrown
land subtypes, that which contains a high proportion of young coni-
fers is used less, the mixed hardwoods and pure stands of popple
prove to be used more than the type as a whole. Likewise, hardwood
woods that have been "spot-lumbered" have a use rating nearly
double that for the entire type group. As would be expected, the
younger woodland stands are much more used by broods than are
the mature woods. In hardwoods the comparison is about four to
one for stands under twelve inches d.b.h. compared to older stands,
while in mixed woods the use intensity for the younger stands is over
twice that for mature woods.

Corollary to the predominant use of the brushy types by broods
is the relation between the location of edges of brushy or open land
for the broods that are actually found within woodland types. Most
of them are found within fifty feet of such an edge and records of
broods over a hundred feet from an edge definitely are unusual.
Thus the interspersion of the more open types is of the greatest
importance in determining the productivity of the range.

The changes in cover type use by broods from birth to the time

Shelter 79

when they reach maturity are in accordance with the relations of
brood cover to adult cover. The younger the brood, the more strongly
is its tendency to adhere to brood-cover selection habits, and the
older the brood the greater the chance that the cover types selected
will follow adult habits. The high use intensity of overgrown land
is most prominent in the first six weeks, and drops off rapidly after
that time. Similarly, the use of the woodland types increases after
midsummer, except for that of coniferous woods, which does not
increase materially in use intensity until autumn. Slashings are used
rather consistently at all ages by the chicks, as would be expected
since the use intensity of slashings by the adults is about the same
as it is for the broods.

CHANGES IN COVER TYPES UTILIZED AT DIFFERENT
TIMES OF DAY

As is the case with the adults, the cover types used by the broods
vary considerably at different hours of the day. These changes are
the result of the needs of the birds for food, for roosting cover, for
dusting, or for sunny spots, all of which vary from morning to night;
and in many instances are the results of reactions to weather con-
ditions which vary according to time of day, notably the cold at
night and the heat at midday.

The most important brood-cover type, overgrown land, shows a
marked increase in use intensity throughout the day, the use in
early morning being a little below, and that in late afternoon cor-
respondingly above, the mean. In slashings the tendency is toward
highest use at midday and early afternoon, and lowest use at late
afternoon. These trends in cover-type use by broods are at some
variance with the habits of the adults ( see page 92 ) . The old birds
use overgrown cover most in early morning, least at midday, and
normally in the afternoon; and slashings least in the morning, high-
est at midday.

The use of hardwood woods increases from morning through mid-
afternoon, then drops off at evening. The latter trend is opposite to
the habit of adults of increasing the use of hardwoods towards eve-
ning. With mixed woods and conifers the trend is complementary
to the rest, a higher use intensity in the early day and falling off
later.

80 The Ruffed Grouse

THE EFFECT OF SLOPE ON SELECTION OF COVER TYPE

Many factors enter into the choice of cover made by grouse broods.
Under most conditions the steepness and aspect of the slope is prob-
ably less important than some other factors. However, slope is of some
importance, directly or indirectly, in its relation to cover, plant
species, and weather (see page 93). The effect of slope on brood-
cover selection is essentially the same as for adults at the same season.
Steep slopes are generally avoided, even more so than with adults.

COVER SELECTION IN RELATION TO OPENINGS

The great importance of open cover types and openings (open
land, overgrown land, slashings, and roadways are classed as open
types or openings) to grouse is most strongly shown by the young
birds. While grouse are usually thought of as birds of the wood-
land, their need for openings and brushy cover is clearly indicated.
Three quarters of all grouse broods are found in or within a few
feet of an opening or open cover type, and almost all stay within
fifty feet of openings.

Thus the young biids utilize a relatively small proportion of the
cover in most areas, especially forest areas. The need for intersper-
sion of the open types with woodland types is apparent if the range
is to have a high grouse production capacity.

COVER TYPES CHOSEN AS AFFECTED BY W^EATHER
CONDITIONS

Cover-type selection by broods is somewhat more stable under
varying weather conditions than is the case with the grown birds.
The most important type used, overgrown land, does not vary sig-
nificantly in use under any conditions of temperature, sky, or wind.
It is thus an all-weather type for birds in summer, which is prob-
ably one of the factors bearing on its importance.

Changes in temperatures affect cover type selection the least of
the three weather attributes. As usual the most significant changes
occur in the use of confferous woods. These are used much more
than average on hot days, and correspondingly less on cold days.
Although this relative preference for conifers as a place of escape

shelter 81

from the heat is significant from the standpoint of use of this type,
it is of httle importance to the birds, since the type is so Uttle used.

Sky conditions produce no significant changes in brood cover
selections in the types most used except for a pronounced avoidance
of slashings in the rain. On the other hand, there is a very pro-
nounced increase in the use of coniferous woods during rain. This
indicates the need for heavy shelter during summer rains.

Differences in wind conditions produce several deviations in
cover selections worthy of note, mainly in the use of conifers. Here
we find the pines and hemlocks used primarily in windless weather,
and used well below average under moderate and strong winds.
The explanation probably lies in the association of windless weather
with hot days in the summer, and breezy weather with cooler days.
A reduction in use of slashings during strong winds and a high use
intensity of hardwood woods on windless days are both significant.

Probably the two most important items in these weather rela-
tionships are ( 1 ) the indicated need of broods for coniferous cover,
even in the summer, for escape from heat and rain; and (2) the
weakness of the slashing type as an all-weather cover in compar-
ison witli overgrown land. It is notably less used during rain and
strong winds.

RELATION OF SHELTER TO ADULT GROUSE

Cover Type Preferences of Adults. The cover types in which adult
grouse were flushed in the several seasons of the year on Connecticut
Hill for 13,553 records taken from 1930 to 1936 are shown in Fig. 3.
These do not include females with broods in summer.

Let us first consider the preferences of grouse during each season.
In order properly to evaluate the data it is necessary to weight them
according to the availability of each of the cover types. Taking the
winter (January to mid-March) records first, we find that the num-
ber of records per acre of each cover type, when reduced to parts
of the major type taken as 1.0, are:

Coniferous woods

1.00

Mixed woods

.62

Hardwoods

.46

Overgrown land

.38

Slashings

.22

Open land

.02

82

The Ruffed Grouse

The preference for coniferous woods during the winter is shown
to be one of the outstanding grouse cover requirements. No other
consideration at this season can compare with this need for heavy
protection cover. The second preference, that of mixed woods over
all hardwood types is also significant, and is merely a repetition of
the need for conifers. The difference between the various hardwood
types— the hardwood woodland, overgrown land, and slashings— is
less important, except that slashings are even less useful than the
others. The open land, as at all seasons, is not significant as a cover

E EH F FH
TYPE OF COVER

Fig. 3. Analysis of Certain Cover Requirements of the Ruffed Grouse in New
York State (taken from G. Bump, 1938). Symbols used for type of cover are:
A— open land; B— overgrown land, single species; C— overgrown land, mixed
hardwoods; D— overgrown land, hardwoods and conifers; E— yoimg hardwood
woodland; F— mature hardwoods; G— spot-lumbered woodland; EH— young
mixed hardwood-conifer woodland; FH— mature mixed hardwoods and conifers;
H— coniferous woods; I— briar-stage slashing; J— sapling-stage slashing.

type although very important in improving adjacent types by pro-
ducing edge. Thus, for the grouse, there are three distinctive win-
ter cover types: First, and most vital, is the confferous woodland;
second in importance comes the mixed hardwoods and conffers;
and thirdly the various types, brushy and woodland, that are pre-
dominantly hardwood. The latter are important in winter primarily
in proportion to their proximity to coniferous or to mixed woods.

Shelter 83

They are predominantly food-producing types in the winter and,
except in the portions close to conifer shelter, are not used inten-
sively at this season.

In the spring the choice of cover types is substantially the same
as in winter except for the hardwoods type as we see from the
following comparison of records. In addition to the predominant
use of hardwoods, the great preference for coniferous woods, in
proportion to mixed woods, is much reduced. The differences be-
tween the intensities of use of overgrown land and slashings is not
significant.

Hardwoods 1.00 Use intensity ratio

Coniferous woods .73 when reduced to

Mixed woods .69 parts of major type

Overgrown land .55 taken as 1.0

Slashings .45

Open land .03

In comparison with the use made of the same type groups in
winter, we see that both the mixed woodlands and the several hard-
wood types have greatly increased in use in proportion to conifers.
The coniferous woods is now second to hardwoods as the most in-
tensively used type and is used much less than during the winter.
The use made of overgrown land and slashings is increased con-
siderably as compared with the winter period and during the last
half of the spring period assumes great importance. For the spring
period we may summarize by saying that an approximately equal
requirement exists for coniferous types and for hardwood types, and
that mixed woods is extensively used as a compromise containing
both of the required elements well balanced. As has been shown
elsewhere in the discussion of nesting cover, the female definitely
makes use of the hardwood types for the most part after the middle
of April.

By the arrival of summer, the intensity of use of the various cover
types has changed immensely. No longer is the coniferous element
of great importance. In its place shrubby, brushy, sunny cover-
the open types of overgrown land and slashings— is preferred. Coni-
fers, while used less extensively than any other type group, except
open land, are still used some, mainly to escape excessive sun heat
and summer storms. The intensity of summer use of the type groups
is as follows:

84

The Ruffed Grouse

Hardwoods

1.00

Slashings

.72

Overgrown land

.61

Mixed woods

.25

Conifers

.17

Open land

.03

Use intensity ratio
when reduced to
parts of the major
type taken as 1.0

The significance of the difference in use intensity between slash-
ings and overgrown land is questionable. Together they constitute
the most important type of summer cover— the woody cover with-
out a tree canopy or crown— although the hardwood type is most
used as a single type. The difference in use made of mixed woods
and conffers is probably not significant. It is worthy of note that
in spite of the trend toward more open cover in the summer, the
use of open land is less than at any other season.

Conifers

-

q^

jy^^

Mixed Woods

-

'^•-•-. .y^

^CX^

^ Hardwoods

>-
t—

cc

UJ

-

€^^^

g Slashings
o

"

<^ K'-^—-^

\.^^ \

"~^'~-- —

Overgrown

-

2^ 5PR\NG^,.\,..^'

FALL ]r- = «

Open

-^^'^

fT^"— — —

I.. . J 1

1.0

0 -1 .2 .3 .4 .5 .6 .7 .8 .9

PROPORTION OF MOST INTENSIVELY USED TYPE (TAKEN AS 1.0)

{From Connecticut Hill, N.Y. data)

Fig. 4. Intensity of Use of Cover Types by Adult Grouse by Seasons.

In connection with all these summer records, it should be noted
again that only males and broodless females are included. The
brood mothers, whose choice of cover is presumably determined by
the needs of the brood, are omitted. Since the brood records are
even more predominantly in the overgrown-land and slashing types,
the addition of these to the adult records given here would only
further accentuate the trends shown. We may sum up the summer
cover requirements of the adults by dividing them into two pref-
erence groups: the open types, slashings and overgrown lands, that
are preferred under normal weather conditions; the crown types that

shelter 85

are used some for normal conditions (mainly hardwoods), and the
mixed woods and coniferous types used particularly for escape from
extremes of heat or storm conditions.

With the advent of autumn the choice of cover types begins the
trend back toward winter conditions with a greatly increased use
of coniferous and mixed woods. The slashing, with its abundance of
brambles for summer food, wanes in importance but overgrown
land reaches its highest degree of use due to the abundance of
fall-fruiting shrubs in these types. Once more, as in spring, there is
a significant increase in the use of open land although this never
rises to a position of great importance. The use intensities of the
cover types in the fall are:

Overgrown land 1.00 Use intensity ratio

Hardwoods .88 when reduced to

Mixed woods .80 parts of the major

Coniferous woods .76 type taken as 1.0

Slashings .52

Open land .04

The great importance of coniferous cover at this season is prob-
ably a reflection that the last of the fall season as used here, all
or most of December at least, is really winter so far as the grouse
is concerned. The choice of cover in the early fall tends more to the
hardwood groups, particularly overgrown land where such de-
sirable fall foods as hawthorn are to be found.

It appears that the intensity of use of cover-type groups in the
fall is significant from each other except between conifers and mixed
woods. However, here as in all of the cover-type records, the use of
hardwood woodlands probably ranks somewhat lower than is
actually warranted. The fact that the hardwood stands were by a
wide margin the most extensive stands on the Connecticut Hill area,
coupled with the second fact that these several stands included the
largest continuous blocks, results in a lower use intensity than the
character of the type itself would warrant. As with all cover types,
the edges are used most, hence the types having the greatest pro-
portions of area far from edges will be used less, other things bemg
equal. If the prevalence of the hardwood-woodland types in an area
were small in comparison, let us say, to the coniferous or overgrown-
land types, then the intensity of use of the hardwoods in relation
to the others would rise markedly.

We have analvzed the shelter type selections of the adult grouse

86 The Ruffed Grouse

during the four seasons and we find them to vary widely according
to the season. These variations are exhibited graphically in Figure 4.
The most marked contrast is in the summer type use intensities in
comparison with the other three seasons. Now let us consider the
same data from the standpoint of the variation in use made through
the year of each major cover type group.

Coniferous Woods: It is rash to say that one type is more im-
portant than any other. The truth is that one type— any type— by
itself loses much of its importance. The potential value of any type
can only be attained when its quantity, quality, and distribution are
in harmony with the same qualities of other important types of
cover. It is likewise true that no type of cover is indispensable. Some
grouse may exist within the geographical range of the species even
if one type, however important, is completely absent. This is also
true, but progiessively less so, if two or more types are absent. Keep-
ing these generalizations in mind, it is clear that coniferous wood-
land is the most intensively used type during the colder months and
hence, with a reasonable balance of other types, the more important
of the several type groups during the period from mid-fall through
mid-spring. The marked preference for this type, especially in the
winter months, was enhanced in the records given here by the mod-
erate quantity and good distribution of this cover on the study area.
This factor tends to widen the use intensity ratio between conifer
cover and the mixed- and hardwood-woodland type groups because
of the greater area and more extensive blocks of the latter. Thus,
while coniferous woodland is an immensely important— almost vitally
important— component type for satisfactory grouse range, it should
be in moderate proportion and well distributed in order to be most
effective.

There were two main confferous types on the Connecticut Hill
area, hemlock woods and pine (white and red) woods. The hem-
lock is definitely superior to the pine type. Over the northeastern
range the coniferous-type group may conveniently be divided into
four types, hemlock, spruce-fir, white and red pine, hard pine (pitch,
banks, or Virginia). These are arranged in order of quality, with
the hard pine type definitely inferior to the other three. The qual-
ity of shelter in each is affected greatly by the age class and by the
amount of admixture of hardwood trees and shrubs. The highest
quality is obtained in the stands having all ages of trees, including

shelter

87

a moderate proportion of young reproduction. It is also desirable
to have a small proportion of hardwood species.

The variation in seasonal importance of coniferous types may be

Winter Spring Summer Fall

Fig. 5. Cover Types Used by Adult Grouse at DiflFerent Seasons.

noted from the percentages of total flushes per acre in these types
in each season: winter, thirty-seven per cent; fall, nineteen per cent;
spring, twenty-one per cent; summer, six per cent.

88 The Ruffed Grouse

Mixed Conifer-Hardwood Woods. The mixed woods is an impor-
tant year-round type providing a significant amount of conifer shelter
interspersed with hardwoods that furnish food and serve other pur-
poses. It is the one type group that, with interspersion of open land,
can support a fair grouse population without the presence of the
other major types. In winter it ranks second in use intensity only
to coniferous woods, and in fall and spring it ranks third behind
conifers and hardwoods. In the summer it is used less intensively
than any of the other crown- type groups except conifers.

The primary subdivisions of the mixed-woods type on Connecticut
Hill are distinguished by age class. Mature mixed woods has a high
proportion of the trees in the twelve inches and above d.b.h. class,
and a rather sparse understory. The submature mixed woods has a
large majority of trees in the four inches to 12 inches d.b.h. class
and the understory is usually well stocked. The latter is superior
to the mature stands for grouse cover. The mature stands can
usually be improved by selective cutting. When the density of ma-
ture trees is reduced so that reproduction of both conifers and hard-
woods establishes a good understory, the stand is changed to the
more desirable grouse cover type.

Over the northeastern states there are a number of subtypes of
mixed woodland varying considerably in value as grouse cover. The
subtypes are combinations of the two age classes just noted with the
several types of conifer and hardwood associations that occur as mix-
tures. The most important subtypes, in approximate order of value,
are as follows (each of these may occur as mature or submature
stands ) :

Hemlock— beech, birch, maple

White pine, red pine— beech, birch, maple

Spruce, fir— northern hardwoods

White pine— oak, hickory

Hard pine— oak, hickory

Hard pine— oak

The seasonal use of mixed woodland varies little except in sum-
mer, varying from nine per cent of all flushes per acre in summer to
twenty-three per cent in winter. The percentage of flushes for each
season are: winter twenty-three per cent; spring, twenty per cent;
fall, twenty per cent; summer, nine per cent.

Shelter 89

Hardwood Woods: The various types of hardwood stands are
ratlier mediocre grouse cover unless interspersed with one or both
of the conifer or mixed-woods types. Witli this interspersion, the
hardwoods are important as nesting cover in tlie spring, and as
feeding cover most of the year. It is not good roosting cover, except
as the birds sleep on nests and drumming logs in the spring. The
hardwood types are most important in spring and summer, least
important in tlie winter. The proportion of use for each season on
Connecticut Hill was: summer, thirty-six per cent; spring, twenty-
nine per cent; fall, twenty-two per cent; winter, seventeen per cent.

The hardwood group, apart from species composition, may con-
veniently be composed into three types, according to age class and
density: mature hardwoods, with a high proportion of the trees over
twelve inches d.b.h., and sparse understory; submature stands with
few trees over twelve inches, and a good understory. Either of the
former may have scattered large trees recently removed leaving
small openings in the crowai, usually with brambles filling in these
spots. The latter may be called "spot-lumbered." The mature stands
are definitely inferior to the other two as grouse cover and receive
little use. Theoretically, owing to the small scattered openings, the
spot-lumbered type should be superior to the subclimax hardwoods
from spring to fall. This proves true in the fall, but for some reason
fails in spring and summer. In the spring it proved to be less inten-
sively used than the regular type. The explanation for the failure of
the statistics to support logic may be inadequate data for the spot-
lumbered type.

The hardwood type most prevalent on Connecticut Hill was the
beech-birch-maple association. This seems to be the most satisfac-
tory of the hardwood types for giouse. In order of their value, the
other common hardwood types in northeastern grouse range are:
northern hardwoods, oak-hickory, and oak. All these types mav be
subdivided according to age class and density. These hardwood
types all exist extensively without appreciable conifer admixture,
and as such are poor grouse range.

Slashings: Areas that have been clearcut in recent years furnish
excellent cover for summer use. At all other seasons it is relatively
unimportant as a cover type. Slashings have a second use, however,
that may be even more important than its cover use, certainly so
when considered apart from summer. Within wooded areas slashings
serve to create openings, hence edges, and thus serve grouse range

90 The Ruffed Grouse

in the same manner as open land does. Extensive areas of con-
tinuous woodland range may be greatly improved by the judicious
creation of slashings in accordance with this principle.

As a cover type, slashings serve somewhat the same use as over-
grown land, especially in the summer. Both have an abundance of
summer food of berries, largely brambles, and insects. During this
season it is an intensively used cover type. In fall, winter, and spring
it is the least-used type excepting open land, largely due to poor food
conditions. The percentage of flushes per acre found in slashings in
each season were: summer, twenty-six per cent; fall, thirteen per
cent; spring, thirteen per cent; winter, eight per cent.

Slashings may be conveniently divided into two subtypes as
grouse cover, largely on the basis of length of time since cutting:
areas cut over from five to ten years or less, depending upon rate
of regeneration, that show considerable giound covered by herba-
ceous species, and that have many brambles and few saplings;
areas cut over five to ten years or longer, depending upon rate of
regeneration, that have little open herbaceous cover, and that have
many saplings both from sprouts and seedlings and relatively
fewer brambles than the first. A slashing grows from the first type
into the second. As cover, the type is most valuable during the lat-
ter half of its first stage and for the first few years of its second stage.
As an opening it is most valuable in its first stage. As the slashing
grows toward the pole stage, toward becoming either a hardwood
or mixed-woods type, its value as slashing cover gradually dis-
appears.

Overgrown Land: Shrubby areas grown up from old fields are an
important cover type group from spring through fall, but definitely
most important in fall. At this season it serves as a feeding ground
abounding in fruits and succulent greens. During the summer this
type produces an abundant supply of berries and insects and is
much used. In winter it is little used, for most of the fruit is gone
and winter shelter is usually lacking. The percentage per acre of
flushes in this type group by season was: fall, twenty-five per cent;
summer, twenty-two per cent; spring, sixteen per cent; winter, four-
teen per cent.

There are three primary subdivisions of this type group in the
Northeast, all occurring on Connecticut Hill in significant quanti-
ties. These are: pure stands of temporary hardwood species; mixed

shelter 91

hardwoods, mainly climax and subclimax species; mixed hardwoods
and conifers.

The first category were primarily popple {P. tremuloides) al-
though areas of hawthorn were common and stands of alder and
pin cherry also occuned. In all of these subtypes brambles were
plentiful in the ground cover. The use of these three categories
varied considerably both by type and seasons. Most intensively used
year round were the pure stands of short-lived species, next the
mixed hardwoods and lastly the mixed hardwoods and conifers. By
seasons, the popple and hawthorn stands were used most inten-
sively throughout the year, except in summer. During summer the
mixed hardwoods showed a little the heaviest use. The overgrown
lands with mixed conifers were used more intensively than mixed
hardwoods only in the spring; at all other seasons this subtype was
least used of the three. This relative unimportance of the coniferous
mixture in overgrown-land types as compared with woodland types
is interesting and reflects the use made of the several overgrown-
land types— feeding rather than shelter. The significant superiority
of the pure-temporaiy stands from fall through spring is probably
due to the value of the hawthorn fruit in the fall and of the popple
buds in winter and spring, but there may be a further explanation.

Open Land: Grouse are found in open-field cover most often in
fall, and least in winter. However, open land is not an important
cover type at any time for grouse. As already noted, its importance
lies in creating edge, and raising the quality of adjacent coverts
thereby, rather than in its own intrinsic value. A large proportion of
the records of grouse actually flushed in open fields are due to the
presence of a bushy fence-row, an isolated apple tree, or some sim-
ilar attraction.

THE EFFECT OF OPENINGS ON COVER SELECTION

The grouse is well-known as a bird of the edges. We have already
seen that in nesting, the grouse mother generally selects a location
near the woods edge or an interior opening. Likewise the broods
were found mainly in the open cover types or near the edges if in
the woods. The same principle follows in the selection of cover by
the old birds, although not so completely. I reported on the utiliza-
tion of coniferous reforestation by grouse (Edminster, 1935) and

92 The Ruffed Grouse

found a general limit of three hundred feet, beyond wliich the birds
did not penetrate the solid coniferous stands. In this case, few grouse
ventured more than two hundred feet from the outside edges. In
ordinary range we find this three-hundred-foot distance to be a good
rule of thumb for all woodland types, although most types are used
regularly in the two-hundred-to-three-hundred-foot zone. Even the
brushy cover types are relatively little used at a distance of more
than three hundred feet from the edges when present in large uni-
form blocks.

We conclude that cover types should be so interspersed that no
point is over three hundred feet from an edge (between open or
brushy type and woodland type) if the range is to be fully and
regularly used. The optimum zone of woods types is from the edge
to a depth of two hundred feet.

VARIATION IN COVER TYPE PREFERENCE AT
DIFFERENT TIMES OF DAY

There are numerous factors that combine to determine the type of
cover selected by a grouse at any given time. We have just observed
the marked variation in the different seasons. Less marked, but still
highly significant, are the differences in cover needed at various
hours of the day within the seasons. Seasonal requirements must be
met within the limits of the bird's yearly range— a reasonably flex-
ible limitation— but the hourly changes required must be available
within the daily cruising radius.

During the fall and winter the most significant trends are: a high
early-morning-use intensity of coniferous woods, hardwoods, slash-
ings, and overgrown land, all generally declining in use intensity
through the day; a low intensity in the early morning for mixed
woods, which rises through the day. All these trends are logical,
except that it would seem that the trend for use of coniferous woods
should parallel that for mixed woods instead of the hardwood
group. The fall and winter season use of the several deciduous
types at early morning feeding time, most notable in the fall, are
linked with the birds' feeding habits.

In spring the hourly cover-type-use trends shift somewhat from
those of winter. The use of mixed woods rises from early morning
until midafternoon as before, but then significantly drops again

Shelter 93

toward evening. The variations in use of other types are less notable.
A rise in use of conifers toward evening is contrary to the previous
season's trend.

Summer adjustments complete the shifts in daily cover type use.
Slashings show a notable increase in use intensity from morning to
night. Mixed woods has a high use intensity at midday with
markedly lower use in the early day and again towards evening.
This is complementary to the trend in use of overgrown land, which
is high in morning and evening and low at midday. In summer, the
use of conifer woods is not important.

The reasons for these movements from one cover type to another
during the day are linked with the birds' habits, although often
conditioned by the weather. Thus the habit of feeding in early
morning takes the bird from its night roost, usually conifers or mixed
woods except in summer, into the types that furnish food, usually
the deciduous types. However, if the weather is inclement, this nor-
mal habit will likely be altered temporarily; the bird either gets
some food in its protection cover or delays feeding for some hours,
or even days, until tlie weather is better. The requirements for sun-
ning, dusting, drumming, and nesting similarly stimulate quick
movements from one cover type to another for the purpose of the
moment. To provide adequately for these vai-^ing needs within easy
daily range, the range must provide not only the needed types of
cover, but also have them interspersed intimately enough for ready
accessibility. Upon these needs must be built any effective plan for
range management.

THE EFFECT OF SLOPE ON SELECTION OF
COVER BY ADULT GROUSE

Slope bears numerous relations to cover and to the acceptability
of cover under various conditions. The steepness of a slope may
impede the bird's mobility, or aid it. When on level ground, a grouse
moves about on the basis of its horizontal flight capability. With
increasing steepness of slope, it requires more power to go uphill
and maintain speed and less power to go downhill; that is, it loses
speed when ascending, and gets additional velocity when "power
diving." Whether a bird desires to go uphill or down when on a
given slope may depend considerably on the location of cover types.

94 The Ruffed Grouse

Considering these factors, it is at least possible that grouse may
choose to avoid areas because of their degree of slope.

Similarly, the direction of the slope may be important. A north
slope, that is one sloping downward predominantly between north-
west and northeast, is a cool slope, whereas the south and west
slopes are comparatively warmer. Cover composition often varies
considerably according to aspect, and may affect grouse distribu-
tion. For example, in south-central New York, mountain laurel is
found only on south slopes. When winds come predominantly from
certain directions, particularly winds accompanied by storms, oppo-
site slopes are tlie most protected, other things being equal.

Substantially the same relationships among the slope-use inten-
sities prevail at all seasons. Steep slopes are used much less than
others. This partial avoidance of steep slopes— those in excess of
twenty degrees— is probably due to a disinclination to climb up and
down, even in flight. Just how steep a slope has to be before a
grouse begins to shun it is not at all clear.

The other significant preference is for east and west slopes over
flat (less than five degrees), south, and north slopes.

COVER TYPES CHOSEN AS AFFECTED BY
WEATHER CONDITIONS

Every hunter of grouse knows that the birds will be found in dff-
ferent situations under varying weather conditions. It is natural
that there should be differences in selection of cover with changes
of temperature, sky conditions, wind, and possibly other elements.
We ourselves seek the most protective abode in a storm, and are
inclined to sun ourselves under clear, balmy skies. So why should
not the grouse do likewise? And they do.

Effect of Temperature on Cover Used. Temperature is relative as
it affects grouse behavior. Fifty degrees Fahrenheit in the summer
would be cold, in the winter warm, and in spring or summer it
would likely be about normal. Conceivably, it would be pos-
sible to record the temperature and precise measurements of other
weather factors in the field as each grouse is flushed, but this is un-
necessary. The temperature at the moment involved may be gauged
as warm, normal, or cold, where normal means that it is within the

shelter 95

regularly anticipated range of temperature for the time of year,
and the others indicate an abnormal extreme in one direction or
the other. Analyzed on this basis we are able to discuss numerous
ejffects of temperature on selection of cover by adult grouse, but it
should be made clear here that temperature is less important as
the determining element of the weather than either sky condition
or wind.

During the fall and winter, especially in the winter, there are more
shifts from the normal in cover type use as a result of temperature
extremes than at any other season. Most significant are a high-use
rate of overgrown land in warm weather and an avoidance of slash-
ings, and an increased use of hardwood woodland and coniferous
woods in cold weather. The increase in use of hardwood stands on
cold days is mainly in the autumn, that of conifers in the winter.
Also important are the complementary effects— an avoidance of
brushland in cold weather, and of hardwoods and conifer woods
in warm days.

In the spring, the tendency to deviate from average habits under
extremes of temperatures is less pronounced than in the winter. Par-
ticularly significant at this season, however, is the marked avoidance
of hardwood stands on cold days, and of coniferous woods when it
is warm. Peculiarly enough, mixed woodlands are selected more
than normal under both extremes, hot days and cold. Hardwood
stands are used intensively in warm weather too, while slashings
seem to attract most in cold weather, and less than normal in warm
weather.

During the summer, deviations from average are less than at other
seasons. Most important is a clear trend to the use of coniferous
cover on hot days (the opposite trend is true in winter), and a
corresponding avoidance of hardwood stands. Of some significance
is the greater-than-average use of slashings in warm weather and of
mixed woods when it is cold.

Thus, the other things being equal, the selection of cover type
is considerably affected by temperatures. But it may be repeated
that temperature is less important as a determining element than
wind and sky conditions.

Relation of Sky Condition to Cover Used. The condition of the sky
plays a very important part in determining what cover a grouse will

96 The Ruffed Grouse

utilize. While there are endless variations that might be analyzed in
relation to grouse habits, the major sky conditions are predominantly
sunny, cloudy, rainy, and snowing.

Except for the spring, there are many marked variations in cover-
type use according to atmospheric condition. In spring, the prob-
lems of courtship, mating, and incubation probably have a leaven-
ing effect on quick shifts in cover types used.

During fall and winter, the most notable deviation from average
on sunny days is a marked increase in use of slashings and a de-
crease in use of confferous cover. There is no significant shift from
normal during cloudy weather at these seasons, but precipitation
brings the most marked changes of all. In the autumn, rain brings
a strong shift to overgrown land and slashings while the heavier
woodland types are ^1 used less, though the decreases are dis-
tributed evenly. In winter, simny skies bring a very different shfft,
an immense increase in use of coniferous cover, and reduction below
average in all other types. Thus there are many extremes in cover
selection at these seasons according to the sky, more than at any
other season. Generally the trend is to the more protected areas in
stormy weather and to the more exposed (and food-producing)
coverts in mild weather.

Spring is a season of more stable cover type use habits. The one
really significant deviation from the norm is the gi-eat shift to conif-
erous cover during snowstorms, and a corresponding reduction in
all other types. The only appreciable deviation in sunny weather is
a shift in use of heavy cover from conifers to mixed woods. Under
cloudy skies there is an increase in use of overgrown land and
conffers and a decrease in use of mixed woods. In the rain there is
a general reduction in use of all types from overgrown land to
mixed woods in favor of coniferous cover.

During the summer the greatest deviations occur during rain-
storms, and these are very pronounced; a great increase in use of
mixed woodland (the heaviest protection of confferous woods is
apparently unnecessary at this season ) , and a large decrease in use
of overgrown land and hardwood woods. There is also a notable
shift to mixed woods and overgrown land in cloudy weather and a
reduction in use of slashing cover, and evergreen woods. On sunny
days the deviations are not so pronounced.

At all times of the year there is a generally increased rise in the

Shelter 97

use of heavy shelter types in stormy weather and a greater use of
the more open type^ in clear weather.

Cover Type Use as Affected by Wind. The strength of wind, with-
out respect to direction, may be noted as either still, moderate, or
strong. When the air is entirely quiet or with just an appreciable
movement, it is called still; when gusty or an average light blow, it
receives the designation of moderate; when the wind rises to stormy
strength, probably exceeding twenty miles per hour, it is strong.
The evidence shows clearly that a strong wind, as compared to a
lack of vraid, produces very significant differences in predominance
of cover types selected. While it is not possible fully to separate the
relative effects of the different elements of the weather, all acting
at once as they do, it is probable that wind is nearly as important as
sky condition in determining grouse use of cover types. Surely it is
a more prominent factor than temperature at its extremes.

More than that of any other factor, the effect of wind is con-
sistent throughout the year. Strong winds bring an increased use
of overgrown land (particularly popple stands) at all seasons and
of coniferous woods from fall through spring, and lessened utiliza-
tion of mixed woodland at all seasons and other types at some
seasons. In still weather, especially in summer, the most notable
shift is to hardwoods.

In analyzing the effect of weather on cover type selection, we
have separated the three most effective weather elements and con-
sidered them separately. This simplified the analysis but actually, of
course, the weather factors all occur together, and hence we should
consider what cover is used when the several factors occur in the
various combinations. To do this would require long and relativelv
unproductive analysis. We have noted that sky condition appears
to be the more dominating element, wind following closely, and
temperature trailing. However, it is also true that extremes, whether
of one factor or another, tend to be the determining element. Thus,
on a clear, quiet, and hot summer day, the temperature may de-
termine the cover a bird will select; or on a very windy day, with
cloudy skies and normal temperatures, the wind will be the guid-
ing element. When extremes of similar direction occur together, as
a snowy, windy, cold day, it makes little difference which elements
dominate the consideration since all bring the same result.

98 The Ruffed Grouse

RELATION OF COVER TO GROUSE MORTALITY BY PREDATORS

It is generally recognized diat cover conditions affect predation.
Other things being equal, quality of shelter and loss from predation
are inversely proportioned— the better the cover, the lower the mor-
tality from this cause. It would logically follow then that different
cover types, since they vary in their shelter quality, would produce
different predation rates. Possibly otlier aspects of cover too may
affect predation mortality. Let us consider this problem from the
point of view of the locations where grouse are found killed.

In the first place we must make a hazardous assumption: That
the locations where grouse remains are found are the places where
they were killed; or at least the occasions of difference will balance.
However, that assumption is invalid. Since a large number of grouse
kills are picked up in open fields, it is quite certain that the majority
were carried here by the predators from the location of the kill in
some other cover type. And from the habits of many of the predatory
species, especially in the spring of the year, we know that it is a
common habit to carry a kill some distance before consuming it,
and there leaving the waste parts.

While we cannot depend too much on the locations where grouse
remains are found as an indication of cover type vulnerability, there
are some fairly dependable conclusions that can be drawn when
supported by field observations. Probably the most important cor-
relations are the relatively low vulnerability of the birds in conff-
erous woodland and overgrown land, in both cases a reflection of
high protective character of the cover types.

Shelter as a Limiting Factor. We have seen that the ruffed grouse
has a rather complicated shelter requirement to satisfy its needs at
all seasons, under various conditions of climate and other physical
conditions and for all of its activities. Yet at the same time we are
impressed repeatedly with the remarkable adaptability of the bird
to different conditions and to the changes that bring about new
environments. With all its flexibility in meeting successfully a variety
of conditions, it is still limited very definitely by the shelter factor
of the environment. Shelter is probably the limiting factor in the
long run throughout most of the Northeast.

shelter 99

We may better understand the importance of shelter if we con-
sider the two extremes: no appreciable shelter available; and a cover
composed of too-complete shelter. In the first instance we may con-
sider a large area of open farmland devoid of woodland or brush.
It may be of any type of fields— hay, pasture, or farm crops— they
are all alike to the grouse in that they are almost completely devoid
of shelter of the kind required by these birds. Of course, this type
of range produces no grouse, can maintain no grouse, even though
many grouse foods ( herbaceous ones ) might be present in quantity,
and other needs may be present. Without shelter, grouse cannot in-
habit an area. On the other extreme we have the situation where
shelter cover is so dense and extensive that little of it can be utilized
by grouse. Extensive areas of pure stands of some conifers at cer-
tain age periods are in this condition. The birds are able to use only
the exterior margins and very rarely will penetrate more than three
hundred feet into such an area ( Edminster, 1935 ) . Here again shel-
ter has proved to be a limiting factor for grouse, limiting by virtue
of excluding other needed components of the range.

Between these two extremes we have all manner of variations in
shelter conditions and in the degree of limiting effect it has on
grouse populations. If shelter is excellent on a piece of grouse range,
food conditions may well be good also, in which case some other
factor would limit the numbers of grouse, possibly the saturation
point of four acres per bird. Generally this ideal state will not pre-
vail and the quality, quantity, and arrangement of the shelter com-
ponents of the range will play a considerable part in determining
the level of grouse abundance.

REFERENCES AND CITATION SOURCES ON RUFFED
GROUSE SHELTER REQUIREMENTS

Bump, Gardiner. Analysis of Certain Cover Requirements of the Ruffed

Grouse in New York State, Trans. Third N. A. WildUfe Conf., 1938.
Edminster, F. C. The Effect of Reforestation on Game, Trans. 21st Am. Game

Conference, 1935.
Fisher, L. W. Studies of the Eastern Ruffed Grouse in Michigan, Mich. State

College, Tech. Bui. 166, 1939.
Leopold, Aldo. Game Management, 1933.
Palderboer, Emmett B. Cover Requirements of the Eastern Ruffed Grouse

in Northeast Iowa, Iowa Bird Life, XII, pp. 50-55, 1942.

100

The Riiffed Grouse

Studholme, A. H. RufiPed Groiise Environment in the Scrub Oak-Pitch Pine
Forest Type, M.S. thesis, Pennsylvania State College, June, 1941.

. Game Cover Types in New York State, N. Y, S. Cons. Dept., Bureau

of Game Circular #4 (mimeo.), March 23, 1936.

New York State Conservation Department Annual Reports for 1930

and 1933.

— . Forest Cover Types of the United States, Joum. Forestry, April, 1932.
(By S. A. F. Committee on Forest Types.)

Food and Water

Types of Food Utilized. Within tlie limits of its physical ability
the rufiFed grouse may be said to be practically omnivorous in its
food habits. Vegetable foods of nearly all types are taken: leaves,
buds, twigs, fruits, seeds, flowers. Its animal foods are more restricted
in type due to the size and structure of the bird; insects and other
related invertebrates are about the only forms it can readily catch
and handle. Within these limitations almost any kind of insect,
spider, or similar type is accepted.

Winter

Spring

Summer

SEASON

Fig. 6. Seasonal Utilization of Food by Adxilt Ru£Fed Grouse According to Type
of Food (Northeastern States)

For purposes of simplification we may include all animal foods
as one type, twigs and buds as another, flowers and leaves as tlie
third, with seeds and fruits as the fourth. Thus arranged, the sea-
sonal food habits of the adult grouse according to type of food are
given graphically in Fig. 6.

101

102

The Ruffed Grouse

This information apphes to the northeastern states, from Viiginia
and West Virginia to Maine, and is of necessity somewhat ideahzed
since records for all seasons for all parts of the region are not avail-
able in adequate quantity. Thus, the percentages at any exact time
of the year for any specific area are not set forth as exactly correct.
However, the diagram does indicate the trends in seasonal use of
v^arious types of food.

It is clear that animal food furnishes a significant proportion of
the volume of adult food only during the summer. Leaves, includ-

—1

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FRUITS AND SEEDS

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MONTHS

August

September

Fig. 7. Utilization of Food by Young Ruffed Grouse According to Types of
Food (Northeastern States)

ing fern fronds, flower parts, and other vegetative parts except buds
and twigs, are by a narrow margin the most staple type of food used
throughout the year, increasing considerably in the spring and sum-
mer, but furnishing from fifteen per cent to forty-five per cent of the
food at all times. Fruits and seeds furnish nearly as much, generally
from fifteen per cent to fifty per cent, of the diet, with greatest use
in summer and fall, while buds and twigs vary widely, from an
almost negligible quantity in summer to nearly three quarters of all
food in late winter.

The types of food utilized by the immature grouse are widely at
variance with the adult diet during the early summer but gradually
change until by the end of the juvenile moult they agree with those
taken by ad\ilts. These trends are illustrated in Fig. 7.

Food and Water 103

It will be noted that the chicks start off with a full insect diet, but
soon add fruit to their fare. By the time they are a month old they
are consuming green food and by midsummer insects make up less
than half the diet. Toward the end of the summer, buds are added
to the menu and the use of insects decreases rapidly. As with the
case of the diagram of adult food types utilized by seasons, the
trends in food habits of the young birds had to be interpolated freely
in order to get a smooth trend. Except for occasional quick changes
in food habits resulting from violent weather, or the sudden ripen-
ing of a certain fruit, trends through the months take place quite
gradually and therefore properly appear in a graph as smooth
curves.

Utilization of Food Species by Seasons. Compared with that of
most wild creatures, the ruffed grouse's diet in the Northeast is well-
known. The records given here and the conclusions drawn are based
on around 4,000 stomach analyses of birds collected from Virginia
and Ohio to Maine as well as upon many supplementary observa-
tions. However, because the majority of the specimens are obtained
as a by-product of hunting, the records are largely from fall and early
winter, and the late winter, spring, and summer material is relatively
scant. The New York investigation made collections of adult birds
throughout the year, except during the hunting season, in order to
obtain the needed food habits information. Young birds as well as
adults were collected through the summer. With a scattering of rec-
ords from other sources, the year round food habits of the species are
now pretty well established.

The diversity of foods taken by the grouse is indicated by the
some 374 kinds of plants and 131 kinds of small animal life already
known to be eaten by it in measurable quantities in the Northeast.
And yet this list is without doubt far from complete. The author
picked up a grouse (killed by a broken neck) in Sullivan County,
Pennsylvania, in December, 1940 while deer hunting. Its crop con-
tained several fruits of mountain holly {Ilex montana)—a species not
previously recorded as eaten by grouse. But this normally rare plant
happened to be common on this particular mountain, and the adapt-
able grouse appeared to be making good use of it.

The number of species of insects known to be taken by grouse
is doubtless only a small fraction of those that the birds actually eat.
Experiments conducted by the State Conservation Department at

104

The Rufted Grouse

Ithaca ill 1936 with captive grouse chicks indicate they wiU take
almost any kind of insect. The main limitation is their abihty to
locate the prey and then to capture it.

Even though grouse eat parts of plants of hundreds of species and
in many cases two or more parts in different seasons, the bulk of their
food at any one season comes from a relatively small number of
species. The species that are predominantly represented in the food
at any given time and place depend upon availability and the degree
of preference by the grouse. There is wide variation between Vir-
ginia grouse food habits and those in New York, or between those
in New York and those in Maine. But at the same time, insofar as
staple food plants are available in each part of the range, they are
consistently used.

The sources of food habits records given are mainly as follows:

Author

Year of Publication

No. of Analyses

Judd, Sylvester D.^

1905

208 adults, 18 chicks

Smyth, Thomas

1923 (unpublislied
thesis )

279 adults, 8 chicks

U. S. Biological Survey ^

1933

111 adults

Kelso, Leon ^

193,5

80 adults

New York State Conser\'ation De-

partment's Annual Report -

1937

—

Gross, A. O.

1937

1055 adults

Hosley, Niel ^

1938 (unpublished
thesis )

925 adults

Nelson, A. L., Clarke, T. E., &

Bailey, W. W.^

1938

185 adults

Van Dersal, W. R.

1938

—

Darrow, Robert W.^

1939

485 adults, 332 chicks

Kuhn, Tracy M.

1940

230 adults

Allen, A. A.

not published

321 adults

Kuhn, Tracy M.

1941

207 adults

MacGregor, Arthur E.

1941

102 adults (some chicks
included)

Merritts, Helen V.

1943

112 adults

GilfiUan, M. C., & Bezdek, li.

1944

42 adults

Total

4300 1 adults, 358 chicks

^The records used in these studies overlap to a considerable extent; hence the
totals of specimens used are not wholly original vAth each study after the first, and
the grand total of analyses is larger than tlie actual nmnber of birds involved.

^ Darrow's records include all the specimens covered by the 1937 N. Y. S. Cons.
Dept. Ann. Rep.; hence the number of specimens is omitted from the latter reference.

By seasons, the adult records were approximately: Fall— 2,113; vvdnter— 392;
spring— 125; summer— 85; fall and vdnter— 1,318.

By area they were collected: Nortlieast (in general)— 319; New England— 1,818;
New York-995; Pennsylvania— 642; Virguiia-205; Ohio— 42.

The chicks were largely from New York.

Food and Water 105

WINTER FOODS

Insects are a negligible item of food during the winter months.
The greatest proportion of winter food consists of buds and twigs,
with fruits and mast taken as available, and a surprising amount of
green leaves taken, considering the general difficulty in finding ex-
posed ground vegetation. In the region from mid-Pennsylvania,
northward, the buds of cherry, birch, apple, and popple are abun-
dantly used, with those of hophombeam vying with them in impor-
tance in New York. Fruits of dogwoods, apple, and grape are very
important foods throughout the Northeast, while those of greenbriar,
acorns, and laurel leaves are among the most important items of diet
from Pennsylvania southward. Others indicated as of considerable
importance by the leading food studies are the leaves of ferns, Can-
ada mayflower, strawberry, sheep sorrel, and partridgeberry; fruits
or seeds of sumac, rose, partridgeberry, hawthorn, and winter-
green; and buds of maple, blueberry, and blue beech. Beechnuts are
used intensively when available but have not amounted to a primary
item in any recent analysis (see under relation of food to cover
types ) .

Taking the Northeast as a whole, the records show that from
forty-seven to sixty-four per cent of the winter food normally comes
from plants of twelve genera that average two per cent or more each
in total volume of food taken.

The remainder is derived from plants of a large number of genera.
In each individual study from ten to all twelve of these genera were
recorded in significant quantities, and from six to nine were found to
compose two per cent or more of the total food. These twelve genera
and their contribution to the winter food are summarized in Table 3.

SPRING FOODS

Actual analyses of spring-collected grouse are confined to those
made by the New York investigation and a scattering from the rec-
ords of Smyth, Judd, and others. It is, therefore, necessary to inter-
pret somewhat our knowledge of foods palatable and food available
in order to present a broad summary of the birds' diet at this time
of year for the whole Northeast. There is a gradual transition from
staple bud and twig foods available all wdnter, regardless of snow.

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Food and Water 107

to a larger volume of succulent green-leaf food exposed by melting
snow and newly sprouted with the advent of the new growing season.
As spring merges into summer, buds practically disappear from the
diet, fresh fruits become more important, and insects are taken to a
significant extent. Flowers, particularly strawberry blossoms, are
sometimes eaten in considerable quantity.

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and blueberry continue to furnish a large per cent of the food well
into May. Leaves of ferns, grass-like plants (particularly Carex),
strawberr\% hawthorn, partridgeberry, sheep sorrel, wood sorrel,
mountain laurel, Canada mayflower, and wintergreen are all of con-
siderable importance. Owing to scarcity, relatively little fruit is eaten
until the advent of summer brings on the strawberries. The fruits
most commonly eaten during the spring are those of sumac (includ-
ing poison ivy), rose, greenbriar, hawthorn, wintergreen, and par-
tridgeberry.

SUMMER FOODS

The summer diet is composed mainly of fruits in season as they
ripen, green leaves, and insects. For the adults the emphasis is on
fruits, with green material a close second, and insects a poor third.
For the chicks insects are predominant during June and part of
July, with fruit second and leaves third. During the latter part of
the summer the young gradually take the same proportions and types
of foods as do the adults.

Outstanding among the fruits eaten by adults are strawberries,
raspberries (including blackberries, dewberries, etc. of genus
Ruhus), cherries, sedges (seeds), and blueberries (see Plate 21).
Most prominent of the plants providing green materials are poplar,
touch-me-not, buttercup, apple, hawthorn, and partridgeberry.
Fruits of Juneberry (Amelanchier) and some dogwoods are also
eaten to a considerable extent in the summer, the former during the
early part and the latter during August and September. Insects con-
stitute from one per cent to three per cent of the total food. Orders
of insects and related animals most commonly utilized by adult
grouse in order are: Hymenoptera (mainly ants), Coleoptera (bee-
tles), Arachjiida (spiders), Lepidoptera (caterpillars), Orthoptera
( largely grasshoppers ) , and Opiliones ( harvestmen ) .

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cc « a;

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s

to

-^

110 The Ruffed Grouse

For the chicks, insects, spiders, etc. made up forty-six and eight-
tenths per cent of the food in the Adirondacks, sixty-eight and three-
tenths per cent in the Catskill part of New York and about sixty per
cent over the rest of the state. In July the proportion of animal food
in the diet has dropped to between ten and twenty per cent and in
August it has further declined to from one and three-tenths per cent
in the state outside the mountains to six and nine-tenths per cent in
the Adirondacks. In records on eight stomachs, Smyth found sixty-
six and nine-tenths per cent animal food in June, ele\'en per cent in
July, ten per cent (one stomach) in August. Thus, the young birds
change their food habits rapidly; from an almost one hundred per
cent insect diet in the first week or two, they shift to only a small
per cent of insect foods within tlu"ee months. Other foods are siiuilar
to those of the adults.

Fruits eaten by the young birds are mainly the same as with the
adults: raspberries, strawberries, Juneberries, and cherries. How-
ever, raspberries assume vastly more importance than with the
adults, comprising over thirty per cent of the total summer food.
Elderberries also are considerably used.

The young birds partake of a larger quantity of seeds in summer
than do their parents. Seeds of maple, violet, jewelweed, and sedge
are taken in considerable quantity, while leaves of sedge, buttercup,
and sheep sorrel are staple foods.

Orders of insects used by the chicks in order of importance
are: Hymenoptera, Coleoptera, Lepidoptera, Opiliones, Hemiptera
(bugs), and Diptera (flies). Thus the insects and other invertebrates
taken by the chicks come mainly from the same orders as with the
adults and nearly in the same order of importance. With the chicks,
the true bugs {Hemiptera) and flies (Diptera) were significant but
not for the adults; for the adults grasshoppers (Orthoptera) and
spiders {Arachnida) were taken in numbers but not by the chicks.

FALL FOODS

As summer gives way to autumn the quantities of fruits and green
leaves in the diet gradually decrease, insects nearly disappear, and
buds are taken in constantly increasing abundance. It is not until the
arrival of winter, however, that buds become the dominant type of
food. As long as fruits and tender leaves are available, they are pre-

Food and Water 111

f erred. Outstanding among the fall foods taken are acoms and haws.
Finaits and buds of cherries, leaves of sheep sorrel, and wild grapes
come next in general importance. From fifty-six to sixty-nine per cent
of the total fall food normally is derived from plants of fifteen
genera that each average two per cent or more of total fall food.
These are summarized in Table 4.

While these fifteen genera of plants are listed in order of volume
of food furnished by each, obtained by arithmetically averaging the
records from the various studies of fall food habits, it is not safe to
attach very much importance to the exact order. The relative vol-
umes of food gleaned from plants of the various groups depends on
availability as well as preference, and even in a given locality the
availability of some species varies widely from year to year. Beech-
nuts are an excellent example— a primary and preferred fall food in
a good crop year, but relatively unavailable during most years. It
is also true that the averaging of the statistics from different studies
is not a proper statistical procedure. However, if the order is not
taken strictly, this procedure does seem to bring out the group of
primary foods.

In certain areas, a food plant may be of great importance, and
yet not be one of the fifteen groups listed above. For example, in the
Adirondacks of northern New York wild raisin (Viburnum) , wood
ferns {Thelypteris=Dryopteris) , violets (Viola), and Juneberries
(Amelanchier) are among the ten most used fall foods— but all of
these are not among the fifteen most important for the Northeast as
a whole and do not average two per cent of the fall food over the
Northeast.

SUMMARY OF FOODS UTILIZED

Arbitrarily taking the twenty-five genera^ of plants that furnish
the most food to grouse in the Northeast year in and year out, we
find that their contribution to the total food of the species is from
sixty-five to eighty-eight per cent, varying with locality, year, and
season. These we may consider to be the primary foods of the ruffed
grouse in this region. No attempt is made at this point to say which of
these twenty-five is first, second, third, etc.; there is too much

■"^ One of the groups named is a family (Polypodiaciae), that of the ferns— an as-
semblage of genera, of which those most used are Thelypteris and Pohjstichum.

Fig. 8. TV/ENTY-FIVE MOST IMPORTANT FOOD PLANTS USED BY
THE RUFFED GROUSE IN THE NORTHEAST

FROM 65% TO 88% OF THE FOOD OF THE ADULT GROUSE AT ANY TIME OF THE
YEAR WILL USUALLY COME FROM THIS GROUP

PLANT GROUPS

MONTHS UTILIZED

Jan. I Feb. | Mar. | Apr. | May | June | July | Aug. | Sept | Oct | Nov. | Dec.

BIRCHES

(Betula)

SEDGE .r

(Carex) ^

DOGWOOD ■

(Cornus) ■

HAZELNUT

(Cory! us)

HAWTHORN r

(Crataegus) L

BEECH ■

(Fagus) ■

STRAWBERRY ■

(Fragaria) *

WINTERGREEN

(Gaultheria)

MOUNTAIN LAUREL..

(Kalmia)

CANADA MAYFLOWER

(Maianthemum)

APPLE

(Malus)

PARTRIDGEBERRY

(Mitchelia)

HOPHORNBEAM

(Ostrya)

FERNS

(Polypodiaceae)

POPLARS

(Populus)

CHERRIES

(Prunus)

OAKS

(Quercus)

SUMACS

(Rhus)

ROSES

(Rosa)

BRAMBLES _..

(Rubus)

SHEEP SORREL

(Rumex)

GREENBRIERS.

(Smiiax)

BLUEBERRIES_

(Vaccinium)

VIBURNUMS

(Viburnum)

GRAPE

(Vitis)

Period of use as a primary food
Period of use as a secondary food

112

Food and Water 113

variation in different areas to make such a ranking of much value.
However, it is apparent that some are of greater utility than others.
The question of palatability is discussed later. Suffice it here to say
that if adequate quantities of these twenty-five groups are readily
available, the food situation on a grouse range is not likely to be a
limiting factor. Figure 8 shows the months during which these foods
are of primary importance (solid bar) and of secondary importance
( dashed bar ) .

THE TWENTY-FIVE PRIMARY GROUSE FOODS IN
THE NORTHEAST

The BmcHES (Betula)

Species utilized: yellow birch {B. lutea), black or sweet birch
(B. lenta), paper birch {B. papyrifera) , giay birch (B. populifolia) ,
river birch {B. nigra), listed in approximate order of importance
(see Plate 20C).

Seasonal importance: most important as a winter and early-spring
food, reaching the height of utilization in February and March;
taken in increasing quantities from late September on to the mid-
winter peak, and in rapidly decreasing quantities after midspring.

Parts used: Almost entirely leaf buds; slight use of catkins and
seeds.

Geographical importance: Northern New England, New York and
Pennsylvania constitute the region of greatest use of birch. The
range south of Pennsylvania and in southern New England pro-
vides little of the three more valuable birch species and reflects
this deficiency in low utilization records. The gray birch, most preva-
lent in southern New England, is relatively unimportant as a source
of grouse food.

The birches, particularly yellow, black, and paper, are one of the
groups of trees and shrubs which through their readily available and
palatable buds enable the grouse to survive the rigors of winter in
the northern portions of its range without difficulty. Utilization
within the genus is partly determined by availability, partly by selec-
tivity. Yellow and black birches, and to a slightly lesser degree paper
birch, are highly palatable. Their use by grouse is mainly determined
by their abundance and distribution. Gray birch, and probably river

114 The Ruffed Grouse

birch, are less palatable; the latter is not well distributed in good
grouse range, and both are relatively of little importance.

The yellow, black, and paper birches are long-lived species. All
three are found abundantly in the climax or subclimax types of wood-
land and forest, mainly in the Appalachian and St. Lawrence plant-
growth regions of the Northeast.^

Paper birch finds its optimum conditions in the more northern
St. Lawrence zone in association with the spruce-fir type, while
black birch is more prevalent in the Appalachian area, particularly
its northern portion. Yellow birch is an important species in the
northern hardwoods type of the St. Lawrence region, and in the
beech-birch-maple-hemlock type of the Appalachian area. These
tliree birches practically never appear in pure stands, although paper
birch sometimes constitutes a high percentage of the crown species
mixture. They are all found in mixtures of both hardwoods and con-
ifers. Gray birch is a temporary, short-lived species that pioneers
on old field areas.

Paper birch seeds in abundantly following burns and is moisture
loving but does not thrive in swamps. The yellow and black birches
seed in readily in the understory of rich, well-drained, upland wood-
lands but do not tolerate swampy soils so well.

The birches are also used as food by the white-tailed deer, cotton-
tail rabbit, beaver, moose, and snowshoe hare. Since these mammals
all feed from the ground, the grouse receives no significant competi-
tion from them for the tree buds.

Chemical Composition: ^ Yellow birch— fresh buds: water— 45.0
per cent; protein— 5.2 per cent; fat— 2.4 per cent; nitrogen-free
extract— 25.5 per cent; fiber— 20.4 per cent; ash— 1.5 per cent (Hos-
ley, 1938).

Sedges (Carex spp.)

Species utilized: Many, including the following seven that have
been specifically identified: C. crinita; C. flextiosa; C. gracillima;
C. intumescens; C. lupulina; C. plantaginia; C. tenella. There is in-

^ Van Dersal, 1938, gives a map of the plant growth regions.

^ The significance of these analyses, given when available, presumably lies in
showing nutritive values. Since little is known of the specific nutrition of the grouse,
no attempt is made here to interpret the figures.

Food and Water 115

sujBBcieiit information concerning the use of the different species to
warrant any comparisons of degree of use. Parts of these plants are
usually identified only to genus.

Seasonal importance: Most important as a summer and fall food,
reaching greatest use in summer by both adults and chicks. Utiliza-
tion begins with the growth of the new leaves in the spring and
continues until seeds are gone and leaves dried up in the fall.

Parts used: Seeds and leaves; leaves from spring through fall,
seeds in summer and fall.

Geographical importance: Since the seeds and leaves of sedges
are only important from spring to early fall, it is not surprising to
find only small quantities recorded in the majority of food studies—
those made of fall and winter specimens. However, records are
available from Canada to Virginia, even in late fall stomachs. It
seems likely that Carex is an important spring and summer food
throughout the region as it is in New York, especially for the young
birds.

Of more than two hundred recognized species of Carex, repre-
sentations of only seven have been specifically identified, five in
New England by Gross, one in New York by Smyth, and one by
Judd. However, it is likely that many species are taken by grouse
and that use is primarily dependent on size of seed, distribution,
and abundance. Sedges are low-growing, perennial, grass-like herba-
ceous plants ( see Plate 20B ) found mainly in old fields, along woods
edges, in marshes and clearings. Some species are found in open
woodlands, still others in thickets. Many are particularly adapted
to wet soils although some will grow in very dry situations.

The Dogwoods {Cornus spp.)

Species utilized: Flowering dogwood (C. forida); gray dogwood
(C. paniculata) ; bunchberry (C. canadensis); red osier (C. stoloni-
fera); roundleaf dogwood (C. circinnata); silky cornel (C. amo-
mum); roughleaf cornel (C. asperifolia) ; blue cornel (C. alterni-
folia); in approximate order of importance.

Seasonal importance: Fruits of some species of dogwoods begin
to ripen in midsummer and are taken by both adult and young
grouse. Most species are primarily fall food producers but flowering
dogwood fruits are also important as a winter food.

116 The Ruffed Grouse

Parts used: Mainly the fruits altliough buds are taken to a sUght
degree.

Geographical importance: Dogwoods are an important fall grouse
food throughout the region. C. canadensis is primarily important in
the northern portions of New England and New York, C. paniculata
from Pennsylvania northward. C. florida is important both in fall
and winter from southern New England to Pennsylvania and Ohio
southward. The Virginia dogwood records of Nelson et al. (1938),
even if entirely C. florida, are low in volume although considerable
in rate of incidence. However, many of Smyth's winter records
were from Virginia and showed a high incidence of flowering dog-
wood, which indicates that the species is used where available.
Gilfillan and Bezdek ( 1944) found flowering dogwood fruits favored
highly in Ohio.

The dogwoods vary widely in type of plant and in their position
in the habitat. Bunchberry is a small, almost herbaceous plant
found in the ground cover of northern woodlands, usually in associa-
tion with hemlock or spruce. Silky cornel and red osier (see Plate
23D ) are medium-sized shrubs usually found on open streambanks
or in damp swales. Gray dogwood, roundleaved dogwood and
roughleaf cornel are medium-sized shrubs usually located in dry
or well-drained thickets in old fields or along woodland borders.
Flowering dogwood (see Plate 23C) and blue cornel are large
shrubs or small trees found in the woodland understory, mainly along
the edges.

The fruits of the dogwoods are eaten by many other birds includ-
ing the quail, ring-necked pheasant, wild turkey, and numerous song
birds, and by Virginia deer, gray fox, beaver, cottontail rabbit,
eastern chipmunk, and gray squirrel. Since the available dogwood
fruits are to a considerable extent exhausted each season, these ani-
mals compete with the grouse for this food. This competition may
cause the grouse to shift to a greater proportion of less palatable
foods, but is rarely an important factor in maintaining grouse
numbers.

Chemical composition: Gray dogwood fruit: water-3.6 per cent;
protein-10.1 per cent; fat-18.8 per cent; nitrogen-free extract-30.0
per cent; fiber-34.8 per cent; ash-2.8 per cent (Hosley, 1938).

Food and Water 117

Hazelnuts {Corylus spp.)

Species utilized: American hazelnut (C. americana), beaked
hazelnut (C. rostrata), the former probably constituting the bulk
of food produced by this genus in most areas. Selection between
species is probably determined by availability.

Seasonal importance: Twigs and buds are taken mainly in winter
and early spring, somewhat in late fall; the nuts are taken mostly
in the fall.

Parts used: Twigs and leaf buds, mainly nuts, and catkins to some

extent.

Geographical importance: Both species of hazelnut are used
throughout the northeastern region apparently with no marked
choice.

The hazelnuts are medium-height shrubs found in dry or well-
drained thickets along woodland edges and in old fields. The nuts
are eaten by many species of mammals, particularly gray squirrels,
and by several of the larger birds. The browse is also eaten by deer.
Competition for this food is of little importance to the grouse.

The Hawthorns (Crataegus spp.)

Species utilized: The species of Crataegus, of which over fifty are
recognized in northeastern United States, are difficult to identify.
So it is not surprising to find that the fruits and seeds in crops and
stomachs of grouse are determined only to genus. Two species,
C. coccinea (Judd, 1905), and C. crus-galli (Judd, 1905; Hosley,
1938 ) definitely have been recorded as used. It is probable that most
species are taken as available. Those that hold their fruit on the
branches well into the winter, and those whose fruit remains plump
and firm on the ground until spring, are probably most useful.
C. crus-galli, which occurs throughout the East, is one of the most
widely distributed natives of the Northeast that holds its fruit well.
C. phaenopyrum in Virginia, C. canbtji in Maryland and southern
Pennsylvania, and C. arnoldiana in Massachusetts and Connecticut
are of more limited range. There are others, introduced as well as
native, that also are useful in winter and spring.

Seasonal importance: The fruits are primarily important in fall

118 The Ruffed Grouse

and early winter, and to a lesser extent, as available, throughout the
winter and earh- spring. The leaves are a staple food in the late
spring and throughout the summer.

Parts used: Fruit and leaves, both very important.

Geographical importance: An important food in New York and
New England, except possibly along the southern New England
coast; of secondary importance from Pennsylvania southwards.

The hawthorns are tall shrubs found on old fields, along wood-
land borders, and in poor pastures, where they are often weeds.
They are valuable as ornamentals for their flowers and fruits, but
many species are subject to fungus diseases. The fruits are eaten by
many species besides the grouse, including the bobwhite, ring-
necked pheasant, at least thirty-six other species of birds, the gray
fox, white-tailed deer, and cottontail. Their competition for this
food, however, is unimportant to the ruffed grouse.

Chemical analysis: (Fresh fruit): water— 75.8 per cent; protein—
2.0 per cent; fat— 0.6 per cent; nitrogen-free extract— 20.8 per cent;
fiber-2.1 per cent; ash-0.8 per cent (Hosley, 1938).

Beech {Fagus grayidifolia)

Seasonal importance: Important only as a fall and early winter
food but probably one of the most highly selected of all preferred
foods of the grouse ( see Fig. 28 ) . Its importance, however, is weak-
ened by its irregular availability. It also appears to be considerably
less important in recent years than formerly due to the changed char-
acter of the beech composition of many farm woodlands ( see "Ef-
fects on Food of Cultural Operations-Lumbering"). These points
are well illustrated by Kuhn's (1940) records of grouse stomach
contents from Bradford County, Pa., the first week of November,
1911, the first week of December, 1919, and November 2-9, 1938.
The volume of beech nuts in the food was fifty and nine-tenths per
cent, seventy-two per cent, and zero, respectively, for these years.
There was a complete failure of the beechnut crop in 1938. The
trend toward elimination of the "worthless" old beech "wolf" trees
in woodlands has the effect of eliminating the only mast-producing
trees of the species. Nevertheless, the beech remains a very impor-
tant source of grouse food. Birds that have been feeding extensively
on beechnuts have a superior flesh flavor.

Food and Water 119

Parts used: Nuts, and to a negligible extent buds and leaves.

Geogiaphical importance: Ohio, Pennsylvania, New York, and
Northern New England, in areas where the beech-birch-maple-
hemlock woodland type is prevalent is the area of importance for
beech.

The beech is a large tree of the northern-hardwoods association
found mainly on sUghtly acid, loam soils in well-drained situations.
Its tree associations are mainly maples, birches, hophornbeams, black
cherry, hemlock, and white pine. It grows slowly, reaches a very
old age and sprouts profusely after cutting. It is considered by many
to be a forest weed because its wood is not as valuable as that of
leading timber species.

The triangular nut is borne in a prickly bur. Trees do not ordi-
narily fruit until they are from forty to fifty years old and then very
irregularly. It is eaten by at least fourteen species of birds in addi-
tion to the ruffed grouse, including the bobwhite, wild turkey, and
ring-necked pheasant. It is also taken by many mammals, among
them all the squirrels, foxes, raccoon, opossum, white-tailed deer,
black bear, porcupine, and cottontail.

Chemical analysis: (entire nut: water-2.3 per cent; protein-
13.0 per cent; fat-34.0 per cent; nitrogen-free extract-7.8 per cent;
fiber-40.8 per cent; ash-2.1 per cent; calories per pound-1820
(Hosley, 1938).

Wild Strawberries {Fragaria spp.)

Species utilized: There are two species of wild strawberries in the
Northeast, both of them common. Stomach analysis reports give all
strawberry as Fragaria sp. No doubt both F. virginiana, and F. vesca
var. americann are utilized although only the former has been
identified.

Seasonal importance: The fruit is a major summer food of both
adults and chicks. The leaves are eaten throughout the year, often
constituting a significant item at any season for young and old.
They are most important in summer, secondly, in winter.

Parts used: Fruit, leaves, and flowers.

Geographical importance: From the records there is some indica-
tion that the strawberry is less important in New England than in
New York and Pennsylvania. This is probably not actually true.

120 The Ruffed Grouse

since the New England grouse specimens were almost all taken in
the fall when the prevalence of many fruits makes strawberry tem-
porarily less important. The small quantity of strawberry leaves
taken in Virginia in early winter, even though the incidence is
fairly high, indicates that it may not be a higlily important winter
food there. It is probably more important in summer. We are quite
safe in saying, however, that Fragaria is important throughout the
region in summer, and from Pennsylvania northward throughout
the year.

The strawberries are low trailing herbaceous plants foimd in old
fields and in the ground cover of open woodlands. They flower in
May, fruit in June and July, and the leaves remain somewhat suc-
culent throughout the winter. Numerous birds and mammals par-
take of the ripe fruit but offer no serious competition to the grouse,
since the abundance of the fruit at the season of ripeness almost al-
ways exceeds the demands upon it.

WiNTERGREEN {Gaultheria procumbens)

Seasonal importance: The wintergreen, or checkerberry, is a food
consistently used throughout the year but which assumes significant
volume only from fall to spring. It is not generally taken in very
large quantities, but the regularity of use enhances its importance.
In spite of its relative unavailability under the winter snows, it is
most important as a winter and early spring food.

Parts used: Leaves, fruit.

Geographical importance: An important fall to winter food
throughout the Northeast, possibly used somewhat more in the
South due to greater availability through the winter.

The wintergreen is a low, evergreen plant of the woodland floor,
with creeping stems that are somewhat woody. It fruits only when
sunlight is available through openings in the crovm. The fruit is
eaten by the ring-necked pheasant, bobwhite, and chipmunks, and
the browse by white-tailed deer.

Chemical composition: (dried leaves) water-6.9 per cent; pro-
tein-6.0 per cent; fat-6.1 per cent; nitrogen-free extiact-60.1 per
cent; fiber-16.5 per cent; ash-4.4 per cent (Hosley, 1938).

Food and Water 121

Laurel {Kalmia spp.)

Species utilized: Mountain laurel (K. latifolia), and narrow-leaved
or sheep laurel (K. angustijolia) ; the former is more important by
far from southern New York and south, the latter is more important,
if only a secondary food, in New England.

Seasonal importance: Eaten during fall, winter and early spring,
by far most important during the winter when green leaves of most
plants are lacking.

Parts used: Leaves primarily, buds and flower capsules to some
extent.

Geographical importance: Mountain laurel is an important winter
food from the Catskills of southeastern New York southward, in-
creasing in importance to the south. In Virginia it was found in
nearly half the late fall and early winter stomachs and amounted to
almost eight per cent of the volume of food. It probably increases
in importance later in the winter. The narrow-leaved, or sheep
laurel, is used from New York through New England but apparently
is not ordinarily taken in significant amounts.

The laurels are low-to-medium evergreen shrubs found in acid
soils in the woodland understory and along woods margins. The
leathery leaves are poisonous to domestic livestock but apparently
have no ill effects on grouse or on deer if not taken to the exclusion
of other foods (Van Dersal, 1938).

Canada Mayflower {Maianthemum canadense)

Seasonal importance: Fall, winter and spring, with somewhat
greater volume taken in winter even though less available at that
time.

Parts used: Fruits.

Geographical importance: From the stomach records it appears
that Maianthemum is used only in New York and New England, al-
though the plant ranges much farther south. The fact that Penn-
sylvania and Virginia records taken during the season when the
plant is readily eaten farther north fail to show any trace of it
substantiates this conclusion.

Canada mayflower (also called two-leaved false Solomon's seal,

122 The Ruffed Grouse

and false lily of the valley ) is a low herb of the woodland floor, fre-
quently associated with conifers ( see Plate 20D ) .

Wild Apple {Mains pumila)

Species utilized: The records identify apple only to genus, but
the common wild apple (M. piimila) unquestionably supplies most
of the apple food taken. Cultivated apple is also eaten, as is evi-
denced by the damage done to apple orchards in New England,
and the native crabs are probably utilized too although there are no
specific records to prove this.

Seasonal importance: Primarily a fall, winter, and early spring
food, although taken somewhat throughout the year. Fruit is of
some importance from late summer to winter; the buds are taken
all winter and the blossoms and leaves when available.

Parts used: Leaf buds and twigs, fruit pulp, seeds, leaves, and
blossoms, in order of importance (see Plate 20C).

Geographic importance: An important food from fall to spring
from Maine to Pennsylvania, with greatest use in New England.
The New York records show some inconsistencies. Darrow found it
among the first ten important foods in only two seasons ( one of them
summer), and only in the Adirondack region of the state. The 1937
New York report failed to list it at all among the first ten for any
season, and Kelso's records show a sharp break in January that is not
logical, perhaps due to deficiencies in the material examined. In
Pennsylvania and Southern New England apple appears to be
somewhat less important while the species is little represented in
the Virginia records.

An interesting sidelight on the use of apple by grouse is the oc-
casional damage to cultivated orchards from budding by grouse.
This reached such serious proportions in parts of New England that
bounties were declared and damage claims paid. In the middle of
the nineteenth century certain Massachusetts townships paid a price
of twenty-five cents a head for "partridge." With the declining im-
portance of the apple industry, the bounties were abandoned but
damage has not entirely ceased. In 1924 New Hampshire paid out
nearly a quarter of the income from sportsmen's licenses (about
$25,000) on valid grouse damage claims (Bartlett, 1924), and as
recently as the mid-thirties, Massachusetts has paid orchard dam-

Food and Water 123

age claims on account of grouse. New Hampshire continues this
practice today, paying thirteen claims totaling $338.44 in 1940.
Highest annual payment was in 1926-$26,000. In the 1930-40
period, the highest was $3488.07 on seventy-five claims in 1935; the
lowest was $12.57 in 1938.

The wild apples are small-to-medium-sized trees, rather short-
lived and very hght demanding. They grow best on fertile soils but
often seed in on abandoned fields with acid and worn out soils.
They are subject to damage by numerous insect and fungus pests.

The browse and fruit of the apple are among the most important
wild-life foods in the Northeast. Hosley (1938) says "If all the ani-
mals using the apple were known it would probably rank as the most
universal wild-life food of the northeast." In addition to the grouse,
it is a staple food of the white-tailed deer, raccoon, red fox, and
skunk; is also eaten by tlie bobwhite, ring-necked pheasant, many
song birds, opossum, woodchuck, gray fox, red squirrel, snowshoe
hare, and cottontail. In spite of this competition for the fruit and
foliage of the apple, the buds that are most important to the grouse
are relatively secure.

Chemical composition: (fruit) water-63.3 per cent; protein-.3
per cent; fat-.3 per cent; nitrogen-free extracts— 10.8 per cent;
fiber-25.0 per cent; ash-.3 per cent; calories per pound-220 (Hos-
ley, 1938).

Partridgeberry {Mitchella repens)

Seasonal importance: Eaten rather consistently in a small way
throughout the year, the fruit from fall until spring, the leaves at all
times.

Parts eaten: Fruit and leaves.

Geographical importance: Mitchella is used about evenly through-
out the northeastern range.

A small, evergreen, trailing, herblike shrub, partridgeberry (see
Plate 21C) is found in the ground cover of dry woodlands, often
associated with conifers. The fruits are eaten by several other
species of birds besides the grouse including the bobwhite and by
the red fox. Their competition is of no significance to the grouse.

Chemical composition (dry leaves and stems): water-4.8 per
cent; protein-7.3 per cent; fat-2.2 per cent; nitrogen-free extiact

124 The Ruffed Grouse

—64.4 per cent; fiber~14.5 per cent; ash— 6.9 per cent (Hosley,
1938).

HoPHORNBEAM {Ostryo virginiana)

Seasonal importance: Winter and early spring; eaten to a lesser
extent in the fall, most important as a winter food ( see Plate 20C ) .

Parts used: Leaf buds, and, to a small degree, seeds (nutlets) in
the fall.

Geographic importance: Hophombeam is primarily important in
New York and Pennsylvania. It is apparently of slight importance
in New England and negligible south of Pennsylvania.

A small slow-growing tree of the northern-hardwoods and beech-
maple associations, it is found in the woodland understory on most
upland soils. The nutlets are eaten by ring-necked pheasant and
bobwhite quail as well as by grouse, and the browse is taken by the
white-tailed deer and cottontail rabbit. The competition is of no
significance to the ruffed grouse.

Ferns (Polypodiaciae)

Species utilized: For convenience, and because the food records
often show identity only to family, all genera of the fern family
are lumped together. Most important is that of the shieldferns
{Thelypteris) , which would rank in the first twenty-five food-
producing genera by itself. Spiny-toothed shield fern {T. spinu-
losum, including var. intermedium ) is probably used most, but mar-
ginal shield fern (T. marginalis) too is regularly eaten. Probably
other species also are used. Christmas fern {Polystichum acro-
stichoides) and polypody fern (Pohjpodium vtdgare) rank next in
importance. Several other species are known to be utilized, includ-
ing sensitive fern (Onoclea sensihilis) and royal fern (Osmunda
regalis), some being locally important. The grape ferns {Botrychium
ohliqiium, B. dissectum, and possibly others), while not in the fam-
ily Polypodiaciae, are closely related plants that are eaten to some
extent.

Seasonal importance: Ferns are eaten throughout the year and are
a staple food in all seasons except summer. While the spring rec-
ords are inadequate for complete analysis over the whole region, it

Food and Water 125

is probable that ferns are of greatest use at this season. In the New
York state-wide average (N.Y.S. Cons. Dept. Ann. Rep. 1937) ferns
appear in the first ten only in the spring. Their importance in winter
is substantiated by abundant data. The relative inaccessibility of
the plants at this season in the northern states adds even greater
weight to the evidence of their importance. In autumn ferns are
taken in significant amount but in lesser quantity than from De-
cember to May.

Parts used: Fronds.

Geogiaphical importance: The ferns are an important food
throughout the Northeast. There is evidence that shield ferns
(Thelypteris) (see Plate 20A) are of greatest value in the northern
range, and that Christmas fern (Polystichum) is more important in
Virginia, and rattlesnake fern (Botrychium) in Ohio. This differ-
entiation is probably the result of local variation in abundance.

Christmas fern is found in the ground cover of dry woodlands
while the shield ferns are adapted to the floor of damp woods. Both
are most prevalent on the banks of ravines.

Popples, or Poplar {Populus spp.)

Species utilized: Quaking aspen (P. tremuloides), large-toothed
aspen (P. grandidentata) , and balsam poplar (P. balsamifera) are
eaten, the first two as staple foods, the third usually incidentally.

Seasonal importance: An important food all year round, used
most intensively in the spring (buds and catkins). The buds are
also one of the most important foods all through the winter. During
the spring and summer the catkins and leaves are eaten by adult
grouse, while in the fall the birds gradually take to the buds again.

Parts used: Leaf buds, flower buds, catkins, and leaves, the buds
being by far the most important ( see Plate 20C ) .

Geographical importance: From Pennsylvania and Ohio north-
ward for P. tremuloides and P. grandidentata. The range of P. bal-
samifera restricts its use to New England, northern New York, and
northward.

The popples are short-lived, small-to-medium-sized trees that es-
tablish themselves quickly on old fields and bums when seeding
conditions are right. In climax or subclimax stands they are re-
stricted largely to the forest edges.

126 The Ruffed Grouse

Popples are cut by beavers as one of their most important foods.
Locall)^ this competition may well affect tlie grouse. These trees are
also browsed by the white-tailed deer and snowshoe hare but with-
out significant competition for the grouse.

Chemical analysis—?, grandidentata, fresh buds: water— 49.8 per
cent; protein— 6.4 per cent; fat— 5.1 per cent; nitrogen-free extract—
24.1 per cent; fiber— 13.3 per cent; ash— 1.4 per cent (Hosley, 1938).

Cherries {Prunus spp.)

Species utilized: Black cherry (P. serotina), pin cherry (P. penn-
sylvanicum) , chokecherry (P. virginiana), all these three provide
staple foods in approximately that order of importance and furnish
most of the subsistence derived from plants of the genus. However,
wild plums (P. americana), sweet cherry (P. avium), plum (P. do-
mestica var. ), and peach (P. persica var.) also contribute more or
less to the diet of the grouse.

Seasonal importance: The cherries furnish an important part of
the food supply throughout the year. Both adults and young con-
sume the fruit during summer, fall, and into the winter, as long as
available. Buds are taken in fall, winter and spring. The cherry
genus probably assumes greatest importance as a food resource in
winter, increasing steadily from December to March, but ranks
high also in early spring, summer, and fall. Use is less in late spring
than at other times.

Parts used: Fruit, buds (see Plate 20C) and twigs, and leaves.

Geographical importance: The cherries are among the most im-
portant grouse food producers from Pennsylvania northward. From
Nelson's report they seem to be of little value in Virginia; however,
this finding from a limited area and period may not hold in general.

Black cherry (see Plate 26A) is a large tree of the Appalachian
highlands, reaching its best growth in the hemlock-hardwoods type.
To the north and south of its optimum range it is usually found
considerably dwarfed. The fruit ripens in August and September
and is taken from tree or ground throughout the fall and into the
winter.

Pin cherry ( see Plate 26B ) is a small, short-lived tree, most com-
monly associated with recent burns. It is often found in nearly pure
stands in bums or old fields, but is ultimately entirely displaced by

Food and Water 127

subclimax and climax species. The fruit ripens from July to Septem-
ber and is eaten until early winter from tree and ground.

Chokecherry ( see Plate 22C ) is a large shrub or small tree found
abundantly in hedge-rows and along woodland borders. As with the
other two species above, it is widely adaptable to a variety of soils.
The fruit ripens in late summer and is eaten through the fall and
into the vmiter.

All of these cherries provide food also for the bobwhite, ring-
necked pheasant, numerous song birds, and the cottontail. Black
cherry is also a food plant of the white-tailed deer, red fox, raccoon,
squirrels, and black bear; pin cherry for the white-tailed deer, chip-
munk, and beaver; and chokecherry for the black bear. As far as
the fruit is concerned, this competition may sometimes aflFect grouse
food habits locally, but not seriously. Competition for leaves and
buds is of no importance, except in spots where beavers may destroy
small stands of pin cherry.

Oaks (Quercus spp.)

Species utilized: White oak (Q. alba), northern red oak ( Q. ho-
realis), red oak (Q. rubra), bear oak {Q. ilicifolia), pin oak (Q.
palustris), chestnut oak {Q. montana), and chinquapin oak {Q.
prinoides), in the approximate order of their importance. As most
of the records show identity only to the genus, it is impossible to
appraise accurately the relative importance of the several species
with the information now available ( see Plate 22A ) .

Seasonal importance: The oak group is one of the most important,
and at time?, and places the most important source of fall food.
Where snow conditions permit, it is also a valuable winter food.
Acorns appear more prominently in the written records than is war-
ranted because of the predominance of fall records, that is, from
grouse killed in the hunting season. That is really the only period
when oak mast is of great importance. Even though not available
throughout the year, acorns still are a very important grouse food.

Parts used: nuts (acorns).

Geographical importance: The area of greatest use for acorns is
middle and southern New England, and from Ohio and Pennsyl-
vania southward. The acorns are of moderate importance in northern
New England and in New York outside the Adirondacks. The value

128 The Ruffed Grouse

of the oaks varies immensely in different years according to which
species, ff any, have a good crop of acorns. This results in somewhat
inconsistent records from a single area, depending on years of
collection. Smyth in 1919-23, with a considerable proportion of
New York records, found relatively little utilization of acorns from
October to March, and none in November when it should have been
at its peak. Kelso's winter records taken prior to 1935 show very
little consumption of acorns, and the New York State Conservation
Department's Annual Report in 1937 pointed out the notable scar-
city of foods of this group in the specimens taken by the study from
1930 to 1937. But Darrow two years later in summarizing the same
records, together with new accessions, reported oak foods in ex-
cess of three per cent in the whole year food averages in New York
outside the Adirondacks.

White oak is a large tree of the climax association of the Appala-
chian and Transition zones. It tolerates most soils except wet ones,
but prefers fertile sites. It is among the least dependable of the oaks
in acorn production, having a good crop only once in every three
years or longer. However, white oak acorns seem to be most palata-
ble of all and are small enough for the grouse to swallow whole.

Northern red oak is a large tree of the hemlock-hardwood associa-
tion. It is more shade tolerant than white oak and, therefore, repro-
duces better under old-field white pine. The acorns are large and
this probably hinders their utilization by grouse to some extent.
Crops are usually produced in alternate years. South, and at lower
altitudes, northern red oak is replaced by the common red oak which
is a very similar tree.

Bear oak and chinquapin oak are shrub species found on dry,
sandy barrens and rocky hillsides. Their habitat keeps them out of
the better grouse coverts in most regions, but their patronage is
considerable where available to the birds.

Chestnut and pin oak acorns are known to have been taken by
grouse, but probably are not very important. It is probable that
those of other oaks too are taken by grouse in varying quantities.

The oaks are an important food-producing group for many species
of wild life. Acorns are a staple for the bobwhite, wild turkey, squir-
rels, white-tailed deer, wood duck, and mallard. They are also eaten
by the ring-necked pheasant, mourning dove, numerous song birds,
and by the opossum, muskrat, raccoon, black bear, gray fox, red

Food and Water 129

fox, and no doubt other species of mammals. The supply of acorns
often being exhausted by this heavy drain, these competing species
no doubt affect grouse food habits by forcing the birds to shift
locally to less palatable foods. Over most of the northeast range, this
does not materially aflFect grouse numbers or general distribution,
but in the oak-hickory type range from Pennsylvania southward, it
probably is a factor in determining tlie carrying capacity for grouse.
Chemical analysis of acorns ( Hosley, 1938 ) : water— 2.4 per cent;
protein— 7.1 per cent; fat— 4.9 per cent; nitrogen-free extract— 81.0
per cent; fiber— 2.2 per cent; ash— 2.5 per cent. Q. rubra (Wright,
1940): water— 1.8 per cent; protein— 7.6 per cent; fat— 22.6 per cent;
nitrogen-free extract— 61.0 per cent; fiber— 4.8 per cent; ash— 2.2
per cent.

Sumacs {Rhus spp.)

Species utilized: Dwarf sumac {R. copallina), smooth sumac {R.
glabra), poison ivy {R. toxicodendron), staghom sumac (R. ty-
phina) (see Plate 24), and poison sumac (R. vernix). All five are
staple food producers in parts of the northeast range, and the group
as a whole is consistently important over the whole Northeast. Other
species also patronized are fragrant sumac {R. aromatica) and
lemonade sumac (R. trilobata).

Seasonal importance: All of the sumacs, including both the poi-
sonous and nonpoisonous types, are of value to the grouse in late
fall, through the winter, and in early spring. They are utilized to a
minor extent in October and May, at the extremes of the season of
greatest consumption.

Parts used: The fruits, heads of which are called "bobs."

Geographical importance: The sumacs as a group are important
throughout the Northeast, and generally to the same degree. The
total of Rhus products taken seems to average quite consistently,
during the seasons used, from one to six per cent of the food. Geo-
graphical variation in importance of the five key species is probably
determined in the main by availability. From the records it appears
that R. copallina is most used in New York, Pennsylvania, and Vir-
ginia, R. glabra in Connecticut, Ohio and Virginia, R. toxocoden-
dron in New Hampshire, Massachusetts, and Rhode Island, R. tij-
phina in Maine, New Hampshire, New York, Pennsylvania, Ohio

130 The RufiFed Grouse

and Virginia, and JR. vernix in Maine and Massachusetts. Sum-
marized by states, the species most patronized are: Maine, R. ty-
phina, and R. vernix; New Hampshire, R. toxicodendron, and R.
typhina; Vermont, none; Massachusetts, R. toxicodendron, and R.
vernix; Rhode Island, R. toxicidendron; Connecticut, R. glabra; New
York, R. copallina, and R. typhina; Pennsylvania, R. copallina, and
R. typhina; Virginia, R. copallina, R. glabra, and R. typhina; Ohio,
R. aromatica, R. typhina, and R. glabra.

Dwarf sumac, contrary to its name, is not always a small shrub,
but sometimes grows to be a large shrub or even a tree. It is a
sparsely-branched, thicket-forming plant, usually found on acid
soils on dry or well-drained sites, either in full sun or partial shade.

Smooth sumac is a small-to-large shrub, sparsely branched, some-
times thicket-foiTning. It is found only in full sunlight, but is adapted
to either dry or wet soils.

Poison ivy is a vine, or sometimes a shrub, very bushy, and at
home in both sun and shade. It is adapted to dry or well-drained
soils. The fruit is a smooth wax-covered drupe, quite unlike the hairy
drupes of the non-poisonous species.

Staghorn sumac is a large shrub or small tree tliat often grows in
loose thickets. It is found mainly in full sun, but grows occasionally
in partial shade. Dry or well-drained soils on old fields and woods
borders are its usual habitat. Poison sumac is a large shrub or small
tree, rather sparsely branched, found on most soils from open sun
to rather overgroMoi bogs or swamps.

Fruits of all the sumacs are eaten by a gieat variety of birds and
mammals. Those of poison ivy alone are known to be consumed by
more than sixty species of birds. Poison sumac berries are an impor-
tant food of the bobwhite and ring-necked pheasant. Drupes of
most of the species are eaten by these two game birds, mourning
dove, wild turkey and by cottontail rabbit and white-tailed deer.
Staghorn sumac is important as a deer browse food and smooth
sumac as a rabbit food. Competition for products of the abundant
sumacs is of no particular significance to the grouse.

Chemical analysis (Hosley, 1938, species not named): of fresh
bobs: water— 9.6 per cent; protein— 6.9 per cent; fat— 17.2 per cent;
nitrogen-free extract— 38.1 per cent; fiber— 22.9 per cent; ash— 5.3
per cent. Rhus toxicodendron (Wright, 1940): water— 3.4 per cent;
protein— 6.8 per cent; fat— 26.1 per cent; nitrogen-free extract— 32.2

Food and Water 131

per cent; fiber— 30.1 per cent; ash— 1.5 per cent. R. copallina (Wright,
1940): water— 4.1 per cent; protein— 4.9 per cent; fat— 11.2 per cent;
nitrogen-free extract— 46.6 per cent; fiber— 31.0 per cent; ash— 2.3
per cent. The amount of fat is surprising, being exceeded in only a
few other species such as Fagus and Juglans. This may explain why
this seemingly rather unpalatable food is so well liked by wild life.

Wild Roses (Rosa spp.)

Species utilized: Because of tlie difBculty of identifying the spe-
cies of rose represented by the remains found in grouse crops or
gizzards, and because the classification of these plants is rather con-
fused, the records give only the genus. It is probable that fruits of
any of the roses found in grouse habitat are eaten; at least we do not
know that any species are not used ( see Plate 25C ) .

Seasonal importance: Fall, winter, and early spring.

Parts used: Fruit (hips) and, to a minor extent, the leaves.

Geogiaphical importance: The records indicate that this genus
is more important as a food producer for grouse in Virginia than
farther north. In Pennsylvania it appears to be definitely secondary,
while in New York and New England it is faiily consistently used,
and composes around one to two per cent of the fall to early spring
food.

The wild roses are shrubs, often climbing to some degiee, or low-
growing. The flora includes both natives and acclimated exotics,
often difficult to identify. The fruits vary immensely in size from the
pea-size of those of R. multiflora to the small tomato-sized hips of
R. rugosa; most of the species produce intermediate-sized fruits
about half an inch in diameter. They are usually retained on the
plant the year round. The roses generally require full sunlight and
fairly fertile, well-drained soil, although some thrive on poor or wet
soils.

The fruits, or hips, are eaten by many birds, including the bob-
white, ring-necked pheasant, and wild turkey, and by the opossum,
white-tailed deer, red squirrel, porcupine, snowshoe hare, and cot-
tontail rabbit. Their competition for this plentiful food is of no
importance to the grouse.

Chemical composition (Hosley, 1938, for R, mtdtiflora, fresh
fruit): water-29.5 per cent; protein— 7.9 per cent; fat—3.5 per cent;

132 The Ruffed Grouse

nitrogen-free extract— 41.2 per cent; fiber— 14.9 per cent; ash— 3.0
per cent.

Brambles, or Blackberries and Dewberries {Rubus spp.)

Species utiUzed: The majority of records identify bramble foods
only to genus but a number are specifically known to be eaten by
grouse. These are blackberry {R. allegheniensis) (see Plate 21A),
trailing blackberry (R. hispidus), wild red raspberry (R, idaeus
aculeatissinius) , thimbleberry (R. occidentalis), salmonberry (R.
parviflorus) , and dewberry (R, villosus). No doubt other species
also are used. We can surmise that the most important species are
those most prevalent in grouse habitat. These are mainly R. alle-
gheniensis, R. idaeus aculeatissinius, and R. hispidus.

Seasonal importance: A very important summer food for both
chicks and adults, and also of value during the fall and early winter
months. Use of the fruit begins in July and continues through
August. The leaves are eaten in the summer, fall, and early wdnter,
and the buds in fall and winter.

Parts used: Fruit, leaves, and buds.

Geographical importance: One of the highest ranking summer
foods of adults and chicks tluoughout the Northeast, in New York
the most important. During the fall and winter their consiunption
is much reduced, but still remains significant. The records indicate
that they may be less important in the extreme North (Maine) and
the extreme South (Virginia) of the grouse's northeastern range in
the fall and winter, but this may not be a finally dependable con-
clusion.

The brambles are low-growing, sometimes trailing, shrubby plants
found in old fields, slashings, and bums (see Plate 21A). They often
form dense thickets. Their adaptability to poor soils enables them
to pioneer on abandoned, worn-out soils. When other woody plants
seed in, it is not long before the brambles are killed off by shade
of which they are very intolerant.

Rubus products are one of the most important of wild-lffe foods.
They are staple for the bobwhite, ring-necked pheasant, cottontail,
skunk, and white-tailed deer as well as for the ruffed grouse. Alto-
gether, more than 140 species of nongame birds in addition to
the opossam, cliipmmik, red fox, gray fox, red squiirel, black bear.

Food ajid Water 133

l^eaver, and snowshoe hare are known to eat them. There is no indi-
cation, however, that this heavy competition adversely a£Fects the
grouse.

Chemical composition (Hosley, 19-38, average figures for black-
berry fruit) : water— 86.3 per cent; protein— 1.3 per cent; fat— 1.0 per
cent; nitrogen-free extract— 10.9 per cent; fiber— 2.5 per cent; ash—
0.5 per cent. (Hoslev, 1938, dry leaves of R. hispidus): water— 7.8
per cent; protein— 9.8 per cent; fat— 4.1 per cent; nitrogen-free ex-
tract—60.8 per cent; fiber— 13.7 per cent; ash— 3.9 per cent.

The high water content of the fniit of Rubus may explain its great
importance as a summer food. During the hot summer montlis
grouse obtain much of their water requirement through foods of
this type.

Sheep Sorrel {Rtimex acetosella)

Seasonal importance: From late summer or early fall imtil mid-
spring, with greatest use probably in late fall and early winter.

Parts used: Leaves.

Geographical importance: Sheep soitcI is eaten by grouse through-
out the Northeast but assumes its greatest importance in the south-
em portion, from Pennsylvania to Virginia. Here it ranks as one of
the most important fall and winter foods whereas farther north it is
used somewhat less, particularly in winter when snow limits its
availability. It is taken to some extent by grouse chicks in late sum-
mer. Darrow found it to rate tenth among chick food plants in the
Catskill portion of New York.

Sheep sorrel (or field sorrel) is a common weed (see Plate 20D)
that has been naturalized from Europe. It is a perennial herb which
spreads hv rootstocks. The leaves have a sour taste. It is a low plant,
from five to fifteen inches high, found along field borders in sparse
cover, in old fields, on roadsides, and in poor soils.

Greenbriars (Smilax spp.)

Species utilized: Most of the records give the identification of
foods derived from this group only to genus. Products of three spe-
cies have been positively identified in grouse stomach records but
it is highly probable tliat those of other Smilax also are used. Cat

134 The Ruffed Grouse

greenbriar (Nelson, 1938) (S. glauca) and common greenbriai* (S.
rotundifolia) are among die most important species patronized.
The fruits of the carrion flower (S. herbacea), an herbaceous species,
also are known to have been taken.

Seasonal importance: A fall and winter food, probably most valu-
able in deep winter although used in greatest volume in late fall and
early winter.

Parts used: Fruit, mainly, and leaves.

Geographical importance: The greenbriars assume their greatest
importance as food producers in the soutliem part of the grouse's
northeastern range, notably Virginia and Ohio, where they rank
first as a fall and winter food. They are also valuable in Pennsyl-
vania and Rhode Island. Elsewhere they are of secondary ranking,
especially in the higher altitudes and more northern areas.

The important greenbriars aie spiny, shrubby, thicket-forming
vines. The nonwoody S. herbacea, although patronized by grouse,
is not of great importance. Of the two leading species, S. gluuca is
found on dry or well-drained soils, while S. rotundifolia occurs in
moist to well-drained sites. Both prefer fertile bottom lands. They
are primarily plants of the woods, being most prolific along wood-
land edges. The young plants produce good crops of fruit, while old
growth is usually barren.

Greenbriar berries are known to be eaten by the bobwhite, ring-
necked pheasant, wild turkey, and about forty other kinds of birds;
also by the opossum, gray fox, white-tailed deer, and cottontail rab-
bit. This competition is normally not serious to grouse but might
affect their local distribution, and possibly even abundance, in the
southern part of their range.

Chemical analysis (Wright, 1940): S. rotundifolia (dried fruit):
water— 2.8 per cent; protein— 9.3 per cent; fat— 5.3 per cent; nitrogen-
free extract— 60.8 per cent; fiber— 18.8 per cent; ash— 3.1 per cent.
S. glauca ( dried fruit ) : water— 3.7 per cent; protein— 10.9 per cent;
fat— 7.7 per cent; nitrogen-free extract— 57.1 per cent; fiber— 18.4
per cent; ash— 2.9 per cent.

Blueberries (Vaccinium spp.)

Species utilized: Lowbush blueberry (V. angustifoliinn) , Canada
blueberry (V. canadense), highbush bluebeny (V. corymhosum) ,

Food and Water 135

cranberry (V. macrocarpon) , deerberry (V. stamineum), dryland
blueberry (V. vacillans), cowberry (V. vitis-idaea) , and rock cran-
berry (V. vitis-idaea minus) are all known to be eaten by grouse.
It is entirely probable that numerous other species of blueberries
also are taken. The relative importance of the several species is
probably determined largely by their distribution and abundance.
The lowbush blueberiy is most important in southern New York,
followed in order by highbush blueberry, deerberry, and dryland
blueberry.

Seasonal importance: There are two distinct seasons of use for
Vaccinium: July to August for the fruit by both adult and young
grouse; and fall, winter, and early spring for the buds and twigs,
with the most intensive use from November through January. The
leaves also are eaten to a considerable extent in summer and fall.

Parts used: Fruit, in summer, by both adults and young; buds and
twigs in fall, v^rinter and early spring; and leaves in summer and fall.
The fruit and buds are the parts most used.

Geographical importance: As a summer food, the fruit is impor-
tant to both adults and yoimg throughout the Northeast. In New
York it ranked higher in the Adirondacks than elsewhere. During
fall and winter the buds are used in all parts of the Northeast, but
with apparently less intensity in New York than elsewhere. Possibly
western New England is also included with New York in this area
of restricted importance.

The blueberries are low-to-medium-height shrubs ( see Plate 21B )
found mainly on acid soils in open fields and along roadsides and
woodland edges. The more important species for grouse are those
growing in dry sites, but several species are confined to bogs. They
are pioneers in plant succession and often become established fol-
lowing a burn. Some are quite intolerant of shade and competition.
The native, wild blueberries are highly prized for human food and
are an important crop in some areas. Blueberry products, mainly the
fruit, are eaten by many species of songbirds, by the ring-necked
pheasant, bobwhite, Hungarian partridge, skunk, white-tailed deer,
opossum, cottontail, black bear, raccoon, and moose. Most of this
competition comes in the summer when food is plentiful. There is
normally little possibility of its having a serious effect on either
distribution or abundance of ruffed grouse. Harvest of blueberries
by man probably affects grouse distribution locally.

136 The Riiffed Grouse

Chemical analysis (Hosley, 1938) for fresh fruit: water— 83.4 per
cent; protein— 0.6 per cent; fat— 0.6 per cent; nitrogen-free extract
—13.9 per cent; fiber— 1.2 per cent; ash— 0.3 per cent; calories per
pound— 310.

Viburnums (Viburnum spp.)

Species utihzed: mapleleaf viburnum (V. acerifoliuni) (see Plate
23A); withered (V. cassinoides); nannyberry (V. lentago) (see
Plate 27A); blackhaw (V. prunifolium) (see Plate 27C); and high-
bush cranberry (V. trilobum) (see Plate 25B) are the species used
most generally and in greatest quantity. Among these, naimyberry,
highbush cranberry, and withered appear to be most important in
New England, mapleleaf viburnum, nannyberry, and withered in
New York, and blackhaw and mapleleaf viburnum to the south.
Other viburnums used to a lesser degree are: hobblebush (V. alni-
folium) (see Plate 27B); arrowwood (V. dentatum) (see Plate
27D); wayfaring tree (V. lantana) (usually cultivated); squash-
berry (V. paucifiorum) ; and hairy naimyberry (V. pubescens) .

Seasonal importance: Fall and winter, diminishing in volume of
use as winter passes.

Parts used: Fruit.

Geographical importance: Some species of viburnums are of sig-
nificance in the fall diet of tlie grouse throughout the Northeast,
with the possible exceptions of Vermont and Rhode Island. How-
ever, if more records from these states were available, it is reason-
ably sure that they would show that at least a small part of the
grouse food is derived from plants of this genus. The highest use
intensities are recorded from New Hampshire, Massachusetts, Con-
necticut, New York, and Virginia.

It is interesting to note a discrepancy in the geographical use of
a food species in connection with highbush cranberry. This species
is readily patronized by grouse in New England and Pennsylvania,
and yet we have observed that it is almost completely avoided in
New York. No explanation for this variation in selection is apparent.

The viburnums are medium-to-tall shrubs found in the woodland
understory, along woods edges, and in fence rows. They usually
grow in well-drained, fairly fertile soils, although nannyberry and

Food and Water 137

some others can endure rather wet soils too. They are eaten by the
lx)bwhite, ring-necked pheasant, and some twenty-odd species of
songbirds, as well as by the chipmunk, cottontail, porcupine, rac-
coon, skunk, gray squirrel, moose, gray fox, and white-tailed deer.
This competition is apparently of no importance to the grouse.

Chemical analysis (Hosley, 1938): fresh fruit, V. cassinoides:
water— 72.6 per cent; protein— 1.7 per cent; fat— 2.9 per cent; nitro-
gen-free extract— 19.8 per cent; fiber- 2.3 per cent; ash— 0.8 per
cent. V. opulus (almost identical with V. trilohum), fresh fruit:
water— 47.3 per cent; protein— 2.7 per cent; fat— 4.9 per cent; nitro-
gen-free extract— 37.1 per cent; fiber— 6.3 per cent; ash— 1.7 per cent.

Wild Grapes {Yitls)

Species utihzed: Summer grape (V. aestivalis), blueleaf grape (V.
argentifolia) y frost grape (V. cordifolia), fox grape (V. lahrusca)
(see Plate 28), riverbank grape (V. vulpina), and probably others
not specifically identified as yet in grouse food habit records. All of
the five species known to be eaten are taken in quantity where avail-
able; hence, their importance depends mainly on distribution and
abundance.

Seasonal importance: Fall and early winter is the season of great-
est use, quickly falling off from January on as the fruit is consumed
by wild hfe or drops off the vines. Some species hold part of their
fruit well into the winter when it is used as long as available, and
some fruit is picked from the ground.

Parts used: Fruit.

Geographical importance: Grapes are an important fall and early
winter food throughout the northeastern states, with the exception
of Maine. The areas of greatest use are southern New England,
southeastern New York, and from Ohio and Pennsylvania to Vir-
ginia. In the north. New York and New England, V. argentifolia, V.
luhrusca, and V, vulpina are most important; to the south, notably
in Virginia, V. aestivalis and V. cordifolia are most used. As a group
the wild grapes are among the top-ranking foods from October to
January, often exceeding all others in volume consumed.

The grapes are high-climbing, vigorous vines found mainly along
woodland borders and fence rows. They generally prefer a fairly

138 The Riiffed Grouse

fertile soil but occur on a variety of sites from dry to wet. They
require lots of Hght to fruit well and climb to the top of the tallest
trees to gain full exposme to the sun.

Grapes are a favorite food of many kinds of wild life besides
grouse. The fable of the fox and the grapes indicates the taste for
grapes that characterizes both the red and gray foxes. These fruits
are also a staple food of the ring-necked pheasant, bobwhite, and
skunk; and are eaten also by the wild turkey, mourning dove ( seeds )
and Hungarian partridge. Nearly a hundred kinds of songbirds are
known to eat grapes. Among mammals, other than foxes and skunks,
that eat grapes are the raccoon, opossum, white-tailed deer, red
squirrel, and cottontail. This competition very likely alters the local
distribution of grouse, and may sometimes affect abundance locally.

Chemical analysis ( Hosley, 1938, frost grape— fresh fruit ) : water
—11.6 per cent; protein— 9.5 per cent; fat— 4.8 per cent; nitrogen-
free extract— 43.7 per cent; crude fiber— 26.4 per cent; ash— 4.0 per
cent. Vitis sp. (dried fruit) (Wright, 1940): water— 8.4 per cent;
protein— 7.4 per cent; fat— 2.1 per cent; nitrogen-free extract— 56.1
per cent; fiber— 22.8 per cent; ash— 4.0 per cent.

SECONDARY GROUSE FOODS

The twenty-five genera supplying the greatest volume of food
throughout the year have been discussed in some detail as the pri-
mary foods of the species. There are numerous other foods that
either furnish a significant amount of food generally or are of special
value in restricted areas. These secondary foods are here arbitrarily
restricted to those that have constituted one per cent or more of the
seasonal food of grouse in the records of a study covering one or
more states. Added to the volume of the twenty-fi^ e primary foods,
from ninety-four to ninety-nine per cent of the birds' normal food
in the Nortlieast is covered by the two groups. The forty-one genera
of plants and the insects and miscellaneous animal foods that are
included in the category of secondary foods are here annotated.

Maples (Acer) : Buds and seeds of at least four species are taken.
Seeds composed two and three-tenths per cent of the summer food
of grouse chicks in the Adirondacks of New York, one and one-tenth
per cent in the rest of the State outside the Catskills, and from
two and four-tenths to three and nine-tenths per cent of summer

Food and Water 139

food at different times in New Hampshii^e (MacGregor). Maple
buds are listed eightli among winter foods and tenth among spring
foods in the Adirondacks (Darrow), ninth among winter foods in
New York (IV. Y. S. Cons. Dept. Ann. Rep. 1937) and as over one
per cent of winter food in the Northeast ( Bi-1297 ) and recorded in
small amounts throughout the region from Maine to Virginia. The
most palatable food producer appears to be mountain maple.

Alders (Alnus): Buds and seeds of two species have been re-
corded as grouse foods, but probably others are sampled too. Alder
products were recorded by Smyth as composing four and eight-
tenths per cent of grouse food in October, three and three-tenths per
cent in November and two and two-tenths per cent in winter in the
Northeast; they are taken in small amounts during the winter in all
parts of the Northeast. Speckled alder is probably the most impor-
tant species.

Serviceberries (Amelanchier) : Buds have been listed as a fall and
winter food. The fruit is used during the summer and fall by both
young and old buds. Only A. canadensis has been specifically identi-
fied but other species no doubt also are patronized. Darrow recorded
Amelanchier products as comprising five and two-tenths per cent of
the summer food of grouse chicks in the Catskill region of New York
and one and one-tenth per cent in the remainder of the state outside
the Adirondacks. Kuhn (1941) records them as forming two and
three-tenths per cent of the November food in Pennsylvania. In Vir-
ginia the buds amounted to one and one-tenth per cent of the early-
winter food (Nelson). Remains of these plants have also been
reported as a fall and winter food in small quantities in Pennsyl-
vania, New York, and New Hampshire ( see Plate 30E ) .

Everlasting (Antennaria): The leaves are taken during fall and
winter but have not been definitely identified as to species. Nelson
found the early winter food in Virginia to have been derived to the
extent of two and three-tenths per cent from plants of this genus.
It has been listed as a moderate food producer in other records from
Pennsylvania to New England.

Chokeberry (Aronia): Both fruit and buds of the two common
species are taken in fall and winter from New York to Maine. In
Massachusetts they composed one and eight-tenths per cent of the
fall and winter food, in Rhode Island one and two-tenths per cent,
and in Connecticut one and seven-tenths per cent (Hosley). Red

140 The Rufled Grouse

chokeberry appears to be used most in New England and black
chokeberry ( see Plate 29A ) in New York.

Aster (Aster): The leaves of asters, species not identified, are used
as a fall and early winter food from New York to Virginia. They
composed three and tliree-tenths per cent of the November-Decem-
ber food in Virginia (Nelson) and appear to be more important in
the South than in the North.

Barberries ( Berberis ) : The fruit of both the European and Thun-
berg barberries is eaten to some extent during the winter in New
York and New England. It composed three and six-tenths per cent
of the total fall-and-winter food in Massachusetts and one and two-
tenths per cent in Rhode Island, with lesser amounts from New
Hampshire to New York ( Hosley ) . MacGregor found it to compose
one and five-tenths per cent of the October food in New Hampshire.

Blue beech ( Carpinus caroliniana ) : Both buds ( see Plate 20C ) .
and seeds are taken in late fall and winter, the buds being the more
important. Parts of this plant composed four and one-tenth per cent
of the winter food in New York ( December eight and seven-tenths
per cent, January three and one-tenth per cent, February four and
five-tenths per cent (Kelso); three per cent of fall-and-winter
food in New York (Hosley), two and five-tenths per cent of year-
round food in New York outside the mountains (Darrow). In other
states blue beech products appear to be Httle used, although they
have been recorded in grouse food from Virginia to New Hamp-
shire.

Chestnut (Castanea dentata): An important food prior to the
sweep of the chestnut blight (Judd, Smyth), now no longer signifi-
cant. As late as 1923, Smyth found that it composed one and four-
tenths per cent of the October food in tlie Northeast. It is noted here
as a matter of history, and a hope for the future.

Climbing bittersweet (Celastrus scandens) : (see Plate 29B). The
fruit is eaten during fall and winter from New York to New Hamp-
shire. It composed one per cent of the fall and winter diet in New
Hampshire (Hosley) and over one per cent of the winter food in
the Northeast (Bi-1297).

Beech-drops ( Epifagus virginiana ) : Seeds composed one and one-
tenth per cent of October food in the Northeast ( Smyth ) .

Trailing arbutus (Epigaea repcns): Leaves and buds are eaten
in fall and winter; they composed one and two-tenths per cent of the

Food and Water 141

November-December food in Virginia (Nelson) and are also taken
throughout the region in small quantities.

Buckwheat {Fagopyrum esculentum) : The seed is a fall food,
mainly in Pennsylvania and New York. It composed one and two-
tenths per cent of the November food in northeastern records,
slightly less in October ( Smyth ) .

Huckleberries ( Gaylussacia ) : Fruit and buds are taken in sum-
mer and winter, respectively. Of the two species definitely identified,
black huckleberry is the more important, but other species probably
are used too. Huckleberries were recorded by Hosley as contribut-
ing one per cent of the fall and winter food in Massachusetts. They
were lumped with blueberries (Vaccinium) in the significant use
records of Kuhn in Pemisylvania and Nelson in Virginia. They have
been recorded as utilized in other New England states as far north-
east as New Hampshire.

Avens {Geum): Seeds and leaves are taken in fall and winter.
Only one species has been definitely identified (G. strictum) but
others also are probably patronized. Composed one and two-tenths
per cent of October food in the Northeast (Smyth); smaller but
significant quantities have been recorded as taken by grouse from
Vii-ginia to New England.

Witch hazel (Hamaiiielis mrginiana): The seeds, flowers, and
buds ai'e taken in rather small but consistent quantities during fall
and winter from Maine to Virginia. They composed one per cent of
the f all-and-winter food in New Hampshire ( Hosley ) ; one per cent
of the November diet in the Northeast ( Smyth ) ; and two and six-
tenths per cent of November subsistence in Pennsylvania (Kuhn,
1941).

Holly (Ilex): The fruit, and to a small extent the leaves, of at
least three species of Ilex are fall and winter foods from Virginia
to Maine. The most important species by far is winterberry (see
Plate 29C ) . Plants of this genus produced over one per cent of the
winter food in the Northeast (Bi-1297); one and one-tenth per cent
in Maine, and one and three-tenths per cent in New Hampshire
( Hosley ) . Smaller quantities are recorded in all parts of the region
south to Virginia.

Jewelweed ( Impatiens ) : The seeds and leaves are an important
summer and early fall food of both chicks and adults. Spotted jewel-
weed is the more important and the only one specifically identified.

142 The Ruffed Grouse

but pale jewelweed also probably is used. It is listed as tenth in
importance as a summer food producer for both adults and young
(IV. Y. S. Cons. Dept. Ann. Rep. 1937). Darrow found that jewel-
weed products composed one per cent of the food of the chicks in
the Adirondacks, two-tenths of one per cent in the Catskills, and
one and nine-tenths per cent in the remainder of the state. In New
Hampshire, MacGregor found this food to rank second from mid-
June to mid- July (five and six-tenths per cent), second also during
August and September (five and eight-tenths per cent), then drop-
ping to a mere trace in fall.

Minnie-bush (Menziesa pilosa): The buds furnished one and
three-tenths per cent of the November-December food in Virginia
( Nelson ) .

Bishop's cap ( Mitella diphylla ) : The leaves are taken in the fall,
and to a small extent in the winter from New England to Pennsyl-
vania. Smyth found them to comprise six and two-tenths per cent of
the October food and seven and three-tenths per cent of the Novem-
ber food in the Northeast.

Bayberry (Myrica): The fruits of two species, M. carolinensis
(see Plate SOB) and M. cerifera, and tlie buds of sweet fern (M.
asplenifolia) are used during the fall and winter. M. carolinensis is
the only one of much importance to grouse. It contributed one and
two-tenths per cent of tlie fall food in New England ( Gross ) . Hosley
recorded bayberries as comprising one and two-tenths per cent of
the fall and winter food in Massachusetts, four and three-tenths per
cent in Rhode Island, and four and five-tenths per cent in Connecti-
cut. In grouse range near the seacoast they become a primary food
from Maine southward to Connecticut, and are used to a lesser ex-
tent in New York and Pennsylvania.

Black gum (Nyssa sylvatica): The fruit, leaves, and buds are
taken during fall and winter from Maine to Virginia, but not usually
in very significant amounts. In Pennsylvania, Hosley records one and
one-tenth per cent of black gum seeds; all other records show less
than one per cent.

Sorrel (Oxalis) : Leaves of at least three species of sorrel are taken
during fall, wdnter, and spring. This foliage ranked eighth in the
spring food records in the Adirondacks of New York (Darrow).
Smyth recorded it as totaling one per cent of October food in the
Northeast, and somewhat less in November. It is also recorded from

Food and Water 143

New England to Pennsylvania, including for the latter state Kuhn's
( 1941 ) record of one and three-tenths per cent for November.

Virginia creeper (Parthenocissus quinquefolia) (see Plate 30A):
The fruit is eaten during fall and winter from New Hampshire to
Virginia. In Connecticut it constituted one per cent of the fall and
winter food (Hosley).

Milkworts (Polygala): The leaves are a summer, fall, and winter
food from New York to Virginia. Darrow records these greens as
tenth among summer foods of adults in the Adirondacks of New
York and eighth (five-tenths of one per cent) in the summer diet of
chicks in the Catskills.

Smartweeds (Polygonum): Seeds of this group are taken during
summer, fall, and early winter from New England to Pennsylvania.
Darrow lists them as tenth (nine-tenths of one per cent) in the
summer foods of young birds in New York outside the mountainous
areas. MacGregor ranks them as seventh (two and one-tenth per
cent) among late-summer foods in New Hampshire,

Cinquefoils (Potentilla) : Taken from New England to Virginia,
the leaves are recorded by Smyth as making up two and two-tenths
per cent of the November food in the Northeast; MacGregor lists
them as seventh (two and eight-tenths per cent) among November
foods in New Hampshire; their use is recorded in smaller but con-
sistent amounts elsewhere by other authors.

Heal-all (Prunella vulgaris) : The leaves are taken during the fall
from New England to Virginia. They composed one and one-tenth
per cent of the November food in Pennsylvania (Kuhn), almost as
much in Virginia, and smaller amounts in New England.

Shin leaf (Pyrola): The leaves are a fall-and-vdnter food, some-
times in quantities that place them for a time among the most im-
portant foods. They amounted to six per cent of the February food
in New York and one per cent of January food (Kelso). Gross re-
cords shin leaf foliage as comprising one and two-tenths per cent of
the fall food in New England; a smaller but significant amount was
taken in Pennsylvania, and a trace in Virginia.

Buttercups (Ranunculus): The leaves are used as a summer, fall,
and winter food throughout the region. Two species have been
identified, but no doubt others are used. Smyth records buttercup
foliage as being four and eight-tenths per cent of the October food
in the Northeast, while Darrow found it amounting to one and eight-

144 The Ruffed Grouse

tenths per cent of the summer food of the young in the Adirondacks
of New York, three per cent in the Catskills, and two and six-tenths
per cent in the rest of the state. He also hsts it as fifth in importance
among the summer foods of the adult birds in the Adirondacks.

Rhododendrons ( Rhododendron ) : The leaves and buds of both
the evergreen and deciduous types of this genus are taken in fall
and winter. Such material composed one and five-tenths per cent of
the fall-and-winter food in EJiode Island (Hosley), and smaller
quantities from New Hampshire to Virginia, mainly near the coast
except in the south.

Willows ( Salix ) : The buds and leaves are eaten in fall and winter
throughout the region. In Maine they composed one and four-
tenths per cent of the fall and winter food (Hosley, 1938), while
the Bi-1297 summary for the Northeast records willow as contrib-
uting more than one per cent of the winter food. Elsewhere it has
supplied food in smaller amounts.

Elders (Sambucus) (see Plate 30C): The fruits of both north-
eastern species are utilized as summer-and-fall food. Darrow found
them to compose two and one-tenth per cent of the summer food of
yomig grouse in the Adirondacks of New York. Other records in New
England and in general northeast summaries are of very small quan-
tities, mainly because information on simimer foods and on the diet
of young birds is so scant.

False Solomon's seal ( Smilacina ) : The fruit is taken as a f all-and-
early-winter food, mainly in the northern part of the region. Smyth
recorded that it amounted to two and two-tenths per cent of the
winter food in his northeastern records and MacGregor as sixth
(two and two-tenths per cent) among October foods in New Hamp-
shire.

Bitter nightshade ( Solanum dulcamara ) : Both fruits and leaves
are used in fall and winter. Smyth fomid them to compose one and
four-tenths per cent of October food in the Northeast, probably
mostly in the north, as other records are from New York and New
England.

Mountain ash (Sorbus) (see Plate SOD): Fruit and buds are
taken of both the native and introduced species during fall and win-
ter. They compose one and one-tenth per cent of fall and winter food
in New Hampshire ( Hosley ) and are recorded in smaller quantities
from New York and other parts of New England.

Food and Water 145

Skunk cabbage {Sijmplocarpus foetidus): The seeds are eaten in
fall and winter from New England to Pennsylvania. In spring the
flowers and leaves also are taken. Darrow Usts products of this plant
as ninth in importance among winter foods in the Adirondacks region
of New York, while Kelso found them to make up over one per cent
of the winter food of New York grouse. Hosley ( 1941 ) lists skunk
cabbage as first among herbaceous foods in Connecticut and second
in Rhode Island. It has also been recorded in smaller but still sig-
nificant amounts in the fall food in New England and Pennsylvania.

False miterwort ( Tiarella ) : Leaves have been recorded as a fall
and early winter food. They are probably taken to some extent
throughout the Northeast but seem definitely most important from
Pennsylvania to Virginia. Smyth fomid them to compose three and
two-tenths per cent of October food in the Northeast, one and four-
tenths per cent in November and one and one-tenth per cent in the
winter. They ranked seventh in the list of November foods in Penn-
sylvania (Kuhn, 1940), amounting to four and seven-tenths per cent
of total food, and sixteenth in 1941 with one and seven-tenths per
cent (Kuhn); and were taken in small amounts in Virginia.

Clovers {Trifolium): The leaves of both wild white and red
clovers are taken during fall and winter, the former being the more
important. Clover is used throughout the Northeast region but is
primarily important in the North, New York and New England.
Nowhere in this region does it approach the use it receives in the
midwest states, however. Smyth found that it composed two and
four-tenths per cent of October food in the Northeast, one and four-
tenths per cent in November, and less in the winter. Gross lists it
ninth among fall foods in New England, where it amounted to three
per cent of total food. Hosley ( 1941 ) records it as first among herba-
ceous foods in Vermont and second in New Hampshire and Massa-
chusetts. MacGregor ranks it second among October foods m New
Hampshire comprising six and eight-tenths per cent of all food, and
ninth among November foods (two and three-tenths per cent).

Violets (Viola): Both the leaves and seeds are used throughout
the Northeast. They are most important as a summer food for chicks
and adults but are used to some extent in the fall. From the existing
records, the area of greatest importance of violets in grouse diet is
clearly New York, but if summer records were available from other
states, it is likely that they would prove to rank highly elsewhere.

146 The Ruffed Grouse

They are Hsted sixth among summer foods of chicks and seventh for
adults in New York (N. Y. S. Cons. Dept. Ann. Rep., 1937). As sum-
mer food for young birds, violets were ranked as ninth in the Adiron-
dacks with one and four-tenths per cent, sixth in the Catskills with
one and three-tenths per cent, and sixth in the remainder of New
York with two and one-tenth per cent; in the Adirondacks they
ranked eighth among fall foods (Darrow). MacGregor placed them
fourth am.ong late-July foods in New Hampshire, amounting to four
and three-tenths per cent of the total subsistence.

Barren strawberry {Waldsteinia fragarioides) : The leaves are
taken during late fall and winter, from New York to Virginia. Kelso
listed them eleventh among winter foods in New York, averaging
one and four-tenths per cent of total food; in January they amounted
to two per cent, and in February three per cent of the food taken.
In Pennsylvania this foliage ranked tenth among November foods
one year and constituted three and eight-tenths per cent of all food
(Kulm, 1940), then twenty-second the next year with one and two-
tenths per cent ( Kuhn, 1941 ) .

Insects : The most important groups of insects patronized for food
by the grouse are Hymenoptera ( ants, wasps, bees ) and Coleoptera
( beetles ) . These two orders are primary contributors to the food of
the young birds from hatching time to mid- July, and are of sec-
ondary importance to the adults. Some of the larger divisions (fami-
lies) of these two orders, and occasionally so small a group as a
genus, may in certain areas provide enough prey to be classed as
secondary foods. Other kinds of insects and insect allies may com-
pose over one per cent of the food of the young birds in some areas,
most commonly Lepidoptera (moths and butterflies), Opiliones
(harvestmen), Hemiptera (bugs) and Diptera (flies).

Animal foods constituted from forty-six and eight-tenths per cent
of the food of the young birds in June in the Adirondacks region of
New York to sixty-eight and three-tenths per cent in the Catskflls,
with the records from the rest of the state totaling about sixty per
cent. In July these percentages dropped to from ten to fifteen per
cent of total food for the chicks. In August the utilization of animal
life by the young birds approached the proportions used by adults,
being highest again in the Adirondacks (six and nine-tenths per
cent), practically the same in the Catskills, and only one and three-
tenths per cent over the remainder of New York ( Darrow ) .

Food and Water 147

For the adults, the animal food amounts to less than one per cent
(in New York six-tenths of one per cent) of the total food. In the
summer it exceeds the annual average, ranging from one per cent
to three per cent in New York, and highest in the Adirondacks ( Dar-
row). Occasionally insects will exceed one per cent of the food in
spring or fall too.

Smyth examined four May-June chick stomachs and found sLxty-
six and nine-tenths per cent of the food to be insects and their
relatives; in three July stomachs the proportion was only eleven
per cent. One August stomach contained ten per cent animal food.
Judd ( 1905 ) reported on four chicks less than one-quarter grovm,
the stomach contents of which were found to be ninety-five per cent
insects, while seven adults taken at the same season contained only
thirty per cent of insects.

Judd's adult records are very notable in that he found insects and
related animal life to constitute ten and nine-tenths per cent of total
food (including Coleoptera— four and six-tenths per cent; Lepidop-
tera larvae one and two-tenths per cent; Orthoptera eight-tenths of
one per cent ) . This is the more unusual because a great majority of
the birds were taken during the cold months. Possibly the grouse
were more insectivorous formerly, but this does not seem likely.
Other fall and winter records are in marked contrast, insects ranging
from one and eight-tenths per cent of the October food (Smyth)
and one and four-tenths per cent of fall food in New England ( Gross )
to a mere trace of the winter food in New York (Kelso). Kuhn
(1940) lists animal food as amomiting to thirteen-hundredths per
cent of the November food in Pennsylvania. Nelson found only
seven-hundredths per cent in November-December stomachs in
Virginia. Smyth gave the November insect diet as totaling eight-
tenths per cent of food taken, but he found none at all in stomachs
collected from December through March.

Occasional Foods. The kinds of plant and animal foods acceptable
to the ruflPed grouse are without doubt more numerous than pres-
ently recorded. The New York investigation is continuing its study
of the grouse food habits and finds new food items with each new
season's collections. The list of foods now known to be taken will
be considerably enlarged with more study, but the importance of the
newly discovered items is likely to be slight (see Tables 5 and 6).

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Table 6
Animal Foods Eaten by the Ruffed Grouse in the Northeast

Scientific and Common Names

Eaten by

References

Phylum Arthropoda
Class Hexapoda
Order Orthoptera
Gryllidae (Crickets)
Gryllus assimilis
(sp.)
Acrididae (Shorthomed grasshoppers)
Melanoplus femur-rubrum
Unidentified
Locustidae ( Longhorned grasshoppers )
Xiphidiujn (sp.)
Unidentified
Phasmidae (Walking sticks)
Diapheromera femorata

Order Neuroptera
Chrysopidae (Lace wing flies)
Chrysopa (sp.)

Adult
Adult

Adult

Adult and young

Adult

Adult and young

Adult

Young

Gross
Judd

Judd

Smyth

Judd
Judd

Smyth

N. Y. S. Cons.
Dept. Ann.

Order Corrodentia

Rep.

Psocidae (Psocids)

Unidentified

Adult

Smyth

Order Homoptera

Membracidae (Tree hoppers)

Unidentified

Adult and young

Judd

Cicadellidae (Leaf hoppers)

Adult and young

Smyth, Kuhn

Aphididae (Aphids)

Unidentified

Adult and young

Smyth

Order Hemiptera

Miridae (Leaf -bugs)

Unidentified

Adult and young

Reduviidae ( Assassin-bugs )

Zelus (sp.)

Adult

Kelso

Unidentified

Adult and young

Judd

Unidentified (Nymph)

Adult

Smyth

Phymatidae (Ambush-bugs)

Vhijmata (sp.)

Adult

Judd

Lygaeidae (Chinch-bugs)

Drymus cra^sus

Adult

Kelso

Blissus leucopterus

Adult

Judd

Eremocoris ferus

Adult

Gross

Unidentified

Young

N. Y. S. Cons,
Dept. Ann.
Rep.

162

Table 6 (Continued)
Animal Foods Eaten by the Ruffed Grouse in the Northeast

Scientific and Common Names Eaten by References

Aradidae (Flat-bugs)

Unidentified Young N. Y. S. Cons.

Dept. Ann.
Rep.
Pentatomidae (Stink-bugs)

Unidentified Young N. Y. S. Gons.

Dept. Ann.
Rep.
Order Goleoptera
Gicindelidae (Tiger-beetles)

Cicindela punctidata Adult Smyth

Garabidae (Ground-beetles)
SphaerodcTus ( sp. )
Harpalus (sp.)
Bothriopterus (sp.)
Anisodactijlus (sp.)
Pterostichus (sp.)
Chlaenhis (sp.)
Unidentified
Staphylinidae ( Rove-beetles )
Unidentified

Lampyridae (Fireflies)

Lucidota corrusca

L. nigricans

(sp.)

Unidentified
Gantharidae (Soldier-beetles)

Cantharis (sp.)

Unidentified

Elateridae ( Glick-beetles )

Unidentified
Helodidae

Unidentified

Buprestidae (Metallic wood-borers)

Unidentified
Ostomatidae

Calitys scabra
Goccinellidae ( Lady-bugs )

Chilocorus bivulucrus

Unidentified

163

Adult Gross

Adult Judd

Adult Gross

Adult Judd

Adult Judd

Young

Adult and young Judd

Young N. Y. S.
Dept.
Rep.

Gons.
Ann.

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Adult and young Smyth
Adult Gross
Adult and young Judd

Adult Gross
Young N. Y. S.

Dept.

Rep.

Gons.
Ann.

Adult and young Judd

Young N. Y. S.
Dept.
Rep.

Gons,

Ann.

Adult Judd

Adult Smyth

Adult Gross
Adult Kuhn

Table 6 (Continued)
Animal Foods Eaten by the Ruffed Grouse in the Northeast

Scientific and Common Names

Eaten by

References

Lagriidae

Unidentified

Young

N. Y. S. Cons.
Dept. Ann.
Rep.

Anobiidae

Unidentified

Young

N. Y. S. Cons.
Dept. Ann.
Rep.

Scarabaeidae (Lamellicorn beetles)

Aphodius (sp.)

AduJt

Kelso

Dichclonycha (sp.)

Adult

Judd

Lachnosterna hirsuta

Adult

Judd

(sp.) (larva)

Young

Judd

Unidentified

Young

F. C. Edminster

Cerambycidae (Longhorned beetles)

. Hapalosalia vibex

Adult and young

Smyth

Chry somelidae ( Leaf -beetles )

Diahrotica vittata

Adult

Smyth

Leptinotarsa decemlineata

Adult

Judd

PJwcdon armoraciae

Adult and young

Smyth

Lina tremulae

Adult

Kuhn

Haltica chalybea

Adult

MacVicar '18

Donacia (sp.)

Adult

Smyth

GaJerucella sexvittata

Adult and voung

Smyth

G. cavicollis

Adult

Gross

G. sagittariae

Adult

Judd

Microrhopala (sp.)

Young

Calligrapha (sp.)

Adult

Gross

Adoxus vitis

Adult

Judd

Chaetocnema (sp.)

Adult

Judd

Systena hudsonias

Adult

Judd

S. blanda

Adult

Judd

Disonycha caroliniana

Adult

Judd

Unidentified

Adult and young

Judd

Curculionidae (Snout-beedes)

Sitones hispidulus

Adult

Judd

Unidentified

Adult and young

Smyth, N. Y. S.
Cons. Dept.
Ann. Rep.

Order Mecoptera

Unidentified

Young

N. Y. S. Cx)ns.
Dept. Ann.
Rep.

Order Trichoptera

Phryganeidae

Unidentified

Adult and young

Gross, N. Y. S.
Cons. Dept.
Ann Rep.

164

Table 6 {Continued)
Animal Foods Eaten by the Ruffed Grouse in the Northeast

Scientific and Common Names

Eaten hij

References

Order Lepidoptera
Olithreutidae

Carpocapsa pomonella (codlin moth)

Carpocapsa larva
Geometridae

Unidentified larva
Notodontidae

Symmerista alhijrons

Schiztira concinna
Noctuidae (owlet-moths)

Alabama argilhcea

Cirphis unipuncta

Noctua (sp.)

Unidentified larva

Unidentified larva
Unidentified eggs

Order Diptera
Tipulidae ( Crane-flies )

Unidentified
Culicidae ( Mosquitoes )

Unidentified
Mycetophilidae ( Fungus-gnats )

Mycetophila (sp.)
Bibionidae (Marsh-flies)

Bibio (sp.)

Dilophus (sp.)
Leptidae (Snipe-flies)

Wiagio mystacea

Unidentified
Asilidae (Robber-flies)

Unidentified

Empididae (Dance-flies)
Unidentified

Syrphidae (Syrphus-flies)
Unidentified

Muscidae

Unidentified pupa
Unidentified larva
Unidentified pupae

Adult

Adult

Adult
Adult

Adult

Adult

Adult and young

Adult and young

Adult and young
Adult and young

Adult and young

Adult

Adult and young

Adult
Adult

Adult and \-oung
Adult

Adult and young

Young

Adult and young

Adult and young

Adult

Young

Smyth

Smyth

Judd
Judd

Judd

Judd
Judd
Smyth, N. Y. S.

Cons. Dept.

Ann. Rep.
Smyth
Smyth

Judd, Smyth

Smyth

Smyth

Gross
Gross

Smyth
Smyth

Kelso, N. Y. S.
Cons. Dept.
Ann. Rep.

N. Y. S. Cons.
Dept. Ann.
Rep.

Kuhn, F. C. Ed-
minster

Smyth
Smvth
Smyth. N. Y. S.

Cons. Dept.

Ann. Rep.

165

Table 6 (Continued)
Animal Foods Eaten by the Ruffed Grouse in the Northeast

Scientific and Common Names

Eaten hy

References

Order Hymenoptera
Tenthredinidae ( Sawflies )

Neodiprion (sp.) larva

Nematus (sp.)

Unidentified
Braconidae ( Braconids )

Unidentified

Ichneumonidae ( Ichneumon-flies )

Ambly teles (sp.)

Unidentified
Serphidae

Unidentified

Pelecinidae

Peleeinus polyturator

P. polyturator larva

Cynipidae (Gall-wasp)

Amphibolips (sp.)

Unidentified
Chalcididae (Chalcid-flies)

Unidentified

Formicidae (Ants)
Formicina (sp.)
Formica (sp.)

Camponotus pennsylvanicus
C. herculeanus

(sp.)
Tetramorium caespitum
Unidentified
Vespidae (Wasps)
Unidentified

Sphecidae
Unidentified

Andrenidae
Unidentified

Adult
Adult
Young

Young

Judd
Judd
Smyth

N. Y. S. Cons.
Dept. Ann.
Rep.

Adult Kelso

Adult and young Smyth

Young

Adult
Adult

Adult
Adult

Yoimg

N. Y. S. Cons.
Dept. Ann.
Rep.

Judd, Gross
Judd

Judd
Judd

N. Y. S. Cons.
Dept. Ann.
Rep.

Adult Gross

Adult and young Gross, F. C. Ed-
minster
Adult and young Judd
Adult Gross

Adult Gross

Adult and young Judd
Adult and young Smyth

Young

N. Y. S. Cons.
Dept. Ann.
Rep.

Adult and young N. Y. S. Cons.
Dept. Ann.
Rep.

Young

166

N. Y. S. Cons.
Dept. Ann.
Rep.

Table 6 (Continued)
Animal Foods Eaten by the Ruffed Grouse in the Northeast

Scientific and Common Names

Eaten

hy

References

Tiphiidae

Unidentified

Young

N. Y. S. Cons.

Dept. Ann.

Rep.
F. C. Edminster

Unidentified cocoon

Young

Class Arachnida

Order Araneae (Spiders)

Lycosidae (Wolf spiders)

Unidentified

Adult

Judd

Attidae (Jumping spiders)

Unidentified

Adult and

young

Judd

Unidentified

Adult

Gross

Order Phalangida (Harvestmen)
Phalangiidae

Unidentified larva

Class Myriopoda
Order Diplopoda (Millipedes)
Julus (sp.)
Unidentified

Phylum Mollusca
Class Gastropoda
Order Pulmonata (Snails, slugs)
Helix (sp.)

Limax (sp.)
Tebennophorus carolinensis

Phylum Chordata
Class Reptilia
Suborder Serpentes (Snakes)
Colubridae
Unidentified

Adult and young Judd

Adult and young Smyth
Adult Judd

Adult and young Judd, N. Y. S,
Cons. Dept.
Ann. Rep.

Adult Judd

Adult

Adult

Smyth

107

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Food and Water 171

References listed for animal food records are cited fully in the list at
the end of this chapter. "N. Y. S. Cons. Dept. Ann. Rep." refers to
Annual Reports of the New York State Conservation Department.
"F. C. Edminster" indicates records obtained by the author: the
identifications for these are by John C. Jones.

I am indebted to Earl C. Murdoch, formerly of the U. S. Soil
Conservation Service for assistance in checking insect names and in
arranging them in order, and to Dr. J. C. Bradley of Cornell Uni-
versity for checking names found in references.

It will be noted that in genera having records of definite species
eaten, there is often also a listing of the genus "sp." This may in
some cases result in duplication since the material identified only
to genus by one author may be the same as one fully identified by
another. Since such duplication is not definite, it was determined
best to list them both when this situation arose.

The above summary of ruffed grouse food-habit records ( Table 7 )
covers 321 specimens received by Dr. A. A. Allen in connection
with his disease studies between 1925 and 1930. The analyses were
largely made by his students. Only incidence of occurrence of each
type of food is available since volumetric measurements were not
made. This material has not heretofore been published.

QUALITY OF DIFFERENT GROUSE FOODS

Wild-life foods have been variously classified according to their
availability, palatability, and physiological effects ( Leopold, 1933 ) .
Terms such as "preferred," "staple," "incidental," "emergency,"
"stuffing," and "salad" have been used to distinguish and describe
them. The first three of these adjectives apply to many grouse foods
according to their quality.

A preferred food is one that is selected by the bird when other
usable items are available. It stands among the highest ranks of
palatable foods.

A staple food is one that furnishes a large proportion of the bird's
food in a given season, and probably is highly palatable. However,
it may be lower in the scale of palatability than the preferred foods.

Incidental foods are those taken normally in small quantities and
quite variably in different localities. They either rank below staple
foods in the palatability scale or are less available.

172 The Ruffed Grouse

Since the grouse is a browser, it is doubtful that it ever has to make
a choice of an emergency food. If any foods are accessible, some of
its regular articles of diet will be among them. No evidence of the
consumption of "stuflBng" or "salad" foods was obtained. As a mat-
ter of fact, it is difficult in many cases to distinguish between pre-
ferred and staple foods of the grouse.

Some grouse foods, however, are clearly preferred. Beechnuts, red
haws, apple fruit or foliage, acorns, strawberry fruits and leaves,
bramble fruits, greenbrier berries, and grapes are so utilized, and
birch, apple, fern, popple, cherry, and blueberry are choice among
the leaf and bud foods. Certainly in a sense, birch buds are preferred,
as they are regularly taken in preference to many other equally
available foods of this type, as those from beech, maple, elm, ash,
and hickory, for example. Yet, it is likely that if beechnuts, hawthorn
fruits and certain other foods were available in winter much less of
birch buds would be taken.

Preference must be interpreted with due regard to availability.
It is likely that chestnuts would be a preferred food if available,
since they probably once were of primary rank. Being absent now
over most of the range, they rank merely as an incidental food.

Most of the foods listed above as primary and secondary may also
be considered staple, while those not so classed are ordinarily in-
cidental.

VARIATION IN SUMMER FOOD HABITS OF YOUNG AND ADULTS

The groups of plants and animals providing the bulk of the sum-
mer food are essentially the same for both adults and young. The
quantities taken by birds of these age classes, however, may vary
widely. Young birds in early summer consume a very high propor-
tion of animal food whereas the adults use only a moderate quantity.
The two topmost food-producing groups of plants, brambles ( Rubus )
and sedges (Carex) are the same throughout New York for the
chicks. Rubus ranks first for adults, but one or the other of Carex,
cherries ( Prunus ) and popples ( Populus ) takes second place in dif-
ferent localities. The volume of raspberries used by chicks ( twenty-
nine per cent-thirty-eight per cent summer average ) greatly exceeds
that taken by adults. There are numerous other variations in summer
food habits between the adults and young, mostly, however, of minor
importance.

Food and Water

173

RELATION OF FOOD SUPPLY TO COVER TYPES

There is great variation in the composition of the major cover
types including the species that furnish food for the grouse. Some
types are more satisfactory than others from the standpoint of food,
even that suppHed by the same species. For example, plants as a
rule produce more fruit when in full sunlight than when partly
shaded. Thus grapevines are more productive when in slashings
or other brushy areas than in a woodland. Further, the fruit that is
produced inside a woodland is mostly at the top of the vines up in
the crown where it is rather inaccessible for grouse.

Some vegetational types are more valuable for the food they pro-
duce at one season, than at another, as an example, the pure popple
subtype which has its greatest value in winter, for the buds, and
again for a time in summer, for the catkins. The value of the several
cover types as food or shelter producers is of prime importance when
considering the need for interspersion of types. The perfect mingling
would result in the best feeding and roosting coverts being adjacent
for each of the seasons.

The major relations of food supply to cover types may be summed
up as follows :

Cover type
Open land

Overgrown land

Slashings

Hardwood woodland

Mixed woods

Coniferous

Season

Relative food-producing value

Summer and fall
Winter and spring
Fall and summer
Spring and winter
Summer

Fall, spring and winter
Winter and spring
Summer and fall
Winter and spring
Summer and fall
All year

High

Low

Very high
Medium to high
Very high
Medium

High to very high
Low to medium
Medium to high
Low to medium
Low to medium

As to cover types in which various foods are produced, we may
make the following generalizations, the cover types being listed
about in the order of their productiveness:

Fruits— overgrown land, slashings

Nuts— hardwood woodland, mixed woods

Seeds— overgrown land, open land, slashings

Buds— hardwood woodland, mixed woods, overgrown land

Leaves— hardwood woodland, mixed woods, overgrown land, slashings

Insects— slashings, overgrown land, open land

174 The Ruffed Grouse

DISTRIBUTION OF BIRDS AS AFFECTED BY FOOD SUPPLY

The distribution of grouse is affected by the food supply both
daily and seasonally. Each morning, weather permitting, the birds
leave their roost, selected for its shelter value, and seek the food-
producing covert most readily available. This may be right w^here
they roosted, but more likely it is a little distance away. Again at
evening the feeding coverts are sought, after which the biids retire
to the roosting place. Thus the proper location of feeding coverts
with respect to shelter is of immense importance in determining the
carrying capacity of an area, since both must occur within the daily
cruising radius of the birds and must be suitable for all seasons of
the year.

In a more extensive manner the location of certain types of food-
producing coverts affects the seasonal distribution. Overgrown land,
particularly that with preferred foods such as the fruits of hawthorn
or dogwood, markedly affect the range of grouse in the fall. Slash-
ings and some types of overgrown land draw many grouse during
the summer, while in the spring the hardwood woodlands probably
have their greatest effect on distribution as affected by food habits.
Food supply has least effect in winter, when widely available buds
are the dominant food and shelter the prime need for existence.

EFFECTS OF CULTURAL OPERATIONS ON THE
FOOD SUPPLY

It is well recognized that the workings of man may have profomid
effects on grouse range. What are the effects on the food supply of
the birds? Let us consider this question for each of the important
cultural practices.

Lumbering: Woodland cutting practices change the food condi-
tions of a woodland in proportion to their severity. In some respects
food supplies for grouse may be increased, in others, decreased. The
cutting of a growth of mast-producing trees, as oak and beech, re-
moves it from production for decades. This is probably the most
serious destructive effect of lumbering on the food of the grouse
and is most injurious in the southern part of the range. In the same
way, lumbering takes some of the important bud-supplying species
as cherry, poplar, and birch, out of production for several years. A

Food and Water 175

by-product of lumbering is the common destruction of the so-called
weed species, which often include important food-producing trees,
including hornbeam, beech, and hophornbeam. In a table classify-
ing thirty-seven New York tree species of upland woodlots accord-
ing to usefulness ( Cope, 1933 ) , three important grouse food-produc-
ing trees are among the fourteen valuable species (red and white
oak and black cherry), eight are among the thirteen intermediate
species, and eight of ten inferior species are important grouse food
plants. A similar list classifying forty-six trees of West Virginia
". . . according to their cash value" includes but three important
grouse food-producers (the same three as above) among eighteen
valuable species. The sixteen inferior species included six important
grouse food plants while the twelve weed species had nine ( Smith &
Byers, 1941).

Fortunately, woodland cutting often results in more improvement
than injury. Shrubs are released so that they produce larger crops of
fruit. Brambles, pin cherry, quaking aspen, and other desirable
woody plants germinate in the new openings; the herbaceous food
plants are usually greatly increased. Insect food also is much more
plentiful in slashings and woodland openings than in the woodland
proper.

It is clear then that taking out wood products may readjust the
grouse food situation in many respects. Since the changes depend
upon the type of cutting operation and upon the conditions existing
before cutting, it is apparent that regulation of woodland cuttings
may be made an important method of improving food conditions for
grouse.

Burning: Woods fires initially destroy grouse food. However, the
plant succession resulting from a fire often brings in a more desirable
food supply than formerly existed, or at least adds variety. A light
ground fire temporarily removes most of the herbaceous ground
cover and some of the shrubby understory. The plants that then
spring up are likely to be somewhat but not strikingly different from
those destroyed. A crown fire, or a very hot understory fire, however,
will create an entirely different cover type which at first is practically
useless for grouse. In a few years, however, it may become an im-
portant food-producing area serving the purpose of a slashing. The
vegetation on these bums is often dominated by pin cherry, poplar,
brambles, and blueberry. . .

l^lkC<h-

176 The Ruffed Grouse

Fencing and Livestock: Woodlands adjacent to pastures present a
management problem to a farmer. The desirability of fencing the
woodlands so that the livestock may not have access to them is gain-
ing increasing recognition. It is generally true in the Northeast that
one cannot have a good pasture and a good woodland on the same
ground. Likewise the pasturing of woodland has a detrimental ef-
fect on the area as grouse cover.

The most obvious effect of pasturing a woodland is the reduction
of the understory and ground cover. That materially affects the sup-
ply of grouse food and often entirely eliminates important food-
producing herbs and shrubs. The changes that grazing brings about
in soil conditions may affect the plant succession on the area for a
long time following abandonment of grazing; the effect is to set
back the ecological stage toward the field weed complex. If grazing
is continued intensively it eventually results in the transformation of
the woodland into a park-like area, and ultimately into an open
field.

Ploughing and Cultivating: Since the use of the plough and other
crop field tools are normally confined to the better-quality open
fields, these practices do not materially affect grouse in a direct
sense. It is true that a cultivated field is less desirable to grouse than
a weedy meadow, but their use as cover is too small to be important.
As openings to provide edges, one field type is as good as another.

From the long-time point of view, the maintenance of land in
field crops prevents it from becoming grouse cover and is a restric-
tion on the increase of available grouse foods as well as of shelter.

FOOD AS A LIMITING FACTOR

There are few species of game birds that have as little trouble
finding food year in and year out as the ruffed grouse. If a covert is
of a nature to permit grouse existence from the standpoint of size and
shelter, it is likely to provide enough food for the maintenance of at
least some grouse. This does not in any way imply that food is not
an important factor. It is. But the grouse can get along on such a
wide variety of foods. It uses parts of plants that are almost always
available, regardless of weather conditions or competition of other
animals, that ahnost any type of woody area will provide some food.

Food and Water 177

This may not be true everywhere in the range of the niJBFed grouse
but is generally true in the Northeast.

That the supply of food is a possible limiting factor in areas that
are well stocked with grouse is conceivable although not likely. Let
us consider this prospect under some of the most likely conditions.

Food Shortages: A shortage of food for adults at any time of the
year except winter and early spring, or for chicks except during
the first six weeks of life is altogether unlikely. So far as fruits, seeds,
and succulent leaves are concerned, a drastic food shortage in late
Mdnter and early spring is conceivable. A lack of buds of hardwood
trees and shrubs, however, is out of the question. As grouse are well
able to thrive for considerable periods on a diet of buds, we can rule
out shortage of foods for the adult as a limiting factor on the usual
grouse range.

A shortage of insects in June or early July, at which time they are a
vitally necessary food for the young bird, would be disastrous. Nor-
mally, grouse coverts contain from one hundred to five hundred
thousand insects and other arthropods per acre at this time of year
in the zone from the loose ground litter to a height of one foot above
ground, the area of availability to young grouse (estimates from
measurements taken on Connecticut Hill ) . Variation in normal years
depends upon the cover type and the weather. While we have not
measured the insect populations in severely abnormal seasons, we do
know enough about the effect upon them of exceedingly cold and
wet weather to surmise the possible consequences to young grouse.
Shortage of animal food during seasons of cold, or cold and wet
from mid-May to the end of June is one of the likely contributing
causes of excessive juvenile mortality that often precede years of
greatly decreased grouse numbers ( see effects of weather on grouse.
Chapter VI ) . The question as to whether these losses in young birds
occur primarily as a result of food shortage or directly from the cold
is not clear. Surely the two factors sometimes combine to take a
heavy toll.

Availability during Emergencies: We have already pointed out
that foods provided by low-growing vegetation, mainly leaves, fruits,
and seeds, may be completely covered with snow for considerable
periods in the winter. These, then, are relatively inaccessible al-
though the birds are surprisingly able in seeking them out. But tree

178 The Ruffed Grouse

buds are always available above the snow, and the grouse turn
largely to them for food when snow is deep.

What then, if sleet or freezing rain puts a coating of ice upon all
vegetation? This condition probably renders completely unavailable
foods under the snow. It may make budding difficult but probably
does not entirely prevent it. Even if the ice coating is complete
enough and thick enough to prevent feeding by grouse, it seldom
lasts long enough to cause starvation of the birds. It is normal for
grouse to refrain from eating for a day or two during bad storms
anyway. The fact that not a single normally healthy^ grouse has
been known to die of starvation is proof enough of the ability of the
species to forage satisfactorily under any winter conditions that
have been observed.

If cold or wet early summer weather may be considered an emer-
gency, then we probably have one type of food emergency that
young grouse are sometimes unable to meet. This condition has
just been discussed.

Competition of Other Species for Food: In considering the major
grouse food plants, notes were made concerning competition for
these foods with other forms of wild life. Most of the preferred foods
are also taken by other species in considerable amounts, but in only
a few cases does this noticeably affect the grouse. Even in these
few cases," only the local distribution of the birds is modified. Rarely,
if ever, is the number of grouse over a large area materially affected
by food competition.

Maintenance of Grouse Health: Does the availability of certain
foods affect the health of grouse? More particularly, does the relative
lack of fruits and green leaves in late winter in the northern range
and the resulting dependence on a bud diet so affect the bodily
health of the birds that their breeding-season vigor is impaired? If
that were true, would it result in weakened young birds the fol-
lowing spring or summer? Does the quality of certain types of im-
portant foods used in winter and early spring, such as buds, vary
in different years in a manner that affects the health of the birds in
the spring, resulting in weakened offspring? Conceivably the chemi-
cal content of tree leaf buds, possibly in regard to the representa-

^ We have observed cases of stanation in birds with injuries to their digestive
system.

2 Swanson (1942) notes that grouse are never abundant in an overbrowsed deer
yard due to the destruction by deer of important grouse food plants.

Food and Water 179

tion of some important vitamin, might be affected by changes in the
character of sunHght reaching the earth in different years. And that
change might be brought about by the cycles in sunspots.

These questions and others of similar import have been seriously
asked in recent years. Sunspot cycles have been clearly recognized,
and some correlations with fluctuations in life on the earth seem to
be satisfactorily established. It followed, therefore, that the records
of grouse cycles would be compared with the sun phenomenon. The
correspondence is not conclusive, although there appears to be
enough relationship to maintain one's further interest.

The evidence at hand refutes the probability of any serious re-
lationship between quality of food and high mortality among giouse.
However, the matter is far from settled, and the opportunity for
direct testing of the possible relationships is practically virgin. It
seems probable that if a deficiency in winter food quality resulted
in weakling chicks, it would show a definite effect in one or more of
the leading factors in reproduction: size of clutch, degree of fer-
tility, proportion of females nesting, or percentage of loss of em-
bryos during incubation. None of these, however, show any connec-
tion with excessive infant mortality that we can discern. We must
conclude tentatively, therefore, that the hypothesis of a major con-
nection between food quality and grouse health is unlikely to be
valid.

ECONOMIC ASPECTS OF GROUSE FOOD HABITS

Early investigators rated the ruffed grouse an important economic
asset to the farmer from the standpoint of its insect consumption.
According to Judd (1905) "Bugs . . . are much more often de-
stroyed by bobwhite and the ruffed grouse than by other birds. The
ruffed grouse has been known to prey on the chinch bug, which at
times is the most injurious insect in our country, and seldom de-
stroyed by any gallinaceous birds. Farmers who permit market
hunters to rob them of their game should remember this point."
This inclination to point up the insect-consuming habits of grouse
as a benefit to agriculture was natural but the conclusion reached
has not withstood the test of further study. In the first place, the
grouse are dominantly insectivorous only as chicks, and then only
for a few weeks. Secondly, the insects are taken primarily from

180 The Ruffed Grouse

woodland coverts, not from crop fields where agricultural benefits
might be involved. Thirdly, while many of the insects taken are of
injurious types, many others are beneficial, or neutral in economic
importance. Lastly, grouse are never abundant enough near farm
fields to be a real factor in controlling insects under any conditions.
And so we must conclude that the direct economic aspects of grouse
insect eating are practically negligible. The grouse may be con-
sidered, however, as part of the combined force of all insect-eating
birds that tends to lower the number of all insects— doubtless a de-
sirable accomplishment.

Another phase of grouse food habits has proved to be significantly
destructive to agriculture in some localities. The habit of budding
apple trees sometimes results in sufiBcient damage to the trees to im-
pair their productivity. In Massachusetts and New Hampshire this
habit has resulted in bounty payments and state-paid damage claims
in years past. In recent years the damage claims paid by New Hamp-
shire due to grouse budding apple trees has varied from about
$26,000 in 1926 to a low of $12.57 in 1938. The highest amount for
the past decade was $3,488.07 in 1935 on seventy claims.' It is in-
teresting to note that these claims are based on a twofold damage:
to the succeeding apple crop, and to the tree itself. Grouse have
been known to take as high as ninety per cent of the buds from
individual trees in a single season. Fortunately this damage does
not occur commonly in commercial orchards.

In other respects the food habits of the grouse are of litde or no
significance to man's interests. We may, therefore, summarize their
economic importance as generally neutral.

USE OF GRIT

The grouse is one of the type of birds that have a strong, muscular
gizzard that crushes the food after receiving it from the crop. In
order that the gizzard walls may perform this function satisfactorily,
hard particles must be present in the gizzard to act as a grinding
agent. Ordinarily, small stones are swallowed deliberately by the
birds for this purpose. Occasionally hard seeds assist in the crushing
of other foods. Red haw (Crataegus) seeds are notable in this con-

^ Records kindly made available by the New Hampshire Fish and Game Depart-
ment.

Food and Water 181

nection and they are often retained in the gizzard for considerable
lengths of time, as evidenced by their worn condition.

In Smyth's records for October, the gizzards averaged thirty-four
particles of grit each which composed nine and three-tenths per
cent of the gizzard contents. The extremes ranged from several hav-
ing no grit at all to one having three hundred and ten bits of gravel
(thirty-two per cent of contents). Other highs were two hundred
and ninety-seven ( thirty-six per cent ) and one hundred and seventy-
seven (forty-five per cent) stones. Two having no stones contained
worn Crataegus seeds.

In Smyth's November records, the gizzards averaged thirty-seven
bits of grit, which were nine and nine-tenths per cent of the con-
tents. Eleven had quartz stones and four had worn Crataegus seeds
but no gravel. His winter records showed an average of twenty-
seven stones each (seven and three-tenths per cent of gizzard con-
tents ) . In six spring records, the gizzards averaged fifty-eight stones
each, fourteen per cent of the total contents.

Kuhn ( 1940 ) found gravel in only nine crops out of two hundred
and thirty examined, but he did not report on the gizzard contents.
This indicates the slow rate of ingestion of gravel needed to main-
tain the required quantity in the gizzard. Similarly, Nelson et al.
(1938) found only eight crops out of one hundred and eighty-four
to contain gravel, while of one hundred and seven gizzards from
the same specimens, seventy-six contained gravel averaging eight-
tenths cubic centimeters or one-third teaspoonful each. Of those
not containing gravel, all had hard seeds showing wear, including
those of rose, greenbrier, sumac, dogwood, witch hazel, and black
gum.

Kelso's ( 1935 ) report gives the winter average of grit in the giz-
zards as seven and nine-tenths per cent in New York, ranging from
four and eight-tenths per cent in March to nine and eight-tenths
per cent in January. There is clearly a considerable variation in need
for grit at different times of the year, depending upon the nature of
the foods being eaten. Probably buds and twigs demand the most
grit while soft insects require the least. Hard seeds, if digested, also
would require a lot of grinding, while soft fruits would need little.
We can, therefore, generalize that grit is most needed in winter,
when hardest to get, and least required during the summer.

182 The Ruffed Grouse

WATER REQUIREMENTS OF GROUSE

The ruffed grouse, like all warm-blooded animals, has an ingestion
requirement of moisture. Unlike many birds, the ruffed grouse meets
this need easily through a variety of sources. Open water, snow,
dew, soft fruits, succulent vegetation, and juicy insects all contribute
to satisfy the need for water.

It has been suggested that serious summer drought may impair
the survival of the young grouse due to lack of available water.
However, the evidence indicates that the chicks are as adaptable in
getting their water requirement as are the adults. In fact both chicks
and adults are so independent of open water, that their summer
distribution is not affected materially by the location of water
courses, swamps, or springs. The occasions when the birds are found
near surface water ordinarily result from food or shelter availability
rather than from the seeking of water for drinking. Many grouse
broods and adults have been observed to spend the entire period of
an abnormally dry summer in coverts completely lacking surface
water. In some cases there was even a complete absence of dew for
days at a time.

REFERENCES AND CITATIONS ON RUFFED
GROUSE FOOD HABITS

Anon. Winter Food of the Ruffed Grouse in the Northeast, U. S. Biol. Survey,

mimeo. leaflet Bi-1297, Oct., 1933.
Bartlett, M. L. Ruffed Grouse and Fruit Trees, Bull. Am. Game Prot. Assn.,

Vol. 13, No. 1, 1924.
Bendire, Chas. Life Histories of North American Game Birds, U. S. Nat. Mus.

Bull. No. 1, 1892.
Brooks, Maurice. Ilex collina fruits as bird food, Wilson Bull., Vol. 55, No. 4,

Dec, 1943.
Cope, J. A. Growing Wood as a Crop on New York Farms-Part 1, Trees and

Products, Bull. 270, Cornell Ext. Bull., N. Y. State Coll. of Agr., Nov., 1933.
Darrow, R. W. Seasonal Food Preferences of Adult and of Young Ruffed

Grouse in New York State, Trans. 4th N. A. Wildlife Conf., 1939.
Davison, V. E., and Van Dersal, W. R. Broomsedge as a Food for Wildlife,

Journal WildHfe Mgt., Vol. 5, No. 2, AprU, 1941.
GilfiUan, M. C., and Bezdek, H. Winter Foods of the Ruffed Grouse in Ohio,

Joum'. Wildlife Mgt., Vol. 8, No. 3, July, 1944.
Graham, E. H. Legumes for Erosion Control and WildUfe, U.S.D.A. Misc.

Pub. No. 412, 1941.

Food and Water 183

Gross, A. O. Food of the Ruffed Grouse, Game Breeder & Sportsman, Vol. XLI,

No. 8, Aug. 1937.
Hosley, N. W. Woody Plants Used by Wildlife in the Northeastern United

States, Ph.D. thesis, Univ. of Michigan, 1938.
Hosley, N. W. The More Important Fall and Winter Foods of the Ruffed

Grouse in the New England States, mimeo., 1941.
Judd, Sylvester D. The Grouse and Wild Turkeys of the United States and

Their Economic Value, U. S. Biol. Survey, Bui. No. 24, 1905.
Kelso, L. H. Winter Foods of Ruffed Grouse in New York, U. S. Biol. Survey

mimeo. leaflet BS— 1, Jan., 1935.
Kuhn, T. M. Fall Foods of the Ruffed Grouse in Pennsylvania, Pa. Game News,

Vol. XI, No. 10, Jan., 1940.
Kuhn, T. M. November Foods of the Ruffed Grouse in Pennsylvania, Pa. Game

News, May, 1941.
Leopold, Aldo. Game Management, 1933.
Loomis, Everett M. Summer Birds of Pickens County, South Carolina, Auk,

Vol. 7, p. 36, 1890.
MacGregor, A. E. Summary of Grouse Crop and Gizzard Analyses for 1940,

N. H. Fish & Game Dept., 1940.
MacVicar, Donald. Practical Grouse Preserving, Bull. Am. Game Prot. Assn.,

Vol. 7, No. 2, 1918.
McAtee, W. L., and Beal, F. S. L. Some Common Game Aquatic and Rapacious

Birds in Relation to Man. U.S.D.A. Farmers' Bull. No. 497, 1922.
Merritts, Helen V. What Grouse Eat During the Hunting Season, Penna.

Game News, Feb., 1943.
Nelson, A, L., Clark, T. E., and Bailey, W. W. Early Winter Food of Ruffed

Grouse on the George Washington National Forest, U. S. D. A. Circular

No. 504, Dec. 1938.
Nuttall, Thomas. Birds of the United States and Canada, p. 31, 1903.
Smith, R. B. and Byers, J. Planting Forest Trees, Circular 330, West Vir-
ginia Ext. Service, March, 1941.
Smyth, Thomas. A Study of the Food and Feeding Habits of the Ruffed

Grouse, M.S. thesis, Cornell University, 1923.
Swanson, Gus A. Note on Relationships of Ruffed Grouse to White Tail Deer,

Pittman-Robertson Quarterly, Vol. 2, No. 1, Jan., 1942.
VanDersal, W. R. Native Woody Plants of the United States, U. S. D. A. Misc.

Pub. 303, 1938.
Warren, B. H. Birds of Pennsylvania, p. 108, 1890.
Weed, C. M. and Dearborn, N. Birds in Their Relation to Man, 1903.
Wright, T., Jr. A Study of the Fall Food Supply of the Ring-necked Pheasant

and the Bobwhite Quail in Washington Coimty, Rhode Island, M. S. thesis,

R. I. State College, 1940.

. Annual Report, N. Y. S. Cons. Dept., 1931.

. Annual Report, N. Y. S. Cons. Dept., 1936.

. Annual Report, N. Y. S. Cons. Dept., 1937.

Weather Conditions in Relation to Grouse

From the instant a potential grouse comes into existence as a fer-
tilized egg to the time it meets its usually violent end, it faces the
continuous vicissitudes of the weather. In its incubation period the
primary hazard is that of excessive chilling; w^hen a chick, hazards
from the heat or cold, the wind, and rain all play a part; while in
adult life the bleak cold, snow, ice, and storminess of winter must
be faced as they come. Humans, who also face essentially the same
handicaps, fare well or poorly in proportion to the adequacy of our
shelter and our food supply. To a considerable extent this is true
also with the grouse. The better their food and shelter, the better
may they cope with the dangers that adverse weather brings. In
actuality, since food and shelter are seldom perfect, the weather
plays an important part in limiting or reducing grouse numbers, or
in preparing the way for destruction by predators.

EFFECTS OF W^EATHER ON THE HABITS AND
NUMBERS OF GROUSE

Sunshine Gauges the Time of Nesting. The laying of eggs by a
hen grouse comes about only after a considerable period of internal
physiological change. The dormant ovaries are stimulated to growth
in early spring by the lengthening hours of daylight. Once started
the development of these organs and the laying of fertile eggs (fol-
lowing mating) comes about systematically. However, the date of
the beginning of egg-laying varies considerably in different years.
To illustrate: On Connecticut Hill the average first egg in 1931 was
laid on April 18; in the succeeding year it did not appear until
April 27.

Correlating the variations in nesting dates with the weather factors
brings out clearly that sunshine is the guiding element. When we

184

Weather Conditions in Relation to Grouse 185

compare Weather Bureau records of sunshine with grouse egg-laying
dates, we conclude that the greater the number of hours of sun-
shine in March and early April the earlier egg-laying will begin.
There seems to be no correlation with precipitation and only a slight
one with temperature. The latter is probably a by-product of the ef-
fect of day length and the involved sunniness on temperatures.

The part played by sunlight (or light) in determining the time
of egg-laying is corroborated by laboratory experiments which
show that grouse, as well as other game birds, poultry, trout, etc.,
can be stimulated to egg production months ahead of the normal
time by the use of artificial light.

Egg Losses Little Affected by Cold or Rain. A bird's egg has re-
markable ability to resist the hazards of changeable spring weather.
With eggs in the nest and before incubation has begun, tempera-
tures as low as twenty-one degrees Fahrenheit have been observed
and in no case has any adverse effect been noted. After incubation
the temperature is kept constant most of the time by the mother's
body, but even then the eggs can withstand surprisingly adverse
conditions. Temperatures well below freezing far into May, storms
that leave snow on the ground for as long as four days, and many
gully-washer rains, occur many times with no discernible effect on
the eggs. Eggs taken from a nest two days after the mother had
been killed, when temperatures had been below forty degrees,
hatched in an incubator a few days later.

Evidently the eggs can stand considerable hardship without
themselves being destroved. How much weakening of the young
is caused by such adversity, with resulting delayed mortality, is un-
known. It is reasonable to assume that chicks from eggs that have
been seriously exposed will be somewhat weaker than normal and
may, therefore, have a higher subsequent mortality. However, we
have not been able to correlate egg losses with variations in weather
conditions.

Leopold (1933) referred to the drowning of nests or young in
heavy rain or floods as due to accident. We have observed only one
nest destroyed by water and believe that such losses are ordinarily
of no particular significance.

Brood Mortality Affected by Weather. One of the most baffling
problems in the life history of the grouse is the high mortality of

186 The Ruffed Grouse

the young birds during the first three weeks after hatching (see
page 303 ) . Long an enigma to those interested in the welfare of this
bird, this loss has been conjectured by many authorities to be the
result of adverse weather. The evidence does indicate that unfavor-
able conditions of precipitation and temperature are often con-
nected with abnormally high brood losses, but it does not answer
the question of what causes the normally high losses at that period.
One of the first to suggest that excessive rain might be at the bot-
tom of the infant mortality was Sandys ( 1902 ) . He points out that
young grouse cannot stand a wetting. Conversely, he suggests that
five consecutive favorable seasons, i.e., dry from hatching time till
the chicks are past the critical stage, would mean a grand lot of
birds. He then concludes that the reversed conditions would mean
scarcity. Eaton (1910) says that in cold, wet seasons the eggs
hatch poorly and the young die from exposure. Forbush (1912)
linked the unseasonable weather from April to June of 1907 through-
out the Northeast with the great grouse decline of that year. He says
that eggs got chilled when females left the nest for food; setting
hens died from exhaustion, starvation, cold or disease; and chicks
that hatched disappeared. Stoddart (1918) listed bad weather in
May and June as second in importance among the many factors
contributing to the grouse scarcity in New York in 1916 and 1917.
Bump ( 1932) summarized some of the early experiences of the New
York study on this subject and pointed out instances in which some
grouse broods hatched in a protracted rainy period suffered severe
losses while others fared normally. Such an inequality of effect of
a given period of supposedly adverse weather on different groups
of grouse chicks has been noted repeatedly. On the occasion of one
severe and quite cold rainstorm in early June (temperature down
to 50° F., no sunshine for two succeeding days) we deliberately
flushed and thoroughly dispersed four different grouse broods only
a few days old. Each group was well soaked. Each of the four bands
was observed again during the next two weeks and in no case was
the loss greater than average.

Contrasting with these instances are many others wherein seri-
ous losses immediately followed severe cold and rain in June.
Grouse allowed to hatch their eggs in natural enclosures in cap-
tivity are almost always poor mothers, and the young may often
become chilled and wet. In all such cases observed, most of the

Weather Conditions in Relation to Grouse 187

chicks died within ten days after exposure. Some of the broods
cited by Bump {op. cit.) lost from half to three quarters of their
numbers within a few days after the bad weather. Prompt and ade-
quate brooding by the mother grouse no doubt aids in preventing
such losses.

If we assume that adverse weather in June will cause excessive
chick mortality, we might expect that the reverse conditions— warm,
dry weather in June— would be particularly beneficial, other things
being equal. However, 1933, a good season from this standpoint in
New York, had a very high brood loss. Other instances also provide
a full range of inconsistencies in respect to this factor.

The best opportunity that we had to evaluate the possibilities of
excessive rainfall affecting grouse brood survival occurred in 1935.
On July 7 and 8, at a time when grouse broods were past the usual
period of high mortality, came the most severe cloudburst in the
recorded history of southern New York. During this two-day period,
approximately ten inches of rain fell (a summer's normal supply),
about eight inches of it the first day. This storm caused unprece-
dented floods and destruction over an area of about ten counties.
It was immediately apparent that disaster had overtaken the young
grouse. Observations both in and out of the flood area clearly showed
that the abnormal brood losses were in the area of high rainfall. As a
result of this one storm, an additional twenty per cent of the grouse
chicks were lost over what would have been expected to perish.
This meant that the survival of chicks was only fifty per cent of the
normal, as an increase in mortality from the normal sixty per cent
to 1935's eighty per cent left only twenty per cent of maturing birds
instead of forty per cent {N. Y. S. Cons. Dept. Ann. Rep., 1936). This
extra loss resulted in a marked decrease in grouse on the area that
fall and illustrated how a moderate addition to the normal mortality
will cause a big loss in the proportion of birds maturing.

Hence, while severe cold and rain can prove disastrous to very
young grouse, the average annual early-season losses are only partly
explained. A normal, healthy grouse chick, with proper maternal
care, can get along pretty well in spite of the chilly nights, rain-
storms, and periods of protracted cloudiness.

Effects of Winter Weather on Adult Losses. Throughout the year
the condition of the weather causes marked changes in the selec-

188 The Ruffed Grouse

tion of cover by grouse (see page 94). In a direct sense, adult
giouse practically never succumb to the weather, that is, die of
exposure or freezing. The "practically" is included to admit the re-
mote possibility that death from exposure may conceivably occur.
It has not been observed, however, in all our observations under
extremes of weather of all sorts. In an indirect sense, weather often
makes grouse susceptible to accelerated predation, hence is an im-
portant element in their mortality story.

The literature is full of references to hardships brought on grouse
by winter. Most often these are associated with supposed starva-
tion induced by imprisonment under a heavily crusted snow follow-
ing snow-roosting. Samuels (1870) evidently considered this to be
commonplace, for he wrote: "It is a common occurrence to find them
in the spring dead, where they have been imprisoned under the
crust . . ." Allen (1927) considers this problem from the bird's
point of view, "quoting" a hen grouse in such a predicament thus:
"Rain or snow would freeze, making it impossible for me to get out
and I spent several days in hunger, imprisoned beneath the snow."
Forbush ( 1913 ) recognized that such an event was not usual and
concluded: ". . . usually they are vigorous enough to find a way
out somewhere." It is credible that death beneath the snow may
happen but it seems likely that most authors have overplayed a
theory or misinterpreted their observations. We do not know of a
single authenticated case of this phenomenon.

In enduring stormy winter weather, grouse regularly seek shel-
ter in coniferous trees or beneath the snow, often remaining there
for several days at a time, as attested by the accumulation of their
droppings. Apparently they take little or no food during these "holing
up" periods, yet ordinarily are not adversely affected.

The more snow there is on the ground and the more blustering
the weather, the more will the birds resort to snow-roosting. The
winter of 1935-36 in southern New York was a season of unusual
snow-roosting; deep snows lay on the ground all winter with no
important thaws. No instances of death from imprisonment were
observed, but the conditions did induce a higher than normal loss
from predation. We have already noted the increased vulnerability
of grouse in snow roosts (see page 46). During the 1935-36 win-
ter, it was common to find the story written in the snow of a grouse
pounced upon while at rest beneath its cool, white blanket. The mor-

Weather Conditions in Relation to Grouse 189

tality rate was very high, from twenty-five to one hundred per cent
more than in normal years. The increased losses occurred in Febru-
ary and March, coinciding with the period of unusual snow-roost-
ing. Whereas the normal curve of mortality reaches a peak in April,
or is at about the same height for March and April, the years of in-
creased winter predation brought on by adverse weather show a
peak in February or March. In 1936 it occurred in March.

Weather conditions other than snow and sleet are of little im-
portance in causing or inducing grouse mortahty. Low winter tem-
peratures, unless accompanied by a heavy snow, do not seem mate-
rially to aflFect the birds. The winter of 1933-34 gave ample chance
to learn of any possible connection between temperature and winter
loss. Over half the days of February gave below zero (Fahrenheit)
readings, with an extreme of thirty degrees below. The monthly
mean was thirteen degrees, which is twelve degrees below average.
Yet there was no indication of accelerated predation, and the mor-
tality records indicated only the normal winter losses.

Extremes of summer heat send the birds to coniferous shelter, as
does cold weather. Mousley (1919) attributed a noticeable increase
in grouse in Stanstead County, Quebec, in the summer of 1919 to
a prolonged hot spell. If rain could cause any increase in deaths of
adults, this should have been reflected in the 1935 summer cloud-
burst that so affected the youngsters. No losses of grown-ups attrib-
utable to this storm could be recognized.

Snow and Sleet in Relation to the Food Supply. The deep snows of
the northern winter conceal and largely render unavailable the
fruits and leaves on the ground and on low-growing plants. Under
these circumstances the birds resort to budding but it is remarkable
the quantities of food as ferns, wintergreen, Canada mayflower, and
the like that they manage to obtain under even the worst of condi-
tions. A man may be wholly rmable to see any of these plants, but
the grouse get at them somehow. The ability of the bird to subsist
on buds alone, for how long or how well we do not know, reduces
the chance of its being starved when so much of its preferred
foods is covered by snow.

Occasionally a freezing rain will encrust a snow, making the food
supply on the ground even less available, and at the same time may
sheath the tree buds with ice. Under such a condition grouse may be

190 The Ruffed Grouse

hard pressed to get food, but it rarely lasts long enough to affect
them seriously, as they can go several days without food and suffer
no apparent ill effects.

POPULATION FLUCTUATIONS AFFECTED BY
WEATHER CONDITIONS

We have already noted instances in grouse trends where extreme
weather conditions resulted either directly or indirectly in abnormal
grouse losses. How great a part such meteorological extremes have
played in causing or precipitating the large grouse declines of for-
mer years cannot be gauged by these observations, but they do
indicate the possibilities inherent in climatic factors. As we have
already noted, Woodruff ( 1907 ) gave "the extremely cold, wet, and
late spring of 1907" as second in his list of three factors that caused
the decline of that year. Forbush (1912) likewise correlated the
1907 grouse collapse with bad weather.

Knowing rather accurately when some of the most extraordinary
grouse reductions have taken place in New York and other parts of
the Northeast, we have examined the Weather Bureau records for
New York State in an attempt to ferret out any broad correlations
that might exist and the results seem too significant to be accidental.
The temperature and precipitation records are adequate as far back
as 1890, and sunshine data are available since 1909, The period
from 1890 to date covers four major and three minor grouse declines
in this state.

If a certain weather condition, or a combination of conditions,
could explain important decreases in the grouse population, presum-
ably severe instances of those circumstances would occur during
all years of these decreases and in no others. However, no single
weather factor does this, nor does any combination completely agree.
But the coincidence between periods of grouse decline and certain
severe weather conditions seems too close to be wholly accidental.

Since 1890 there have been four primary periods of sharply de-
creasing grouse abundance, approximating the years 1896-97, 1907,
1916-17, and 1927. Minor declines occurred in 1904, 1924 and 1935-
37. The evidence indicates that serious trouble from adverse weather
occurs from February to July and we may therefore confine our
analysis to these months.

Weather Conditions in Relation to Grouse 191

The temperature and precipitation records for 1896-97 show a
very severe late winter in 1896 and a very cold June in 1897. Febru-
ary and March snows in 1896 exceeded twenty-five inches each
month, more than fifty per cent above normal. Temperatures in
February were normal, but in March the mean was seven degrees
below the thirty-two degree norm. This low March temperature
condition was rivaled in only one year, 1916, another year of grouse
decrease. The 1897 June temperature mean was some three degrees
below the sixty-five degree norm, a condition that has occurred out-
side of grouse decline years only in 1902 and 1903, and in the latter
possibly contributing to the minor decline in 1904. Other conditions
were close to average in these two years.

The next big decline period of 1907 was notable for its cold spring.
April was four and seven-tenths degrees below normal in its mean,^
May five and five-tenths and June two and one-tenth, the latter be-
ing even worse during the first half of the month. This coolness was
accompanied by a marked deficiency in sunshine in April and May.
Precipitation was average though Woodruff (1907) and Forbush
( 1912 ) called it a cold and wet spring. The late winter of 1907 was
normal except for a six degree deficiency in the February mean.

As there is some indication that the 1907 grouse drop-off may have
begun in some areas in 1906, let us examine the records for that year
before proceeding further. A severely cold March (six degrees be-
low norm ) with snowfall well above average succeeded an ordinary
February. The spring was normal throughout.

The decline of 1916-17 concurred with some of the most severe
weather in the history of the state. The February temperatures in
1916 averaged three and five-tenths degrees low, and there was
seven and nine-tenths inches of snowfall in excess of the typical.
During March, an eight and one-tenth degrees deficiency in the
temperature mean, accompanied twenty-one and six-tenths inches of
snowfall above the usual. In the spring, sunshine was low through-
out, especially in April and June. Temperatures were normal in April
and May but very cool in June, being four and two-tenths degrees
below the average. Precipitation was normal in April, but excessive
in May and June. Turning to the winter of 1917, we find that it in-
cluded severe February temperatures and March precipitation, but

^ "Mean" weather records are deemed "average" or "normal," and are the standards
used by the Weather Bureau.

192 The Ruifed Grouse

was otherwise normal. April was abnormal only in being short in smi-
shine, while May was deficient in both sunshine and temperature.
Precipitation in June was five and eight-tenths inches, which is very
high; temperatures were low throughout, especially in the first half
of the month.

The most recent major die-off in New York took place in 1927, but
apparently began in 1926. This two-year period shows essentially
the same weather characteristics from late winter to early summer
as those just discussed. Snowfall in February 1926 exceeded twenty-
five inches, far above average, and was followed by March tempera-
tures that were some five degrees below noiTnal. The other mani-
festations of the winter weather were about typical. The April tem-
perature mean was thirty-nine and five-tenths degrees, the same as
in 1907, both the lowest on record. This cold weather continued
through May, but to a less abnormal degree, then in June maintained
an unfavorable spring record with a mean of sixty and eight-tenths
degrees, the same as in 1916, and next to lowest on record. Other
conditions were not adverse.

The winter and early spring periods in 1927 were average, except
for a small excess of snow in February. In May, however, tempera-
tures were low, rainfall high, and sunshine very much below normal.
Following this came another cold June, though not as bad as the
year before. The precipitation and sunshine factors in this June
were good.

Before examining the weather situation accompanying the minor
periods of decline, we may take note of the elements common to the
four occasions just discussed. In all, there was a condition of late
winter weather severity, with heavy snow and unusually extreme
cold. Likewise there was a cold June; only in 1906-07 were June
temperatures not extreme; in 1907 both April and May, as well as
June, had low thermal records.

The minor decline of 1904 was attended by extremely low Febru-
ary and April temperatures, and March too was colder than usual.
Conditions through other parts of the winter and spring were not
severely adverse. It may well be that the very low June temperature
and high rainfall in 1903, together with lack of sunshine, contributed
to this drop in grouse numbers.

Another intermediate decrease occurred in 1924. Then the weather
record was mostly orthodox but with some extremes. There was

Weather Conditions in Relation to Grouse 193

somewhat more snow in February than usual, but not a serious
amount; April had very high rainfall and was somewhat colder
than usual; May was very cold, its rainfall high, and sunshine lack-
ing; June was somewhat colder than the normal. Its first half was
very cold. The previous year, 1923, was not notable for any extremes,
although June was very cloudy and the late winter was quite cold.

The most recent minor decline in New York began in 1935. Dur-
ing the critical months from February through June all weather
factors were normal or favorable. The cloudburst and flood of early
July (see page 187) offers the only apparent correlation of weather
with this decline. Since this catastrophe was effective over ten
counties in the best grouse range of the state, it probably offers the
explanation for the rather small decrease that was revealed in the
state hunting kill, as well as for the severe drop on Connecticut
Hill within the flood area.

The 1935 decline indicates how an individual short-lived storm
may affect great numbers of grouse. If such a phenomenon was con-
nected with any of the previous drop-offs we were unable to single
it out. It must be recognized though that if such a storm did occur
it might well explain some of the failures to correlate more general
weather conditions with drops in the grouse population.

If there is any certain connection between adverse weather and
grouse decimations, it apparently is tied in with excessive snow in
February and March, usually linked with extremely low tempera-
tiu-es, or with low temperatures in June, especially during the early
part, often supplemented by heavy rains and lack of sunshine, or
both. All of the notable grouse declines in New York since 1890,
when the weather records began, can be explained by these condi-
tions. If the correlations indicated are only accidental, then there
were probably other years having these same weather characteristics
but which failed to show widespread grouse declines.

An examination of the corresponding records for all the years
since 1890 reveals very low June temperatures (mean below 61.6°)
in 1897, 1902, 1903, 1916, 1926, and 1927. Of these 1902 is the only
one definitely not of a poor grouse year. It was not preceded by a
bad winter. Years of low, but not extremely low, June temperatures
(below 63°) were 1907, 1910, 1912, 1918, 1924, and 1928. Of these
1907 and 1924 coincided with grouse decreases and in both, the
early June thermal record was very severe. The years 1918 and 1928

194 The Ruffed Grouse

were seasons of low abundance, when these conditions could only
prolong the existing grouse depression. Both 1910 and 1912 were
preceded by mild winters.

Very bad February or March snow conditions occurred in 1893,
1896, 1897, 1900, 1908, 1910, 1916, 1920, 1928, and 1932. Those not
happening in a period of declining or low abundance, 1893, 1900,
1910, 1920, and 1932, were all followed by favorable June weather.
Several of these periods of excessive snow were of short duration,
as for example in 1932.

Every time severe February-March snow conditions and very low
temperatures, followed by a very cold June, occurred in two suc-
cessive years, a giouse decline followed. And every time there was
an important grouse decline, these weather conditions had prevailed.

A close scrutiny of the details of this record shows that it was not
uniform each time. Each decline was unquestionably the product
of several conditions, each of which varied on different occasions.
Predation varies widely, even when conditioned by weather factors.
We believe that disease played a significant part in the 1927 decline.
Was this epidemic set up by weather factors? We don't know. It
hardly seems that a haK-century record of weather-grouse-trend
correlations could have so much in common, as appears to be the
case, unless there was actually a real functional connection. It is far
from a perfect record of cause and effect; but is so impressive that
the importance of climatic factors in relation to grouse population
trends can hardly be discounted lightly. Weather conditions play a
big part in grouse fluctuations both great and small.

REFERENCES AND CITATION SOURCES ON WEATHER
CONDITIONS IN RELATION TO RUFFED GROUSE

Allen, A. A. The Autobiography of a Mother Grouse, Bird Lore, Vol. XXIX,

No. 6, December, 1927.
Bump, Gardiner. New York's RufiFed Grouse Survey, Game Breeder, Vol. 86,

No. 2, Feb. 1932.
DeLury, R. E. Sunspots and Living Things, Trans. 17th Am. Game Conf.,

1930.
DeLury, R. E., and O'Connor, J. L. Regional Types of Response of Wildlife

to the Sunspot Cycle, Trans. 1st N. A. Wildlife Conf., 1936.
Eaton, E. H. Birds of New York, 1910.
Forbush, E. H. Game Birds, Wild Fowl and Shore Birds of Massachusetts

and Adjacent States, 1912.

Weather Conditions in Relation to Grouse

195

Forbush, E. H. Useful Birds and Their Protection, 1913.

Hatfield, D. M. Aniinal Populations and Sunspot Cycles, Chicago Naturalist,

Vol. 3, No. 4, 1940.
Leopold, Aldo. Game Management, 1933.
Mousely, H. Further Notes and Observations on the Birds of Hatley, Stanstead

County, Quebec, 1919; Auk, Vol. 38, No. 1, Jan., 1927.
Samuels, E. A. The Birds of New England, 1870.
Sandys, E. Upland Game Birds, 1902.
Stoddart, A. M. Ruffed Grouse in New York State, 1918.
Wing, L. W. Wildlife Cycles in Relation to the Sun, Trans. 21st Am. Game

Conf., 1935.
. New York State Conservation Department Annual Reports for years

1931, 1932, 1935, 1936.

Interrelationships of Ruffed Grouse to Mammals and to

Other Birds

SPECIES THAT PREY UPON GROUSE

Discussion of predators is fraught with many dangers. Not the
least of these is prejudice in dealing with creatures that compete
with us. This attitude we must completely discard. The otlier major
difficulty is the tendency to accept simple action and reaction as a
complete story of the relations between predator and prey species.
That is far from justifiable. The subject of predation is one of the
most intricate, if not the most complex, in the field of animal rela-
tionships. But before we consider the drama of predation as it re-
lates to grouse, let us become acquainted with the actors. For
present purpose the grouse is the prey species and we know it
reasonably well. Let us take a look at its natural enemies, and then
some of the other creatures that play subordinate roles to the victim
and the "villains."

A bare list of the enemies of the grouse is impressive even though
we know little of their specific depredations. Study reveals that some
of them exact a greater toll than others. They vary, too, as to the
time of their attacks, some during nesting season as egg eaters,
others in the summer as destroyers of the young, or at any time as
predators on the grown birds. We have listed in three categories in
Table 8 those of significance in the Northeast, those of primary,
secondary, and occasional importance. It has been necessary to list
a few in more than one classification because they require separate
ratings for dijfferent types of predation.

The species in the first group are listed in probable order of im-
portance for south-central New York. This order varies considerably
in diflFerent parts of the Northeast, and some of those listed as sec-

196

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Plate 1. A (Upper). When the white man came, farms were hewn out of the
wilderness. B (Lower) . Where the land clearance work was confined to small scat-
tered units, the range was improved for grouse. However, extensive agricultural
development gradually pushed the grouse out of much of its original range.

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Plate 2. A (Upper left). The drumming of the cock grouse is one of the
wonders of the bird world. The sound is made by striking the wings against air.
B (Upper right). Strutting before the female bird, the male stands stiffly with
tail raised and spread, wings drooping at the side, and the head drawn back into
its erected ruff. C (Lower). Young grouse in a brood dust bath. In addition to ex-
posed dirt, grouse often use well-decayed wood of old stumps for dusting areas.
Photos by A. A. Allen.

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Plate 3. A (Upper left). The nest was built at the base of a big tree, on the
sunny side. B (Upper right). Eleven eggs composed the clutch, an average num-
ber. C (Lower left). The hen grouse watches carefully for enemies before step-
ping onto the nest to continue incubation. Can you pick out clearly the well-
camouflaged bird? D (Lower right). The chicks remain in the nest only a few
hours after hatching. As soon as they are dried off, and the weather is clear, they
leave home, never to return. C and D photos- hij A. A, Allen,

Plate 4. Grouse Chicks at Different Ages. A (Upper left). One day old-a
cute, completely downy little fellow. B (Upper right). At five days of age, the
chick shows marked improvement in wing feathering. C (Lower left). By the
time seventeen days have passed, the wings are well developed, and it can fly
short distances. D (Lower right). At five weeks of age the juvenile feathering is
quite complete with tail and all. It now resembles a small edition of its parents
and can night-roost by itself. Photos by A. A. Allen.

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Plate 5. Grouse roosts are likely to be almost anywhere in spring, summer or
fall, but in winter they usually sleep in trees, or if the weather is bad, in the snow.
A (Upper left) , Grouse going to roost in a tree. B (Upper right) . Grouse just after
landing in the snow. C (Lower left ) . Pheasant and grouse eggs in one nest. This
phenomenon is not common. In all cases observed, the pheasant has been the in-
truder, the grouse the one imposed upon. Photo btj A. A. Allen. D (Lower right).
A snow roost after being vacated, showing the usual pile of droppings.

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Plate 6. Two Types of Grouse Range. A (Upper). Disconnected cover, wood-
lands separated by open fields, typical of much of the Northeast. B (Lower) . Con-
tinuous cover (beyond valley), extensive forests found in the several mountainous
areas from the Appalachians to the Adirondacks and Maine. Photos Courtesy of
Pennsylvania Game Commission.

Plate 7. An aerial view of a portion of the Connecticut Hill submarginal farm-
ing area well illustrates the broken nature of this type of grouse range. Woodland
is interspersed with open fields, the fencerows and woods margins are generally
brushy. Some of the abandoned fields are being reclaimed by woody vegetation.
Note the woodland clearings (squares) and lanes in lower part of area. These
are grouse management cuttings.

Plate 8. Overgrown land as a shrubby border to woodlands provides desirable
interspersion of cover types. A (Upper left) . Open land, overgrown land and hard-
wood woods in close juxtaposition. B (Upper right) . Shrub cover next to coniferous
woods. C (Lower left). Woodlands lacking a shrubby edge lose valuable inter-
spersion; the coniferous type above has excellent shelter but is deficient in handy
food cover. D (Lower right). Pure alder most often occurs on moist soils, and is
much used by broods.

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Plate 9. Overgrown land is of particular value as feeding cover, especially in
summer and fall. A (Upper left). Subtype of mixed hardwoods composition.
B (Upper right). Mixed conifer-hardwood subtype has better shelter. C (Lower
left). Mixed oak composition, common from Long Island and Pennsylvania south-
ward. D (Lower right) . Pure stand of popple along woods border, one of the most
useful types of overgrown land.

Plate 10. Slashings bring summer food cover to woodland areas. A (Upper).
Newly cut subtype is characterized by rank herbaceous growth and briars, lasts
from three to ten years usually. B (Lower). Old cuttings have a stand of shrubs
and sapling trees, but with fewer herbs and briars. It will soon become a pole-
stage woodland, and will then have a very different value as cover for grouse.

Plate 11. Hardwood Woodlands. Of particular value as nesting cover, this
type varies immensely. A (Upper). Pure hardwoods are deficient in shelter.
B (Lower left). A scattering of conifers improves this type for grouse. C (Lower
right). Broadleaved evergreens, such as mountain laurel, serve the same purpose
as conifers, are food plants as well. Mature stands are less valuable to grouse than
second growth.

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Plate 12. A (Upper), Small openings made by removing a large tree, or two
or three, here and there, known as spot lumbering, in hardwood stands is the best
of the many subtypes. B (Lower). The mixed oak association, so prevalent in the
middle Appalachian range, is quite unproductive of grouse where it occurs in
large, unbroken areas.

Plate 13. Mixed Woodland. A (Upper). Hardwood overstory of near-mahire
trees with much hemlock in lower growth-of less use to grouse than the younger
age class. B (Lower) . Young stand of hemlock and hardwoods, the best all-purpose
type of grouse cover.

Plate 14. A (Upper). Overstory of mature white pine with completely hard-
wood understory. The coniferous shelter of this woodland is so high that it is
largely useless to grouse. B (Lower). Hard pines (Virginia pine) and oaks com-
pose a mixed woodland cover type common on sandy soils and on burned areas.

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Plate 15. Coniferous Cover. Hemlock, pine, spruce, fir, cedar furnish the basis
of the best protective shelter for grouse. Its value depends on species, age, density
mainly. A (Upper). Hemlock on left has sheltering branches close to ground,
white pine on right has shelter only in crown. B (Lower). Close-up of low-hanging
hemlock boughs showing their provision of winter protection.

Plate 16. A (Upper left). Some of the broad-leaved evergreens, as mountain
laurel, may substitute for the conifers in furnishing winter shelter close to the
ground. B (Upper right). The hard pines (pitch and Virginia pines shown) are
the predominant conifers in much of the Appalachian range from central Pennsyl-
vania south. C (Lower) . Spruces furnish most of the winter shelter in the northern-
most range, from the Adirondacks and northern New England into Canada.
B and C photos by U. S. Forest Service.

Plate 17. Plant succession must be one of the bases of grouse management.
A (Upper). An old white pine has established the makings of a stand of solid
pine around it while near by (in the foreground) the new woods will be mixed
pine and hardwoods in a few years. B (Lower left). A view of the north end of
the Connecticut Hill area, showing how natural plant succession of hardwoods with
scattered conifers is gradually changing the open fields, first to brush overgrown
land, ultimately to forest. No planting has been done on this part of the area.
Management must maintain openings and overgrown land with ax and plow.
C ( Lower right ) . Among the factors that influence plant succession is the nearness
to an existing woodland or hedgerow. Except for light-seeded species, most plants
invade an old field slowly from the edges, as the hemlock and maple are here.

Plate 18. Grouse nests are usually at the base of some object, most commonly
a tree. A (Upper left) . Hen grouse on nest at base of a large beech. Note the fern
and dead twigs that offer some camouflage. B (Upper right). A successful nest
showing the characteristic appearance of hatched eggs. Note the two unhatched
eggs. C (Lower left). Grouse incubating on nest at base of stump. Photo by A. A.
Allen. D. (Lower right). Grouse brooding chicks. Photo by C. W. SevcringJiaus.

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Plate 19. A (Upper). A single grouse meal. Crop contents of a female grouse
from Sullivan County, Pennsylvania, December 13, 1940. Top: left, leaves of
mountain laurel; right, cherry buds. Below: left, birch buds; right, fruit of moun-
tain holly {Ilex monticola), the first record of this species being eaten by grouse.
B (Lower). A grouse with a full crop, skinned out to show the immense capacity
of this organ when fully distended. The food thus stored passes to the gizzard only
as fast as it can be ground and digested. PJioto by A. A. Allen.

Plate 20. Herbaceous plants provide the grouse with a considerable part of its
food. Among the more important are: A (Upper left). Ferns, especially the woods
fern (shown), the fronds of which are eaten mostly from autumn to mid-spring.
B (Upper right). Sedges (Carex) are used from spring to fall. The leaves are
eaten as long as they are green, and the seeds are taken in summer and fall.
C (Lower left). Important among winter foods of grouse are tree buds. Some of
the most used species are: a. Yellow Birch; b. Black Birch; c. Blue Birch; d. Hop-
hornbeam; e. Trembling Aspen; f. Apple; g. Black Cherry. D (Lower right).
Sheep sorrel (upper plant), a small field weed. Its sour-tasting leaves are relished
by grouse. The Canada mayflower (lower plant), a plant of the woodland floor,
has red berries that are eaten from fall to spring.

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Plate 21. Summer fruits are among the most important foods of grouse for
young and old alike. Two of the most important are: A (Upper left). Brambles, of
which a blackberry is shown here. B (Upper right). Blueberries. The Dryland
blueberry is the one illustrated. Photos by W. R. Van Dersal. C (Lower). The
partridgeberry provides food all year round, but it is especially useful in the spring
when fruit is scarce.

Plate 22. Autumn is the season of plentiful fruits and nuts, and the grouse takes
full advantage of them. Among the important fall fruits and nuts that furnish
food for grouse are: A & B (upper) the acorns, of which those of the northern red
oak (A) and scrub oak (B) are illustrated. C (Lower left). Cherries, the choke-
cherry (illustrated) and black cherry being most important. D (Lower right).
The dogwoods, among which the gray or panicled is most used.

Plate 23. Other important fall food for grouse. A (Upper left). The vibur-
nums. The mapleleaf viburnum is shown here. B (Upper right), Greenbriers, of
which the sawbrier is illustrated. The dogwoods provide fruits for fall and early
winter use. Gray dogwood is among the best (see Plate 22D). C (Lower left).
Flowering dogwood is most important from southern New England and Pennsyl-
vania south. D (Lower right). Red osier primarily used in northern region.

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Plate 24. A ( Upper left ) . Old apple orchards furnish excellent feeding cover
all year, especially if close to a woodland. This orchard with its mixture of briars
and other shrubs, is perfectly situated. B (Upper right). Wild apples near pine
clumps are a perfect combination in autumn. C (Lower left). The beech furnishes
a preferred food when it has a nut crop. It is a component of the woods next to
the orchard above. If the old trees are cut out as "weeds" the food value to grouse
will be lost. D (Lower right). The sumacs are an important source of winter
food. The grouse feeds on the bobs of some species (R. typhina shown in center),
and on the fruit of the two poisonous ones, of which the poison ivy is shown in D.
Note the sumac in the foreground of the orchard above. These shrubs must have
full sunlight to survive.

Plate 25. Winter foods are not a serious problem to the grouse on good range
except for variety. These plants help break up the predominance of buds as a
winter diet. A (Upper left). Mountain laurel, whose leaves are a staple food.
B (Upper right). Highbush cranberry, one of the viburnums. C (Lower left).
Wild rose, whose hips are eaten when accessible. D (Lower right). Teaberry, or
wintergreen, is a small plant of the woodland floor.

Plate 26. The cherries are useful for both fruits and buds. Three species are
used most: A (Upper). Wild black cherry. B (Lower). Pin cherry; and choke-
cherry (see Plate 22C).

Plate 27. The viburnums are an important group of shrubs that furnish fruit
for fall and winter grouse food. Two of the more important species are tire maple-
leaf viburnum (see Plate 23A), and the highbush cranberry (see Plate 25B).
Among other useful viburnums are: A (Upper left). Nanny berry. B (Upper right).
Hobblebush. C (Lower left). Blackhaw. D( Lower right). Arrowwood.

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Plate 28. Wild grape {Vitis labrusca), in this instance, is one of the most im-
portant fruits in the grouse diet in fall and early winter. A (Upper) . Prolific growth
along a woodland edge. B (Lower). Close-up of fruit.

Plate 29. Among the secondary grouse foods are several that furnish fruit that
assumes great importance in certain regions and at some periods. Among these are;
A (Upper left). Black chokeberry, a small bush. B (Upper right). Bittersweet,
a climbing vine. C (Lower). Winterberry, one of the hollies.

Plate 30. Other secondary grouse foods furnishing fruit are: A (Upper left).
Virginia creeper, vine. B (Upper right). Bayberry, most prevalent near the coast.
C (Lower left). Elder, a common fencerow shrub. D (Lower right). Mountain
ash, a small woodland tree. E (Center). Serviceberry, another woodland tree
whose fruits ripen in early summer.

Plate 31. A (Upper), Farming activities greatly affect the grouse food sup-
ply. The type of woodland cutting carried on, care of fencerows (or lack of it),
pasture fencing, ploughing and cultivation all show their effects in this picture.
The ploughed fields have no cover, but maintain edges against the woods. Some
fencerows are clean, others brushy. Pasture is fenced, and some of the former
pasture has been abandoned to grow into brushy land, then to woodland.
B (Lower). When deep snows blanket the ground, grouse resort to buds for most
of their food. When the day is calm, though, they wander through the snow seek-
ing fruits and greens.

Plate 32. The telltale marks of feathers and tracks in the snow reveal tragedies
for the ruffed grouse. A (Upper). Fox evidence. B (Lower). Horned owl work
(whitewash of owl droppings does not show against snow).

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Plate 33. A (Upper). Grouse nest destroyed by a horned owl killing setting
hen, after which a fox ate the eggs. Note feathers and broken egg shells. B (Lower
left). The common skunk loves eggs, and occasionally stumbles onto a grouse
nest. In some areas it may be one of the most destructive of grouse predators.
C (Lower right). Great horned owl, one of the most efficient of all predators, a
prime enemy of the grouse. Photo by Pennsylvania Game Commission.

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Plate 34. The rabbits are among the rodents whose population fluctuations are
correlated with those of the grouse. A (Upper left). The cottontail, found mainly
in the disconnected cover areas of grouse range. Photo by A. A. Allen. B (Upper
right). The snowshoe hare, found mainly in the extensive northern forests. C (In-
sert). The squirrels, of which the gray squirrel is shown, affect the grouse in two
ways, as a buffer between the grouse and its enemies, and as a food competitor.
D (Lower). The squirrels' love for nuts, and their keen ability to find them, often
make the establishment of a direct-seeded oak, walnut or other nut tree planta-
tion impossible. The photograph shows the empty shells of a wahmt planting after
being dug up and eaten by squirrels.

CROP
ESOPHAGUS

PROVENTRICULUS

Crop Worm
(Capillaria annulata)

Plate 35. A (Upper). Diagram of alimentary canal of the ruffed grouse
showing location of some parasite infestations. B (Lower left). Grouse stomachs:
a. normal; b. lightly parasitized with Disphanjnx spiralis; c. heavily parasitized with
D. spiralis. C (Lower right). Stomach worms (Displuirynx spiralis) (small) and
intestinal worm (Ascaridia lineata) (large worm), showing comparative size.
Photos ]nj A. A. Allen.

Plate 36. A (Upper left). The modem hunter pursues grouse as a sport.
B (Upper right). To many hunters the grouse is the king of game birds. When a
spht-second shot brings a clean kill, they feel a sense of real accomplishment.
C (Lower). Man's use of well-trained pointing dogs has done much to make hunt-
ing a fine sport. In the early part of this century most of the pointing dogs of the
northern states were trained on grouse but today the majority have been "spoiled"
on pheasants and a top-notch grouse dog is a rarity. Photo hij J. M. Sloan.

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Plate 37. Man's operations as a farmer affect gfrouse habitat in many ways. His
plougfiing and cultivation of crop fields prevent woody cover from expanding,
and at the same time maintain the valuable woodland edges ne,\t to the fields.

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Plate 38. A (Upper). Man's interest in managing the land to produce grouse
is often directly affected by what he does or does not do to control other animals,
such as deer. In recent years, deer populations in many areas such as parts of
Pennsylvania, Wisconsin, the Pisgah National Forest, North Carolina, have been
permitted to increase to such an extent that their browsing has partially destroyed
the range for deer and grouse alike. B (Lower). When deer populations are too
large for the carrying capacity of the range, it is impossible to establish success-
ful tree plantations. Note damage to white pine seedling from deer browsing.

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Plate 39. As a lumberman, man sometimes improves and sometimes destroys
the cover values. A (Upper). Overcutting destroys shelter, eliminates much of
the interspersion of types, and prevents high quality regeneration by eroding the
forest floor. B (Lower). Woods roads break up the cover, make valuable edges,
feeding lanes, and, in summer, dusting and sunning spots.

Plate 40. A (Upper). Seed stock refuges as a medium of hunting control
are of little value in grouse management except in the most intensely shot areas.
Photo shows wire bounding refuge (on right) on New York game lands. B (Lower).
The woodcock, shown on its nest, is one of the more important game birds com-
monly sharing range with the ruffed grouse.

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Plate 41. A (Upper). Protection from grazing is one of the first essentials of
woodland management, whether for game or timber. Note the good understory
to the right of the fence where protected, and the lack of any saplings or shrubs
on the left where grazed. B (Lower left). Overgrazing in a woodland not only
prevents plant reproduction, but has a detrimental effect on the soil itself. If car-
ried on for many years, the effects of compaction and erosion may last a long time.
C ( Lower right ) . Fire in woodlands may destroy grouse cover almost completely
for a period. If an area is repeatedlv burned, as is so much of the middle Appa-
lachian range, it cannot support grouse at all.

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Plate 42. Interplantings may sometimes be made to advantage in woodland
openings or in partially-seeded old fields. A (Upper). Farm woodlands that have
been heavily grazed by cattle can often be interplanted successfully. B (Lower
left). Norway spruce interplanted in a woodland glade. C (Lower right). An
old field partially taken over by naturally-seeded white pine. The interplanting
of other species could well be done here.

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Plate 43. A (Upper). Brush piles made from the waste tops and branches of
cuttings serve well as temporary winter shelter. B (Lower). The margin of the
woodland lane is a good place to improve food conditions. By releasing shrubs
and herbs where the sun can reach them, fruiting and seeding will improve.

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Plate 44. Plantings may improve existing coverts for grouse. A (Upper). In-
terplanting of white pine in an area of cut-over hardwoods improves the shelter
value. B (Lower left). Shrub border of silky dogwood planted along a woodland
edge. C (Lower right). Shrub borders along the woodland edge may be devel-
oped by cutting out tree species and favoring the shrubs in the woods margin.

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Plate 45. Group-selection harvest of timber provides small but valuable open-
ings for grouse. Here v^e see a quarter-acre clearing resulting from the removal
of three large crop trees. A (Upper). Just after cutting, showing stumps, and sun-
shine reaching the ground. B (Lower). Same after two years, showing thicket
development of shrubs, and tree sprouts and seedlings.

Plate 46. Slashings, or clear-cuttings, oflFer a means of improving and man-
aging woodlands for the benefit of ruffed grouse. A (Upper). Aerial view of a
Connecticut Hill woodland showing both unit and lane cuttings. B (Lower left).
Close-up of a one-acre clear-cutting grown to briars, elderberry and other shrubs
and trees. C (Lower right). A lane slashing, thirty feet wide, providmg needed
brush cover and valuable edges.

^

lep^ - --c

MflMlllW

^ tB^^^^"^

Plate 47. A (Upper) . Clear-cut areas provide edges, so needed by game. Lanes
give much edge for the area cut. B (Lower left). An experimental slashing unit
created by poisoning the trees, leaving the dead trees standing. Note the Virginia
creeper climbing the dead trees upon receiving full sunlight. C (Lower right).
The edge of a unit slashing showing the change in the character of the cover.

>^-^"-'

<SI!l^ k, £^lbi**^-.-^-^^i?^^^^

Platk 48. Wherever the terrain permits, it is desirable to plant in contour-
ploughed furrows. Survival and growth are hoth improved, especially for hard-
woods. A (Upper). Contour-furrowed field being planted. B (Lower). The same
plantation five years later.

Plate 49. Selection of species that are adapted to the planting site is vital to
success. A (Upper). In this ten-year-old plantation, the arborvitae (being pointed
out), while surviving, has grown but a very few inches due to an unsatisfactory
site. B (Lower). In this shrub border planting, the first (on right, next to grain
field ) , third and fourth rows are a success, but the second row is a failure because
of the use of a species unsuited to the soil. In this instance, the misplaced plant was
winterberry in a well-drained soil.

Plate 50. Reforestation provides interspersiou ot cover t\pcs by arrangement
of species. A (Upper left). An aerial view of two fields on Connecticut Hill show-
ing pattern of conifer and hardwood plantings (shrub bands cannot be distin-
guished from hardwood trees). B & C (Upper right and lower left). Same two
fields from the ground, left and right respectively. D (Lower right). Field plant-
ing showing alternate bands of conifers and hardwoods.

y ^^^J^tiik-A

■^'

Plate 51. Coniferous reforestation, or natural pure stands of pines and other
conifers, vary greatly in value to grouse according to age. A (Upper). For the
first few years the young trees affect the old field complex very little. B (Center),
From about five to fifteen years of age the stand is mixed with natural reproduc-
tion, usually hardwoods, and serves as overgrown land. It may be of considerable
value as a feeding area, is likely to be excellent cover for cottontails. C (Lower).
After the crown closes, often between fifteen and twenty-five years of age, the
competing natural vegetation is rapidly driven out. As the lower limbs die, the
shelter \alue is reduced. Food is practically non-existent.

Plate 52. A (Upper left). The trees gradually prune themselves and the tightly
closed canopy prevents any germination of ground cover. It is this stage, lasting
from twenty to thirty years, that is sometimes referred to as a "biological desert."
B (Upper right). At around the forty-to-fifty-year age period, the crown begins to
open and the return of herbs and hardwoods begins on the woodland floor.
C (Lower). At maturity we find a top crown of pine, partially broken, with a com-
plete hardwood-shrub-herb stand underneath. The shelter is out of reach, but
iood has now returned.

Plate 53. Field plantations should strive for a balanced habitat. A (Upper).
Mixed planting of conifers and hardwoods to provide both food and cover.
B (Lower). Border of shnibs provides a desirable thicket type next to a pine
planting.

■W^^*'^'

:^^

■7

V

■'f«

Plate 54. A ten-year-okl planting of northern red oak.

"'■"^>"

Plati: 55. Nursery beds of seedling bayberry, one of the shrubs recommended for
planting. These plants are in a nursery of the U. S. Soil Conservation Service.

''<'^^>'^^;^'''*!:^E»'— '

// 71-

/f

^^4*^.

^•r^

7'

/

Plate 56. A (Upper). Game refuges are generally associated with public shoot-
ing ground areas when established on public lands. The type of marker shown
here is used in New York. B (Lower). Trapping fur-bearers for their pelts on a
sustained-yield basis is the only predator control recommended on grouse range.
The red fox shown is one of the most important ruffed grouse enemies.

Interrelationships of Ruffed Grouse 197

Table 8.

The More Important Pkkdators of Ruffed Grouse in the Northeast ^

Species of Important as Destroyers of:

predator ~ ^gg^" Chicks Adults

I. Those of Primary Importance

Foxes (red and gray) x

X

Great horned owl

X

Weasels (New York and small brown) x

Skunk X

Goshawk

X

X

Cooper's hawk

X

X

Sharp-shinned hawk

X

II. Those of Secondary Importance

Crow X

Raccoon x

Red squirrel x

Weasels

X

X

Great homed owl x ^

X

Foxes

X

III. Occasional Predators

Bobcat

X

Woodchuck X

Chipmunk x

Dog X

Barred owl

X

X

Black snake x

X

Red-tailed hawk

X

X

Crow

X

House cat

X

X

Marsh hawk

X

X

Porcupine (rarely) x

^ "Importance" as used in this section is based on the relative number of grouse
destroyed by one predator as compared to other predators, and does not imply the
same significance as a limiting factor. See Chapter X for relative significance of all
factors.

- Man is intentionally omitted as he is treated elsewhere.

^ The horned owl does not touch the eggs but sometimes causes their loss by killing
tlie nesting female. The other hawks listed may occasionally do tlie same.

ondary predators are of primary importance in local areas, and vice
versa.

The question of the importance of the grouse in the diet of a given
predator, or of a predator as a destroyer of grouse, may be ap-
proached from two directions. Both are needed to bring out the

198 The Ruffed Grouse

complete story. From the standpoint of the prey species, the per
centage of its population that the predator takes is all that counts,
regardless of its ranking in the predator's diet. As a problem in the
food habits of a predator, the volume of food that a prey species
furnishes is all that matters at the moment, aside from the effect on
the status of the prey species. To illustrate let us take a hypothetical
case. We have two situations. In the first we have one family of foxes
and one hundred grouse in a covert. In the second we have one
family of foxes and ten grouse. Let us assume in each case that the
foxes take ten grouse. In the first instance the prey species is little
affected, losing only ten per cent of its numbers; in the second it is
entirely wiped out. Yet in both cases the grouse furnished the same
part of a fox family's diet. Let us assume it was two per cent. It was
relatively unimportant to the predator, but to the prey species in
one covert it was fatal.

Now let us begin over and take a different course with the same
data. Say the foxes take fifty grouse in the first situation and none
in the second. Then the grouse becomes a big item of food, ten per
cent, to the fox in case I, without critically damaging the prey spe-
cies, while in the second case, though the fox has not gained, there
are fewer grouse than are left from the preyed-on population.

Thus the importance of prey species as food and the importance
of predators as enemies are relative matters that must be correlated
with other food habits and with population data to be properly ap-
praised. Since we are concerned primarily with the grouse, we will
examine the effect of the predatory species on this species, even ff
grouse is not an important item in their diets. We will also give some
information on the part that grouse play in the subsistence of the
various predators. It can be said categorically, however, that the
grouse is rarely an item of vital importance in the diet of any preda-
tory species, except possibly the goshawk.

Red and Gray Foxes. Both species of foxes native to the Northeast
commonly take grouse. As predators, one is about as efiicient as the
other, and the relative importance of either one is generally in pro-
portion to its abundance. The red fox is common throughout the
Northeast, whereas the gray is found mainly from central-New York
southward. So far as we know, their effects on grouse are much the
same, therefore we will discuss them together.

Interrelationships of Ruffed Grouse 199

The foxes probably warrant the title of "number one grouse
enemy" over most of the Northeast. In the northernmost range they
may sometimes be displaced by the goshawk, and in occasional
localities by the great horned owl. Their leading position derives
from their adeptness at taking grouse in all stages from the egg to
the adult. As a nest predator they are by far the most destructive,
and as an enemy of adult grouse they are outranked only by the
homed owl and, in the northern range, by the goshawk. They are
unimportant as destroyers of young grouse in summer, although
they do take some.

Our experience indicates that a fox is not only a connoisseur of
grouse eggs but is also quite skillful in finding them. On one occa-
sion an observer had been placed in a house built in a tree at some
distance from an incubating grouse to make a twenty-four-hour
record of her activities. Near dawn on one morning a red fox was
observed at some distance coming through the woods. It disappeared
from sight and returned to view several times as it coursed the cover
back and forth a-hunting. Gradually it approached closer to the
observer and likewise closer and closer to the setting grouse. In
time it came to within a few feet of the hen, and she flushed with
the usual whir of leaves. The fox sprang after her but missed and
ran swiftly in the direction the bird had disappeared. Presently it
returned to approximately the place of action and with deliberate
efficiency sniffed the base of every tree and stump in the vicinity
until it found the nest of eggs.

That fox knew what it was doing. It obviously had flushed grouse
before, had eaten grouse eggs before, and knew the relation be-
tween them. We had evidence that a single fox broke up seven
grouse nests in a woodland of about two hundred acres in one spring.
Whether or not this particular evidence may be taken as the full
truth, it is certain that the fox as a species is a rather efficient nest
hunter and that some individuals become particularly adept. On
the other hand, I recall a grouse that successfully brought off a
clutch of eggs vdthin fifty feet of a red fox den in active use. The
fox kits had played within twenty feet of the setting hen as evi-
denced by bones and skin of kills that they had dropped there.
That did not seem to indicate predatory efficiency.

Foxes consistently destroy more grouse nests than do any other
predators in New York. The proportion of all grouse nests taken by

200 The Ruffed Grouse

foxes on Connecticut Hill has averaged about one in five. The per-
centage of predator-destroyed nests attiibutable to foxes varied
much from year to year but averaged forty-three and five-tenths per
cent over the whole state as summarized for two hundred and thirty-
nine such nests observed from 1930 to 1934 {N. Y. S. Cons. Dept.
Ann. Rep., 1934).

The records of foxes taking young grouse have been very few. It
should be noted that until the birds are nearly half grown they are
probably eaten whole, thus leaving no evidence. But even though
summer food habits of the fox have not been well studied from
stomach contents, circumstances indicate that foxes are not impor-
tant as predators on grouse chicks at this season when other food
is plentiful.

With the advent of winter the fox once again becomes a serious
enemy. From autumn through spring the toll of adult grouse taken
by foxes is significant, the greatest numbers being captured from
midwinter to midspring. The number of grouse that are killed by
foxes in the winter is somewhat dependent upon the weather. With
deep snows, the grouse resort more to the snow for roosting; there
some of them will certainly be found by foxes. The vulnerability of
grouse in snow roosts is great. One of our observers, upon noticing
one day that many birds in snow roosts flushed at very close range,
determined to try to catch one by hand. Of the next three birds he
saw in snow roosts he was able to stalk and catch two, and the third
barely escaped. A fox too can learn to locate the bird by the slight
snow mound at the end of a grouse track, and surely it can outdo a
man at stalking.

The proportion of grouse lost through predation on Connecticut
Hill that is attributed to foxes averaged twenty-one and four-tenths
per cent of one hundred and ninety-one remains for which the enemy
was identified {N. Y. S. Cons. Dep. Ann. Rep., 1933). The propor-
tion of adult mortality ^ resulting from predation is about ninety-five
per cent. As grouse mortality averages some forty per cent for the
same period, foxes kill nearly ten per cent of the total population.
In considering these figures we should not overlook the possibility
that some of this apparent prey may have been carrion, or crippled
or sick birds. However, it is our considered opinion that making
such an allowance would not affect the figures materially.

^ Aside from human destruction.

Interrelationships of Ruffed Grouse 201

We have seen how important the foxes are in grouse mortahty.
How important is tlie grouse to the fox? The answer is that it repre-
sents a food dehcacy, even as to you and me, but not a staple item
of food. In following fox tracks ( mostly red ) for nearly three hun-
dred miles on Connecticut Hill, no evidence was found of grouse
being caught, and apparently only one attempt to catch a grouse
had been made. The event, though rare, does occur. The accumu-
lated animal remains at four fox dens on Connecticut Hill occupied
by one pair of red foxes from March through May contained four
grouse among thirty items of prey (N. Y. S. Cons. Dept. Ann. Rep.,
1934). But all comprehensive records of fox food habits show the
ruffed grouse to be a negligible item in the year-round food supply.

Before we review some typical records of fox food habits, it may
be well to consider certain factors that affect the interpretation of
these records. In the first place, the fox is most destructive as a nest
predator in the month of May, but few indeed are the fox stomach
records representing that season. Further, as the fox eats the egg
contents, taking little if any shell, the stomach and scat records do
not fully bring out the extent of this food habit.^ Another important
factor in food habits records that often leads to misinterpretation,
is the source of material. It is obvious that foxes taken from areas
that have no grouse will have no grouse remains in their stomachs.
To interpret properly the importance of grouse in fox diet, we must
give weight to these facts: Stomach or fecal analysis will not reveal
the extent of egg-eating, even in specimens collected during the
nesting season; summaries of stomach or fecal analyses will ordi-
narily be conservative in revealing grouse in the diet, as a part or aU
of the specimens may have been taken in localities where there were
no grouse.

Hamilton (1937) summarizes the food of the red fox as follows:
"Investigations in the Midwest, Michigan, New England, New York,
and Virginia are all in essential agreement. The studies indicate that
the fox feeds chiefly upon fruit and berries, small mammals— chief
among which are mice— carrion, insects, and an occasional bird. I do
not doubt it prefers a tender grouse to a half-dozen mice, but the
fact remains that mice are far more abundant than grouse, and much
easier to catch." Examination of five hundred and thirty-seven scats

^ Of two fox scats collected in June in Centre Count)', Pa., one contained remains

of grouse eggs (Kozicky, 1943).

202 The Ruffed Grouse

of red foxes from eastern New York (Cook and Hamilton, 1944)
revealed a frequency of occurrence of grouse averaging one and
five-tenths per cent for the whole year; increasing to four per cent
in winter and five per cent in April, by June the incidence fell to
three per cent. No grouse parts were found in summer specimens
but in autumn one per cent of the scats included some grouse re-
mains. Dearborn (1932), writing of Michigan conditions, says of
the ruffed grouse in the red fox diet: "Opposition to foxes is based
largely on the belief that they are very destructive to game, espe-
cially to game birds. The only game bird of importance where most
of this fox material was gathered is the partridge, or ruffed grouse,
Bonasa umhellus, which was common there both years. According
to the evidence collected in this investigation, the average fox eats
not more than two giouse per year." Handley (1934), in reporting
on the analyses of both red and gray foxes in Virginia, lists no grouse
remains in twenty-seven summer and fall collected specimens and
eighty-four winter specimens. As quail were recorded in these rec-
ords, it may well be that the specimens were not taken in grouse
range. Errington (1935) reported on fox stomach analyses from
Wisconsin and Iowa specimens. He found no ruffed grouse in the
forty-six red foxes taken in fall and winter, and but one in the
seventy-two gray foxes taken from fall to spring. Likewise, Erring-
ton found no ruffed grouse remains at the one hundred and thirteen
red fox dens studied or in one-thousand, one hundred and seventy-
five feces samples. Here again the records indicate that the majority
of the specimens were taken from farm land and not in grouse range.
Analyses of one hundred and forty-eight stomachs (those that
contained food out of a total of two hundred and twenty) from
Pennsylvania (English & Bennett, 1942) showed grouse to be a
fairly common item of food for the red fox. Of one hundred and
thirty-six late summer and fall specimens, five contained grouse,
amounting to four and four-tenths per cent of the total volume of
food. In this group, grouse were exceeded in volume by six other
foods, all mammals, except grasshoppers. Only eleven winter stom-
achs were examined, and one of these contained a trace of grouse
remains. Two of the five grouse captured from late summer to fall
were in the twenty August records and constituted fifteen and six-
tenths per cent of the food, ranking second to remains of wood-
chuck. Two more grouse were among the forty-two October sped-

Interrelationships of Ruffed Grouse 203

mens and were ninth in volume of food eaten. One November
stomach out of fourteen had grouse remains, and no grouse were
in the fifty-six September specimens. Bennett & English (1942)
also reported on analyses of twenty-nine gray fox stomachs taken
from August to February. Only one, a winter specimen, contained
a trace of grouse.

Of one hundred and eighty-six fox scats gathered in Huntingdon
County, Pa., in the fall months, three contained grouse remains,
constituting eight per cent of the volume ( Kozicky, 1943 ) .

The food of the red fox in southern New Hampshire was studied
through examination of two hundred and six scats covering the
period from October 1 to April 30 and two hundred and seven for
the May 1 to September 30 period (Eadie, 1943). The author con-
cludes, in reference to the use of grouse by the fox, "Ruffed grouse
were abundant during the period of collection but were only spar-
ingly represented in the scats ( three per cent, winter; two per cent,
summer)."

An examination of food eaten by forty-one gray foxes in Ohio
revealed that only one stomach contained ruffed grouse (Bezdek,
1943).

From these references we can see that the ruffed grouse does not
rate a high place in the scale of fox foods, even considering the con-
servative nature of these records. Nevertheless, foxes are a very
important enemy of the grouse.

Great Horned Owl. The horned owl ( see Plate 33C ) is probably
the most efficient of all grouse predators. With very powerful talons,
soundless flight, a twilight-to-dawn attack, and adequate weight,
it is the most capable of all the common predatory birds in taking
grouse. It is widely distributed throughout the Northeast, and is
generally the most serious predator on adult grouse. It ranks second
to the foxes only because it is responsible for the loss of relatively
few clutches of grouse eggs.

On the Connecticut Hill area the gi-eat horned owl was by a wide
margin the most deadly of the adult grouse predators. More than
half ^ of the mature grouse killed by predators each year were taken

^ The figures given for proportions of grouse killed by the various winged predators
are conservative, since a considerable number of remains were the work of hawk or
owl, with the evidence inconclusive as to species. The total proportion of kills was
75 per cent by avian species, and 25 per cent by mammals.

204 The Ruffed Grouse

by this bird. Since many of the signs could not be distinguished with
complete accuracy from those of other hawks and owls, this figure
is only approximate. It is regularly somewhat over fifty per cent of
grouse adult losses from all causes.

While the prominence of the homed owl as a predator of grown
grouse far overshadows its indirect effect on nests, a considerable
number of grouse eggs are lost due to the owls killing the mother
bird. On Connecticut Hill about one in twenty-five of the nests
destroyed was attributed to avian predators, largely this species of

owl.

We have already noted that predators are not a primary cause of
mortality among the young grouse in the summer. No doubt the
homed owl takes a small toll then, but the fact that feathers of
very young grouse are not left in the woods but are consumed with
tlie birds, makes an accmate survey of the owl predation impossible.
Some remains of immature grouse have been found in horned owl
pellets and the feather remains of a few have been taken as evidence
of horned owl killings. It is probable that this species ranks next,
after the sharp-shinned and Cooper's hawks, as a destroyer of young
grouse.

We have seen that the foxes, though veiy important as grouse
predators, do not depend upon them for a high proportion of their
food. This is also tme of the great homed owl. Although first among
the predators upon adult ruffed grouse, it still does not make a very
high percentage of its food of grouse. We must again conclude,
therefore, that the proportion of a predator's food composed of a
given prey species does not necessarily measure, or even indicate,
the place that that predator plays in the ecolog)' of the prey species.

Typical of the records of horned owl food habits are those sum-
marized by McDowell ( 1940 ) . Ruffed giouse made up one and nine-
tenths per cent of the whole diet of the nine hundred and eighty-
three owls examined. They were taken between November and May.
The grouse were found in only fourteen of the stomachs, or one and
four-tenths per cent of the total, and were far outranked by a num-
ber of other prey species, mainly rodents. Errington, Hamerstrom
and Hamerstrom (1940) examined four thousand, eight hundred
and thirty-eight pellets from Iowa and Wisconsin and found twenty-
two of them, or five-tenths of one per cent, to contain remains of
ruffed grouse. They say: "Ruffed grouse . . . populations in the

Interrelationships of Ruffed Grouse 205

north-central region seem to he in many ways vulnerable to, or rela-
tively secure from, predation much as are bohwhite populations,"
referring to the fact that predation primarily affects that portion
of a population in excess of the carrying capacity of the range.

New York and Small Brown Weasels. We presume that both species
of weasels common in the Northeast are capable of taking grouse
eggs, and occasionally grouse, although evidence is lacking to
identffy the work of each. The New York weasel is probably the
more destructive, owing to its larger size. Most important in grouse
ecology as nest predators, the weasel is also of some significance in
taking the birds too.

The weasels ranked third only to the foxes and skunk in nest de-
struction on Connecticut Hill. They were accounted responsible for
ten per cent of the nests rifled by predators. Only three and one-
tenth per cent, as an average, of the loss of adult grouse from preda-
tors was attributed to weasels {N. Y, S. Cons. Dept. Ann. Rep.,
1933). These animals are no doubt responsible also for the loss of a
few young birds in the summer.

All scientific writings on the food habits of weasels show that their
food is largely rodents, mainly mice. Hamilton (1937) found that
ninety-five per cent of their fall and winter food consisted of small
mammals, and he does not mention grouse at all as a part of their
diet.

Skimk. The common skunk ( see Plate 33B ) is a remarkable, though
not an efficient, predator of grouse. It is unable to prey upon the
birds themselves, but often stumbles upon and robs a nest. When
these animals are abundant, the number of nests broken up by them
may be quite large. On Connecticut Hill the skunk ranked second
among grouse nest predators, and was adjudged responsible for
eleven per cent of all nests destroyed by predators ( op. cit. ) .

As might be expected, studies of skunk stomachs and feces have
not revealed this habit. Skunks crush the eggs thoroughly when
eating them and swallow bits of shell only by accident. The proba-
bility of recovering egg shells in these specimens is limited also by
the short time in the spring during which they are taken. Hamilton
(1936) examined five hundred and seventy fecal specimens and
thirty stomachs representing the spring and summer seasons and
found no remains of grouse eggs. Likewise, Dearborn (1932) says

206 The RufiFed Grouse

"Not a trace of a game bird egg was found." In both of these studies
no grouse either were noted. The grouse eggs that skunks eat are
to them merely an occasional treat, not a dependable or sizable item
of food.

Goshawk. This magnificent hawk is the only one that might dis-
pute the horned owl's pre-eminence as a killer of giown grouse. A
daytime hunter, it has a different type of attack; it simply overtakes
the grouse by superior speed and maneuverability. This is all the
proof needed of this bird's hunting prowess.

It may be fortunate for the grouse that the goshawk is not more
generally plentiful. It is primarily found from northern New Eng-
land and New York northward, although it occurs locally southward
into Pennsylvania. It visited the Connecticut Hill area only in winter
and then only in occasional years. From 1930 through 1933 it was
not recorded at all; then a few were observed each winter for the
next three years. In these winters, the goshawks levied a high toll
of grouse, considering their own numbers. The proportion of all
grouse killed by predators that were taken by goshawks averaged
only about four per cent the first four years ( op. cit. ) . Thus it ranked
fourth among predators of adult grouse, behind the horned owl,
foxes, and Cooper's hawk. But in its more year-round range, the
goshawk probably is the most destructive of all adult grouse enemies.
No doubt where the hawks breed, they take young grouse in the
summer too.

The goshawk is the one species of predator for which the ruffed
grouse furnishes a really big proportion of the food. McAtee ( 1935)
says: Grouse, chiefly ruffed grouse, were determined in thirty-one
of the stomachs out of two hundred and forty-three examined, or
twelve and eight-tenths per cent. McDowell (1941), in examining
one hundred and one goshawk stomachs taken from November
through May in Pennsylvania, records an incidence of thirteen and
nine-tenths per cent of grouse, which amounted to thirteen and two-
tenths per cent of this predator's whole diet. Mendall (1944) found
remains of grouse in five of thirty-one (sixteen and one-tenth per
cent) of these hawks taken in Maine throughout the year. McAtee
concludes: "On the economic side, there is comparatively little that
can be said in its favor." In spite of its destructiveness, I still have
a great admiration for the goshawk. In it, and the duck hawk, nature

Interrelationships of Ruffed Grouse 207

has evoh^ed the "Spitfires," "Mustangs," and "Messerschmitts" of
the bird world.

Cooper's and Sharp-shinned Hawks. These two "blue darters," or
"bird hawks" are generally common over the Northeast. They are
substantially smaller editions of the goshawk, less powerful, and
hence less able to take large prey. The Cooper's is the larger of the
two and the only one that as a rule can kill an adult grouse. Both
birds are notable predators of the young grouse, the sharpshin being
the more important on Connecticut Hill due to its greater numbers.
McAtee (1935) recorded the Cooper's hawk as having taken grouse
in a few cases, but reported no grouse in the nine hundred and forty-
four stomachs of the sharpshin examined. McDowell ( 1941 ) found
the remains of two grouse in one hundred and eight stomachs of
Cooper's hawks from Pennsylvania, these constituting one and eight-
tenths per cent of the total food. On Connecticut Hill, the Cooper's
hawk was third among the predators of adult grouse, averaging over
five per cent of the kills made by predators {N. Y. S. Cons. Dept.
Ann. Rep., 1933).

The data on mortality of the young grouse caused by these hawks
are too meager to summarize. The number of chicks taken by them
is of some significance. They were clearly the first-ranking predators
of the chicks on the study area. The number of remains found in-
creased towards late July and in August, but this was at least pardy
due to the larger size and more substantial nature of the feathers at
that time.

It is unlikely that grouse ever compose a very high proportion of
the diet of the sharpshin, and only in occasional local instances of
that of the Cooper's hawk.

Crow. This "black marauder" is widely renowned as an egg eater.
There is little doubt but that crows will raid any nest of eggs they
discover, that of the grouse being no exception. The number of
grouse nests broken up by crows on Connecticut Hill from 1930-34
averaged six per cent of the predator-destroyed nests, but this pro-
portion declined somewhat in later years. This record entitles the
crow to a place of rather limited importance as a grouse predator.

On one occasion a flying crow was seen carrying a very young
grouse chick. We must thus list the species among predators of the
young grouse too, although this habit is surely of small importance.

208 The Ruffed Grouse

While eggs of all kinds may be an appreciable part of the crow's
spring food, it would be rare indeed for grouse eggs alone to amount
to a very significant item.

Other Mammalian Grouse Predators. Both the raccoon and the red
squirrel have in some years been of considerable significance as
grouse nest predators on the Connecticut Hill area. On the average,
the raccoon was responsible for the destruction of about eight per
cent of the nests taken by all predators, and the red squirrel for only
about two per cent. The bobcat, and no doubt the Canada lynx
where it occurs, are of some importance as grouse predators, pri-
marily in winter. Neither of these wild cats occurred on Connecticut
Hill, however. Of one hundred and forty bobcat stomachs examined
by Hamilton and Hunter (1939) from animals taken from fall to
late winter in Vermont, twelve contained grouse. These composed
five and five-tenths per cent of the bulk of the food. The authors
conclude: "It is evident the bobcat has little trouble in catching
grouse. Dearborn found no evidence of grouse remains in more
than three hundred feces, although grouse were abundant around
the swamps where the wildcats live.'" A trace of grouse remains
was found in the intestine of one mink from among one hundred and
two specimens examined from southern Michigan (Sealandt, 1943).
Woodchucks are known to have broken up a few nests, and the
porcupine has been recorded as eating grouse eggs {Pennsylvania
Game News, May, 1933 ) . Chipmunks have a habit of stealing grouse
eggs and storing them in holes. King ( 1937) found this to be of con-
siderable significance in Minnesota and noted that playful chip-
munks used the eggs like marbles. House cats that roam grouse
coverts no doubt take some grouse. Dogs, likewise, will occasionally
break up a giouse nest, but rarely will catch a grown grouse. It is at
least possible that shrews (Blarina) may take grouse eggs occasion-
ally, since they have been recorded to have eaten whippoorwill eggs.

Other Avian Grouse Predators. Several species of hawks in addi-
tion to the accipitrine trio already discussed occasionally take a
grouse. None are of much significance. McAtee (1935) reported
grouse in three of seven hundred and fifty-four stomachs of red-
tailed hawks, in four of six hundred and one marsh hawks, and,
surprisingly enough, in one stomach of a broad-winged hawk. Men-
dall (1944) reports the remains of one bird, "apparently a ruffed

Interrelationships of Ruffed Grouse 209

grouse," in one stomach of the fourteen from red-shouldered hawks
taken in Maine, and one grouse in thirty-sLx stomachs of marsh
hawks. It is hkely that duck hawks and pigeon hawks may also occa-
sionally get a grouse. Among the owls, the barred and snowy owls
take an occasional grouse. Gross ( 1944 ) records the occurrence of
grouse remains in five of one hundred and twenty-seven stomachs
of snowy owls examined from birds taken in the northeastern states
during winter incursions. However, none of these species are of any
significance as grouse enemies.

It should be noted here that while no specific instances of this
latter group of hawks and owls taking grouse on Connecticut Hill
were noted, it is reasonable to suppose that a few of the cases attrib-
uted to the horned owl and the accipitrine hawks may have been
misidentified and actually were the work of some other species.

Reptilian Grouse Predators. The black snake is the only reptile
that has been recorded as a grouse predator. Records of nests being
destroyed by black snakes are not uncommon, and two cases of
grouse chicks being killed by black snakes have been recorded
(Pennsylvania Game News, August, 1935; West Virginia Conserva-
tion, Dec, 1941 ) . It is probable that other snakes too may take grouse
eggs, even though no records are available. In the southern Appa-
lachians where snakes are numerous, they might even be of some
significance in connection with grouse egg losses. T. E. Clark of
Virginia advised me that snakes, including the pilot black, black
racer, timber rattler, and pine snake, are important grouse nest pred-
ators there and occasionally take young giouse.

RELATION OF PREDATION TO COVER TYPES

The concept of "escape cover" used in wild-life management indi-
cates that certain cover conditions are better in preventing attack,
or successful attack, by predators than others. Generally, it has been
thought that dense shelter, such as that of thorny thickets and
coniferous types, served this purpose best for game birds. The
types of cover in which grouse remains left by predators are
found as compared with the incidence of grouse in the same cover
indicates that mixed woodland is the most hazardous type of cover.
It is somewhat ironical that the mixed-woodland type also is the

210 The Ruffed Grouse

best all-use cover for grouse. Apparently it may also serve a like
purpose for some predatory animals, and in any event they would
tend to hunt wherever their prey is most plentiful. Least hazardous
are coniferous woods and overgrown lands, including thickets.
The usual concept of safe cover, therefore, seems to hold for ruffed
grouse. However, it must be remembered that factors that benefit
grouse, such as escape cover, are effective only for populations
well accommodated by the environment ( that is, below the thresh-
old of security ) .

PREDATORS AS AN AGENT OF SANITATION

It is generally recognized that predators take the prey that is easi-
est to get— limited, of course, by the range of their food habits. This
being the case, it is reasoned that sick, injured, and weak individuals
of a prey species will be the first to succumb, other things being
equal. Thus the predator acts to maintain or increase the virility of
its prey species. Likewise, many predators are also scavengers to a
degree. They clean the range of dead animals that escaped attack
before dying from some cause other than predation. How do these
ideas stand analysis with respect to grouse?

The ruffed grouse normally maintains its full vitality in the face
of affliction until very near death. By the time a bird shows weak-
ness of flight, or sluggishness of activity, it is ordinarily very near
death. The time during which a predator might take a sick grouse
easier than a well one is very limited. Although not very significant,
the sanitation principle probably does hold with grouse within this
limitation. Probably of greater significance than physical weakness,
is the vulnerability of birds that are in unfamiliar habitat. Similarly,
grouse living under strain of intolerance by confreres may fall vic-
tim to an enemy more readily than would otherwise have been the
case.

When it comes to cleaning up carcasses of birds that have met
death through means other than enemy attack, the scavengers are
very efiicient. For example, three birds that were shot and left lying
without being handled by man, were taken the first, second, and
fourth nights of exposure. It is rare indeed for a grouse to lie on the
ground long enough for the flesh to rot appreciably.

Interrelationships of Rutfed Grouse 211

THE EFFECT OF PREDATOR CONTROL ON RUFFED
GROUSE POPULATIONS '

We have aheady observed that any discussion covering the rela-
tions of predators and game is most delicate. That phase of the sub-
ject dealing with control of the predators, and its value, is probably
the most delicate of all. Our object, therefore, must be to evaluate
dispassionately the efiFectiveness of control on the prevalence of the
predatoiy species themselves and the resulting efiFects on the ruffed
grouse populations.

Predator control studies were made on the Connecticut Hill area.
New York, in 1930-32, and continued in 1933-35. Two types of con-
trol experiments were made: complete control, where all predatory
species were taken; and selective control, under which foxes (two
species ) and weasels ( two species ) were taken.

The initial experiment in 1930-32 was designed to appraise the
effects of complete predator elimination. Of course this was far from
accomplished, the degiee of success with any species depending
upon its mobility and its susceptibility to trapping. The trapped
area consisted of one thousand, four hundred and twenty-five acres
of giouse coverts while the untrapped check-area covered one thou-
sand, two hundred and twenty-three acres. The fact that these two
tracts lay adjacent to each other, one being the northern, the other
the southern part of the whole study area, unquestionably acted to
reduce differentials in both grouse populations and mortality. Hence,
any conclusions as to positive effectiveness of predator control in
improving grouse numbers would probably be conservative.

A total of five hundred and fifty-seven predators were taken by
trap and shot during the first two-year period, three hundred and
twenty-one the first year (October 1, 1930-September 30, 1931) and
two hundred and thirty-six the second year (October 1, 1931-August
31, 1932), including fifty-six homed owls, twenty-five foxes, sixty-six
skunks, forty-three weasels and twenty-two accipitrine hawks.

Records of the numbers taken the second year indicate that re-
duction was markedly successful with respect to the long-eared owl,
red-tailed hawk, and small brown weasel; and moderately success-

^ This account is taken largely from an article of the same title, by the author, in
the Journal of Wildlife Management, October, 1939.

212 The Ruffed Grouse

ful with the honied owl, marsh hawk, and crow/ Species which
showed httle or no reduction the second year included the Cooper's
hawk, sharp-shinned hawk, sparrow hawk, red fox, gray fox, skunk,
domestic cat, red squirrel, and raccoon. Considering only the more
important grouse predators, control apparently showed some effect
on the horned owl, crow and small brown weasel; but made little
dent in the Cooper's hawk, sharp-shinned hawk, red fox, gray fox,
skunk, and raccoon numbers.

Even though some of the most important grouse predators re-
mained as abundant the second year as the first, there may have
been an immediate effect on the grouse by the temporary reduction.

The data indicate that the predator control reduced the nest
mortality markedly both years. During the first year the brood- and
adult-mortality records show a small differential favoring the trapped
sections, but in the second year the brood mortality was practically
identical on the two tracts and the adult mortality was actually
higher on the controlled portion. But even though the predator con-
trol proved unable consistently to lower the brood or adult losses,
the reduction in nest loss accomplished might produce a higher
shootable fall population.

In the fall of 1931, while the grouse population of the area was
still well below carrying capacity, the predator-controlled portion
had a markedly higher population density. These additional grouse,
twenty-five per cent more than on the check area, may reasonably
be credited to the control of predators. Then the second year, with
a nearly peak population at hand, the effect of the predator control
was negligible, although still producing a slightly more dense popu-
lation than developed on the untrapped subarea. It appears then
that the increase in adult mortality on the controlled portion the
second year, presumably brought on by the higher population
density the first fall, almost completely offset the saving in nests.
That is to say, the inevitable inverse relationship of productivity to
breeding density canceled any possible benefits of predator control.
Thus the vital part that population level plays in determining grouse
losses indicates the futility of attempts to regulate them.

The value of predator control in high population seems, from this
test, to be negligible.

^ Since these species are migratory, the indicated results may have resulted merely
from variations in the annual flights.

Interrelationships of Ruffed Grouse 213

Following the two-)'ears' study of predator control in 1931 and
1932, a year was allowed to elapse and then the experiments were
resumed. With the possibility in mind that the results of the first
experiments may have been affected by factors peculiar to the two
subareas, the trapped and untrapped areas were reversed in the
second series of tests beginning late in 1933.

Trapping and shooting were carried on from October 1, 1933, to
April 30, 1934, rather than for the full year as previously. The 1934
experiments continued the complete control with a no-trapping
check area. A total of one hundred and one birds and ninety-four
mammals were taken, including thirty-seven homed owls, fourteen
foxes, nineteen skunks, ten weasels, and twenty accipitrine hawks.

As in the earlier experiments, the outstanding species taken was
the homed owl, which is probably the most efficient ( if not always
most important) of the common grouse predators. The mammalian
predators showed practically the same abundance as in the first
experiments, with the two species of foxes being most important.

The nest mortality on the subareas for 1934 was twenty per cent
for the trapped portion and forty per cent for the untrapped. The
following summer, the brood mortalities were fifty-two and three-
tenths per cent and fifty-six and three-tenths per cent respectively.
By calculation, the loss of adults on the trapped area was twenty-
three and seven-tenths per cent and that on the untrapped thirty-
two and two-tenths per cent.

In reversing the trapped and untrapped areas the results, too, have
been reversed. Nest mortality was decreased by one half, brood
mortality slightly lessened, and adult losses lowered appreciably.
We conclude that the predator-control-effect data were not seriously
confused by other factors, although detailed accuracy of the results
cannot be claimed. If comparability of the results of the two experi-
ments was maintained, the 1934 fall population densities on the two
subareas should have shown little difference, as was the case in 1932,
since 1934 was another peak year. The density on September 1 on
the trapped area proved to be a bird to five and six-tenths acres
while on the untrapped portion the ratio was five and five-tenths
acres per grouse. Thus, we find the result repeated that in a peak
year even effective reduction of the nest mortalitv bv predator con-
trol does not appreciably increase the fall population on unshot
areas.

214 The Ruffed Grouse

The main objective of the 1935 experiments on Connecticut Hill
was to learn the effectiveness of fox-weasel control, these animals
being the most destructive nest predators. But with the fall of 1934,
a situation arose that seemed to present an unusual opportunity to
get further valuable data on complete control. One section on Con-
necticut Hill attained a September density of two and five-tenths
acres per grouse (equivalent of one and eight-tenths acres per
grouse of actual coverts, including no open land). This represented
the highest population density any covert had attained. It was de-
cided to subject this section alone to complete predator control to
determine if by so doing this saturated population density could be
maintained. Indirectly, it gave a better opportunity to obtain data
on dispersion or disease that might result from this heavy popula-
tion.

The predatory species taken during the 1935 trapping and shoot-
ing period, October 1, 1934 to April 30, 1935, totaled one hundred
and twenty-one individuals, of which ninety-seven were from the
complete control subarea and twenty-four from the selective control
subarea. These included sixteen homed owls, twenty-five foxes, and
thirteen weasels. Most significant in this record are the takes of
horned owls and foxes.

While the analysis of the data for 1935 is complicated by addi-
tional factors, the broad effects followed the same course as in pre-
vious years. Both complete and selective predator control proved
effective in reducing nest losses. The selective control brought a
reduction of about forty per cent in the nest mortality. Again, little
or no reduction of brood losses was accomplished by either type of
control— complete or selective.

The adult losses ran exactly opposite to the objectives, being
greatest on the complete control subarea, least on the non-control
sections. Because the losses on the complete-control section were
accentuated by a marked dispersal of birds from the high popula-
tion of the fall of 1934, examination of the predator decimation
would seem to give a fairer comparison. Based on grouse remains
found, we find that nearly twenty per cent of the 1934 fall popula-
tion were picked up on the complete control subarea, nearly twenty-
two per cent on the selective control and only thirteen per cent on
the no-control sections. Thus, despite drastic control of predators,
the high population density still induced a higher rate of predation

Interrelationships of Ruffed Grouse 215

than occurred without control on the areas used as a check but
which had much lower populations to start with. Thus, the high
decimation expected in peak populations occurred regardless of
elimination of predators.

From all the data on hand, we must conclude that intensive
predator control of any type, while it may be markedly effective in
reducing nesting loss, will not produce a higher shootable fall popu-
lation of grouse during years of high abundance. During years of
low grouse numbers, the evidence shows that predator control may
increase appreciably the fall grouse population. But even under
these conditions, with the grouse population increasing anyway, the
justiiication for deliberate predator control is very doubtful.

Control measures designed to benefit game populations often take
the form of bounties. In this connection, Gerstell (1937), in report-
ing upon the experience of the Pennsylvania Game Commission,
concludes:". . . as a predator-control measure the payment of boun-
ties has proven grossly inefficient ... it has been impossible to
prove that the operation of the bounty system over a relatively long
period of years has improved game conditions. Furthermore, it was
shown that the annual amount of money expended for bounty pay-
ments was controlled not by the abundance of predators, but prin-
cipally by climatic and general economic conditions." What more
need be added? There is no substantial evidence to justify intensive
predator control for the benefit of grouse populations.

THE ABUNDANCE OF BUFFER SPECIES AS IT AFFECTS
PREDATOR FOOD HABITS AND GROUSE MORTALITY

Buffer species are those animals that furnish the bulk of food for
predatory animals. They are generally abundant, sometimes exceed-
ingly so, have a high-reproductive rate, and wide distribution. They
are the staple animal foods of the foxes, weasels, owls, hawks, etc.
Predominantly they are rodents, and they are markedly cyclic in
their population trends. In the Northeast the more prominent ones
are: rabbits (cottontail rabbit, snowshoe hare) (see Plate 34A, B),
squirrels (red and gray) (see Plate 34C), mice-field (Microtus),
wood (Peromtjscus) , red-backed (Evotomys), and others— shrews,
and woodchuck. Since they are relatively easy to catch, because
of their abundance, their slowness afoot, or inability to protect

216 The Ruffed Grouse

themselves, they tend to act as buffers in reducing predator pressure
on other prey. If, because of a scarcity of buffers, the predatory
population pursues the game birds more intensively, more game
birds will succumb. Thus there is a close theoretical relationship be-
tween the rodents and grouse. How does it work out actually?

Determination of populations of rodent and predatory species on
extensive areas is a trick still to be mastered. However, methods for
determining population trends are available and, we believe, are
reasonably accurate. These include state game kill records; "sign"
recorded on field censuses, reduced to a number-per-man-day basis;
sample-track counts taken over marked trails of standard length un-
der as nearly uniform conditions as possible. All of these methods
have their shortcomings, but using all of them some trends appear.

Several interesting trends come to light. Beginning with the early
1930's, all of these rodents increased to an irruptive peak in numbers
in 1935. Then from 1935 to 1936 they all suffered a catastrophic de-
cline. Three out of four rabbits died, four of five field mice, and nine
out of every ten red and gray squirrels. This fact is supported by
general census data, by the game kill reports ( see Fig. 9 ) for cotton-
tails and gray squirrels (although the general decline in cottontails
apparently began two years before), and by the well-defined and
much-publicized migration and die-off of gray squirrels in the fall
of 1935.

The decline in cottontails was not so complete from 1935 to 1936
as that of the other animals, but it continued for two more years
while recovery of all the others began. This three-year period of cot-
tontail decline is substantiated by the state game kill records.

The uniformity of the rise and fall of these species from 1930 to
1936 is notable. But very significant variations in the recovery rate
followed. As already pointed out, the cottontails continued to de-
cline for another two years; the mice began a rapid recovery the
next year, and the red squirrels a gradual one. The recovery in gray
squirrels began the second year, as the mice and red squirrels com-
pleted their recovery to normal abundance. Then from 1938 to 1939,
as the cottontails began their increase, the others again declined,
the squirrels only moderately and for a single year, the mice signifi-
cantly and for at least two years. The cottontails also showed some
loss in the second year of the latter period.

From the standpoint of the effect of these fluctuations on the

Interrelationsliips of Rufied Grouse 217

predatory species that depend upon them for food, and the indirect
effects upon the ruffed grouse, two very significant facts are perti-
nent. In spite of the gieat coincidence of the periods of major decline
of these rodents, there is enough dissimilarity in tlieir population
drops so that they are not all very scarce at once. Thus, in 1936, the
cottontails had only declined part way when the others "hit bottom."

2.00

1.75

1.50

125

yi.oo

|.75

.50

.25

/

/ \

/ COnONTAIL RABBIT \ /

-- / \ /

\ ^

\

\

V

/

/

A

GRAY SQUIRREL / \

^ '-

^ ^'" RUFFED GROUSE

1930 1931 1932 1933 1934 1935 1936 1937 1938

YEARS (Autumn)

Fic. 9. New York State Game Kill Records 1930-1938.

In 1937, the mice had increased considerably above their low point
and the cottontails had still not reached their low. In these two
worst years for buffer foods, there were still some species in fair
relative abundance. The second significant point is that even when
the rodents reached the trough of their population trends there were
still a lot of them, enough to support a fair number of predators.

The only one of the predator groups that is recorded in the state
game kill take is the foxes. We find a slight decrease indicated in
these records for 1935 (see Fig. 9).

218

The Ruffed Grouse

We may fairly conclude that the numbers of several of the preda-
tory animals are responsive to changes in the numbers of their chief
prey species. The response in decline of predators may take up to
two years following marked changes in rodent numbers to become
significantly apparent.

PREDATORS
(Primarily foxes, weasels,
horned owls and hawks)

RUFFED GROUSE

1930-31 1931-32 1932-33 1933-34

1934-35
YEARS

1935-36 1936-37 1937-38 1938-39

Fig. 10. Generalized Population Trends of Ruffed Grouse, certain Predators,
and certain Rodents New York State 1930-1939 (Based on State Game Kill
Records and field data)

Turning now to the trends in ruffed grouse in comparison with
those of rodents and predators, there are both correlations and lack
of correlations. We find that the Connecticut Hill trends indicate
that the population drop in 1933 was probably not associated with
predator-buffer trends, the 1935 decline is definitely not so associ-
ated, while the 1937 decline is quite probably linked with these
trends. In examining the state-wide game kill records ^ there appears

^ These records are well-known to be inaccurate; but since the sources of error are
the same each year, tlie trends indicated may be quite reliable.

Interrelationships of Ruffed Grouse 219

to be a possible correlation in the 1936 decline in grouse take, which
coincided with decreases in the take of foxes and gray squirrels and
with the completion of the reduction in take of cottontails that had
begun two years prior ( see Fig. 9 ) .

So many factors enter into these population trends, and they are
of such varying importance, that it would not be expected that the
population curves of so many animals would agree in details. There
is enough evidence from the data, though, to support the interrela-
tion theory among these populations. In Fig. 10 we have tried to
generalize the data in order to illustrate what appears to be the cor-
relation. The rodents lead the parade, and change most violently;
the predators and giouse follow with less marked changes.

OTHER ANIMALS THAT AFFECT THE GROUSE WELFARE

Predators and their usual prey, the buffer species, are not the only
mammals that influence grouse. Many mammals, and birds too,
compete with grouse for food, although this does not ordinarily
cause much trouble ( see page 178 ) . However, those mammals that
graze or browse can have a seriously detrimental effect on the wood-
land cover. Over most of the grouse range, the white-tailed deer is
the only one that becomes abundant enough to damage the range.
The Pennsylvania deer problem, much publicized in the past two
decades, will serve as an example. Leopold (1943) in discussing this
situation says: "Many plants important to other game species were
also depleted; thus greenbrier, on which ruffed grouse depend for
cover, was nearly annihilated. Snowshoe hare and wild turkey like-
wise felt the pressure of excess deer."

PREDATION AS A LIMITING FACTOR

Throughout the discussion of the relations of grouse populations
to other animals, it has been apparent that predation is normally a
constant and immediate medium of reduction of grouse, either as
eggs or birds. The manifestation of predation does not, however,
necessarily measure its status as a limiting factor, or potential limit-
ing factor. However, it does deserve the most careful analysis in
comparison with other major factors. Any group of organisms that
normally destroy two eggs out of every five laid by a prey species,

220 The Ruffed Grouse

and half the birds that reach maturity— and that may, under certain
conditions, far exceed these rates of destruction— cannot be dis-
missed hghtly, even though a part of the losses be conditioned by
other factors.

It may be well for us at this time to review the meaning of "Hmit-
ing factor." It is that element which ultimately prevents any more
of a given species from existing or surviving than actually do. To
clarify the matter let us take a couple of simple, and factual, exam-
ples.

The Lancaster County region of southeastern Pennsylvania is an
intensively farmed area. The vegetation on the land is mainly crops
(com, wheat, oats, tobacco, hay) and pastures. The woodlands are
small, scattered, and lacking in conifers. This range contains no
grouse. Clearly the limiting factor is a lack of suitable habitat, shel-
ter first, and food also but probably to a lesser degree. It is obvious
that predation is not limiting the grouse here, since there are no
grouse for them to capture.

There are cases in the Northeast where good grouse habitat exists
almost literally in the shadow of the tall buildings of large cities.
Unless this habitat is protected as a virtual sanctuary, it will ordi-
narily contain no grouse. Man's interference, as a hunter, or in seek-
ing recreation, or cutting the underbrush for a park, or in other
pursuits, eliminates gi-ouse from this potentially habitable range.

These are two cases where a single element serves definitely and
continuously to act as a limiting factor upon grouse— in both cases
an excluding type of action. Under more normal circumstances, rec-
ognition of the limiting factor is more difficult. Moreover it may
change from one year to the next. It is very likely to change as the
grouse population itself fluctuates. There may be times and places
that predation will act as the ultimate determinant of the level of
grouse population, but these will not be general. They will most
commonly be where the number of birds is well below both carry-
ing capacity and saturation point. There will be times at almost any
density of grouse when weather conditions will act as the major
brake to increase. This may merely condition predation, as in severe
winters, or may act more directly, through chick mortality. When
populations are high, disease may limit the birds or condition an
increase in predation, or the natural intolerance of the birds for their
own kind (each requiring so much territory) may be the factor that

Interrelationships of Ruffed Grouse 221

prevents a higher degree of survival so that further increases are pre-
vented.

We have aheady seen that drastic control of predators may bring
about an increase of grouse when the density is quite low, and that
it fails to have that result when populations are high. When the
number of grouse is more than the envii-onment can safely accommo-
date, decline is inevitable. The conclusion seems to be that predators
may limit grouse populations when below the peak of their upward
trend, but still do not ordinarily prevent them from increasing to a
greater density that permits another limiting factor to bring about
declines beyond expected seasonal losses. These relations of popu-
lation densities to changes in population level are fundamental and
override all others including predator-prey relationship.

REFERENCES AND CITATION SOURCES ON RELATIONS
OF RUFFED GROUSE TO OTHER ANIMALS

Bennett, L. J., and English, P. F. Food Habits of the Grey Fox in Penna.,

Penna. Game News, Vol. XII, No. 12, March, 1942.
Bezdek, F. H. (Progress report on research into methods of increasing ruffed

grouse populations), Pittman-Robertson Quarterly, Vol. 3, No. 4, Oct., 1943.
Cook, D. B., and Hamilton, W. J., Jr. The Ecological Relationships of Red

Fox Food in Eastern New York, Ecology, Vol. 25, No. 1, Jan., 1944.
Dean, R. Unusual Abundance of American Goshawk, Auk, Vol. XXIV, No. 2,

1907.
Dearborn, Ned. Foods of Some Predatory Fur-bearing Animals in Michigan,

Univ. Mich., School of Forestry and Conservation Bull. No. 1, 1932.
Eadie, W. Robert. Food of the Red Fox in Southern New Hampshire, Journ.

Wildlife Mgt., Vol. 7, No. 1, Jan., 1943.
Edminster, F. C. The Effect of Predator Control on Ruffed Grouse Populations

in New York, Journ. Wildlffe Mgt., Vol. 3, No. 4, Oct., 1939.
English, P. F., and Bennett, L. J. Red Fox Food Habits Study in Pennsylvania,

Penna. Game News, Vol. XII, No. 11, Feb., 1942.
Errington, P. L. Food Habits of Mid-west Foxes, Journ. of Mammalogy, Vol. 16,

No. 3, 1935.
Errington, P. L., Hamerstrom, Frances, and Hamerstrom, F. N., Jr. The Great

Homed Owl and Its Prey in North-central United States, Res. Bull. 277,

Iowa State College Agr. & Mech. Arts., 1940.
Fisher, A. K. The Hawks and Owls of the United States in Their Relation to

Agriculture, U.S.D.A. Div. Omith. & Mammal. Bull. No. 3, 1893.
Gerstell, Richard. The Pennsylvania Bounty System, Res. Bull. No. 1, Pa. Game

Commission, 1937.
Gross, A. O. Food of the Snowy Owl, Auk, Vol. 61, No. 1, Jan., 1944.

222 The RuflFed Grouse

Hamilton, W. J., Jr. Seasonal Food of Skunks in New York, Journ. Mammal-
ogy, Vol. 17, No. 3, 1936.
Hamilton, W. J., Jr. The Value of Predatory Animals, N. Y. Zool. Soc. Bull.,

Vol. XL, No. 2, 1937.
Handley, C. O. Progress Report of the Study of the Food Habits of the Red

and Gray Fpx in Virginia, Ann. Rep., Va. Comm. of Game and Inland Fish-
eries, 1934.
King, R. T. Ruffed Grouse Management, Journ. Forestry, Vol. XXXV, No. 6,

June, 1937.
Kozicky, Edward L. Food Habits of Foxes in Wild Turkey Territory, Pa.

Game News, Vol. XIV, No. 4, July, 1943.
Leopold, Aldo. Deer Irruptions, Wise. Cons. Bull., Vol. VIII, No. 8, Aug.,

1943.
McAtee, W. L. Food Habits of the Common Hawks, U.S.D.A. Circular

No. 370, 1935.
McDowell, R. D. The Great Homed Owl, Pa. Game News, Vol. XI, No. 8,

1940.
McDowell, R. D. The Eastern Goshawk in Pennsylvania, Pa. Game News,

Vol. XI, No. 11, 1941.
McDowell, R. D. The Cooper's Hawk in Pennsylvania, Pa. Game News, Vol.

XII, No. 1, 1941.
Mendall, H. L. Food of Hawks and Owls in Maine, Journ. Wildlife Mgt.,

Vol. VIII, No. 3, July, 1944.
. New York State Conservation Department Annual Reports for years

1931, 1932, 1933, and 1934.
Sealander, J. A. Winter Food Habits of Mink in Southern Michigan, Journ.

Wildlife Mgt., Vol. VII, No. 4, Oct., 1943.

8

The Diseases and Parasites of Wild Ruffed Grouse

Most species of vertebrate animals are subject to infection by a
variety of parasites, and the grouse is no exception. The proportion of
grouse playing host to one or more forms of parasites varies with
the seasons, in different regions or localities, and changes from one
year to another. The presence of parasites is not necessarily an ill
omen. How serious the infection is to the bird depends upon the
kind of parasite, the number present, and the destructiveness of the
species.

As disease is a serious factor in the mortality rate of many animals,
it has long been suspected that many grouse may die from this cause.
Particular attention has been directed at the possible connection be-
tween epizootic diseases and population fluctuations, or cycles.
Epizootics among the red grouse ( Lagopus lagopus scoticus) in Eng-
land have been shown to be caused by a nematode worm ( Tricho-
strongijlus pergracilis) and Coccidia (Leslie and Shipley, 1911).
One of the early reports attributing disease as a cause of ruflFed grouse
scarcity was that of WoodruflF ( 1908 ) in New York. Of nine theories
"offered as a possible explanation of the grouse scarcity," three re-
lated to disease: "(5) An epidemic disease ... (6) An internal
parasite. (7) An external parasite ('ticks')." He concluded "that it
was due to an unhappy combination of three separate factors, each
one of which alone was serious in its effects." Following winter
predation and adverse spring weather in 1907, number three is "an
epidemic of some disease or parasite, or both, just which we cannot
now determine."

However, the report on the next die-oflP in New York (Stoddart,
1918) deprecated disease as an important factor. After recalling the
importance that Woodruff attached to disease in the earlier report,
Stoddart says: "There is nothing in Mr. Woodruff's report that is

223

224 The Ruffed Grouse

conclusive on this subject. Those who answered the Commission's
questiomiaire are convinced that these agencies (i.e. diseases and
parasites ) are playing little or no part in the present grouse shortage
—'disease' being placed last on the list of causes . . ."

It was not long though before the matter of grouse disease again
received considerable attention. Following the secondary grouse
decline in 1924, the American Game Association arranged a study
of the grouse which from the start gave primary emphasis to the dis-
ease factor. Led by two eminent ornithologists, A. A. Allen and
A. O. Gross, this study inaugurated an era of research in the field of
wild life ecology and management. In their first report Allen and
Gross ( 1926), after a year's work including examination of nine hun-
dred and twenty-three birds, stated that ". . . no definite conclu-
sions or recommendations can yet be advanced." Nevertheless, con-
siderable progress was made. "The common belief is that there is
one disease responsible for the disappearance of the grouse. The
investigation this year uncovered no evidence to support the belief.
On the contrary over twenty different parasites and diseases were
found, any one of which may prove to be important in different lo-
calities. Some are abundant and of general distribution, others seem
to be of only local importance as yet." Later, evidence indicated that
the stomach worm {Dispharynx spiralis) might be the cause of local
grouse declines. Gross (1930) found that "this parasite was respon-
sible for the death of a large proportion of the birds found dead in
Massachusetts and Connecticut and sent to us . . ." No other dis-
eases or parasites of the wild birds showed any likelihood of causing
serious mortality. When Gross reported again a year later (Gross,
1931) he said that "not a single bird was received (during the past
year) which had met death through diseases or parasites."

Thus was indicated the periodic nature of grouse disease as a
lethal agent. In the early years of the New York study, 1930-32, not
a bird found gave any indication of having serious trouble arising
from disease. Then, in 1933, the first indications of possible trouble
showed up at Connecticut Hill. The author found two grouse dead
and in full flesh which proved to have died from Dispharynx infec-
tion. Soon another bird was found which, while killed and decapi-
tated by a predator, was found to have been severely affected with
aspergillosis, which very likely had weakened it before the predator
attack. The grouse population was high and the incidence of para-
sitic infections was much higher than during the previous year. The

The Diseases and Parasites of Wild Ruffed Grouse 225

bird population declined that winter more than usual, but recov-
ered quickly the next year, and there was no conclusive evidence of
disease being a major decimating agency. And so it has been each
succeeding year since in New York.

During the 1930's the results of three other investigations on the
grouse disease factor were reported. Working in Ontario, Canada,
Clarke (1936) concluded that a protozoan blood parasite, Leuco-
cytozoon bonasae ". . . is most probably the organism responsible
for the dying-off of grouse." Boughton ( 1937 ) reported on examina-
tions of five hundred and sixty grouse, mainly from Minnesota, and
was unable to draw definite conclusions as to the part played by
disease in causing grouse mortality. He did feel that ". . . none of
the parasites found during the present survey (which does not in-
clude examinations of adult grouse during the months of June, July
and August ) have been directly responsible for the mortality among
the adult ruffed grouse during the year 1933-34, when a decrease
in the grouse population was reported." However, he added that
"the possibility still exists that parasitism is the cause of 'epidemics'
in adult grouse during the summer months ... It appears more
probable . . . , however, that any mortality that did take place
might have occurred in the young grouse . . ." More than three hun-
dred grouse were autopsied in connection with the Michigan grouse
study by Fisher ( 1939 ) . He concluded ". . . that parasites and dis-
eases of the ruffed grouse appear to be a major factor in the decima-
tion of these birds in Michigan. It is evident that in order to account,
if possible, for the periodic fluctuation in the population of the ruffed
grouse in Michigan, it will be necessary to continue laboratory
studies and field observations on these parasites that infest the
species."

It seems apparent that the several investigators each feel that there
is something vital in the grouse's disease relationships but they are
not quite sure what it is or how it works. Before considering further
the epidemiology of these diseases, let us review those that are the
more prevalent.

CHARACTERISTICS AND OCCURRENCE OF DISEASE AGENTS

The diseases and parasitic conditions in the ruffed grouse, and
their causative agents, may conveniently be grouped in the follow-
ing categories:

226 The Ruffed Grouse

Helminths, which are internal, parasitic worms, divided into three
classes, the roundworms (Nematodes), tapeworms (Cestodes), and
the flukes ( Trematodes ) .

Protozoa, microscopic, single-celled animals.

Bacteria, viruses, and fungi.

Ectoparasites, including flies, lice, ticks, fleas, and mites.

The Nematode Worms (roundworms). At least fourteen species of
roundworms have been found in the ruffed grouse, of which nine
are known to occur in the eastern seaboard states. In the listing be-
low, those of most significance are placed first.

Dispharynx spiralis (Molin, 1858) Skrjabin 1916 b. This stomach
worm (see Plate 35) attacks the biid's proventriculus or glandular
stomach, but is also occasionally found in the gullet and the intes-
tine. It affects a number of hosts in addition to the grouse, includ-
ing the domestic chicken, dove, robin, turkey, and guinea fowl.

It is a small, whitish worm a little less than half an inch long,
which attaches itself by its head to the mucosa of the stomach, and
usually rolls its body in a spiral.^ There is usually a swelling of the
organ, the degree depending upon the severity of the infection.
There is a small, bloody lesion of the epithelial tissue, which spreads
to redden the entire mucous surface ff many worms are present. In
such cases the lumen ( passage ) is reduced by swelling of the walls.
In the most serious cases, the worms penetrate and destroy the gas-
tric glands. This is accompanied by an excessive secretion of a white,
glary mucus. In some cases the proventriculus is perforated, result-
ing in a fatal peritonitis.

In severe infections, the digestion is severely impaired and emacia-
tion results. Death commonly results from the primary tissue de-
struction and starvation. Most of the infections are produced by a
few worms. AUen and Gross (1926) found the usual infection to be
"from three or four up to a dozen." However, they recorded one case
of two hundred and twenty-eight parasites. Evidence indicates that
infections with twenty to thirty worms are suflBcient to affect the
bird's health and those more severe may prove fatal.

The Iffe history of the parasite begins with the eggs passing out
of the grouse in its droppings. These are eaten by sowbugs (Porcellio
laevis) which are the intermediate hosts. Here the embryos escape

1 See Cram ( 1927) for full description.

The Diseases and Parasites of Wild Ruffed Grouse 227

and develop into infective larvae in the body cavity. The sowbugs
are then eaten by grouse (or other primary host species). In the
bird's digestive tract, the larvae mature and attach themselves to
the proventricular wall. When mature, they lay eggs free in the
digestive tract to be carried out in the droppings, thus completing
the round.

The occurrence of Dispharynx is quite local, and varies greatly
in different years. So far as is now known, it has been a serious grouse
parasite only in the area from New York and New Jersey through
southern New England. Of four hundred and forty-three birds ex-
amined by Allen and Gross ( 1926 ) from five states, several of which
had been picked up dead, forty-nine (eleven per cent) had Dis-
pharynx infestations. None of these stomach worms were found in
four hundred and four birds from ten other states and provinces.
Of 2,059 adult wild grouse examined during the New York study
(N. Y. S. Cons. Dept. Ann. Reports 1932-42) about three hundred
and fifty contained Dispharynx, an average infection of seventeen
per cent. The rate of infection varied from sLx and five-tenths per
cent in 1933 and seven and seven-tenths per cent in 1932 ( and prob-
ably less in 1931 and 1930 ) to twenty-eight and two-tenths per cent
in 1942. The rate increased generally from 1932 to 1938 when it
reached twenty per cent. Then it declined to six and five-tenths per
cent in 1939, whence it again increased each year until 1942. Serious
cases were not uncommon, as in 1934 when eight out of eighteen
cases "were thought serious enough to have caused death in due
time."

The parasite occurs in young grouse too. The New York State
Conservation Department Annual Reports from 1933 to 1941 (ex-
cept 1936 ) summarize the examinations of nine hundred and thirty-
two chicks, of which fifty-four or five and eight-tenths per cent had
Dispharynx. The annual rates of infection varied from zero (1933
and 1934) to ten and two-tenths per cent in 1940.

Since these birds were collected from various parts of the state
and at all seasons ( chicks only in summer, of course ) , the probability
of obtaining a true estimate of the rate of infection from the number
of cases is slight. However, there is some indication, particularly as
indicated from the records of the adults, that the incidence of this
parasitism fluctuates more or less periodically, building up to a large
number of cases, then falling off, and so on. Further, the high in-

228 The Ruffed Grouse

cidence reached in 1937 coincides with a minor decline in grouse
populations. How much significance this may have is conjectural.
Except for the records in New York and New England, the only
instance of Dispharynx in grouse is in a single bird recorded by
Fisher (1939) from Michigan. It was very heavilv infected. In do-
mestic fowl this parasite is known from many parts of the world.

Ascaridia honasae (Wehr, 1940). This is an intestinal worm that
may be found almost anywhere in the alimentary tract, although
primarily in the small gut, and occasionally free in the body cavity.
Closely related species are parasites of the domestic duck and
chicken.

The parasite is one of the largest infecting the grouse, varying
from about two to four inches in length; it is yellowish in color.^
Ordinarily it does not affect the host seriously and no gross lesions
are produced in most of the cases. The life history probably parallels
that of a close relative in the chicken. Infection is acquired directly,
no intermediate host being necessary. Its primary effect is to deprive
the host of nutrients from the digested food in the intestine.

Infections often are high, two hundred or more worms occasion-
ally being found in a single bird. Such numbers do not seem to
affect the grouse seriously although it has been observed that in
domestic chickens there may be a large enough number of worms
to prevent food from moving through the intestine. If this does occur
in grouse, it has not been obsei*ved and surely is rare.

Allen and Gross (1926) found from one-quarter to one-third of
all birds they examined from various states to be affected with from
one to fifteen worms per bird. Clarke (1936) found this species to
be the commonest endoparasite of Ontario grouse, with incidence
ranging from zero to forty per cent in different areas and averaging
twenty-one per cent. Bough ton (1937) records an incidence of
thirty-seven per cent in five hundred and tl^irt^'-three Minnesota
grouse, with an average number of six worms per infected bird."

Erickson (1944) found eleven of twenty-six birds from Minnesota
to contain this parasite. Fisher (1939) gives the rate of occurrence
in Michigan grouse as from twenty to thirty-seven per cent in differ-
ent years, with the number of worms averaging one to three.

^ See Cram ( 1927) for complete description.

" Boughton lii>ts the species as A. galli in the text although he gives it the name
A. Uneata in the first iiistence on puge 7

The Diseases and Parasites of Wild Rutfed Grouse 229

The records of the autopsies performed by the New York study
from 1932 to 1942 on 2,059 adult grouse showed a great variation
in incidence of Ascaridia, ranging from nine to seventy-one per cent
in different years and averaging forty-one per cent. The incidence of
twenty-five per cent in 1932 ( and probably less in the preceding two
or more years) increased to the highest rate in 1935 and then de-
creased steadily each year to the lowest rate in 1942. This steady
trend may have some significance as an indicator of population re-
lationships.

These worms are commonly found in grouse chicks too, but are not
nearly as prevalent in them as in the adults. The average incidence
in nine hundred and thirty-two chicks reported by the New York
study in their annual reports from 1933 to 1941 was ten and four-
tenths per cent, with variations of from zero to twenty-five per cent
in different years. The highest incidence of infection with this para-
site in the chicks occurred in the same year that it reached a high in
the adults, a year of high grouse density ( 1935 ) .

Thus, while its distribution is general and its incidence high,
Ascaridia is not normally a parasite of much consequence. However,
as several authors have indicated, it might conceivably cause serious
trouble, and may be one of the numerous parasitisms of the grouse
that are associated with high population densities and may con-
tribute to their decimation.

Hcterakis honasae (Cram, 1927). This parasite infects the caeca, or
blind gut, that forms an appendage to the intestine. It is a little less
than half an inch in length. Its development is direct, eggs dis-
charged from the host embryonating and infecting other birds.

The occurrence of this worm has been recorded most commonly
from New York and the species found here and named H. honasae
is somewhat different from the species, H. gallinae, reported as com-
mon to many domestic and wild gallinaceous birds and certain
others throughout the world.

It was found in one bird from Rhode Island and one from Nova
Scotia among four hundred and seventeen examined by Dr. E. E.
Tyzzer and in ten per cent of the birds from New York and fifty
per cent of those from Pennsylvania studied at Ithaca (Allen and
Gross, 1926). Neither Fisher nor Clarke reported any occurrences
from their studies but Boughton (1937) reported three from among

230 The Ruffed Grouse

five hundred and thirty-three birds (one to tliree worms each); he
identified the species as H. gallinae.

In the adult birds examined during the New York study, six hun-
dred and thirty-two or thirty-one per cent had this parasite present.
The incidence varied from zero to seventy-three per cent in different
years, with 1935 to 1937 being the high period. This worm was
found in only four of nine hundred and thirty-two chicks examined
over a nine-year period.

There is no indication that this parasite is of serious consequence
to the ruffed grouse.

Cheilospirura spinosa (Cram, 1927'). This is a slender, whitish
worm that inhabits the gizzard between the muscular walls and
chitinous lining. Its length varies from about three-quarters to one
and one-half inches, and the body usually assumes a twisted shape.
Its life history involves an intennediate stage in a grasshopper
(Cram, 1931).

Its distribution appears to be quite general in the range of the
ruffed grouse. First discovered in 1925 by Stafseth and Kotlan in
Michigan specimens, it was found commonly in the birds examined
by Allen and Gross ( 1926) except in those from New England. They
reported the highest incidence of infection from Michigan birds
( forty-two per cent ) and twenty-one per cent of those from Minne-
sota, twenty-two per cent of Pennsylvania grouse, nineteen per cent
of Wisconsin specimens, fourteen per cent of those from New York,
and three out of eight birds examined from New Jersey, revealed
them. The average number of worms was about twelve to sixteen
but in some cases varied up to forty. Only in the most severe in-
fections was any damage apparent— an abnormal thickening of the
muscular wall, and even this seemed to cause little damage.

Gross ( 1930 ) reported that "This parasite was found to be com-
mon, particularly in specimens received from New England."
Mueller ( 1941 ) again recorded its occurrence in New Hampshire.
Clarke's ( 1936 ) records from Ontario showed an average infection
of Cheilospirura of nine and two-tenths per cent, with variations at
different locations of from none to fourteen and eight-tenths per
cent. He found a few cases severe enough to cause serious lesions
but noted that in these instances the bird's vitality must have been

^ See Cram, 1927, for complete description.

The Diseases and Parasites of Wild Ruffed Grouse 231

impaired. His records showed no pronounced variation from year
to year and he concluded that the parasite has no relation whatever
with the cyclic dying-off.

Boughton ( 1937 ) recorded an infection rate of twenty-three per
cent in Minnesota grouse with an average infection of three and
eight-tenths worms and a maximum of thirty. Erickson (1944) re-
ported four instances among twenty-six birds also taken in Minne-
sota. Boughton included this parasite among those that are poten-
tially dangerous to the ruffed grouse but found no very serious cases
among those examined. The grouse from Michigan reported upon by
Fisher ( 1939 ) were more heavily infected than any recorded, except
the Michigan specimens earlier examined by Allen and Gross. In the
four years from 1933 to 1936 the rate of infection varied from
twenty-eight per cent to fifty-one per cent. Fisher indicates that the
species may be quite significant, for not only was it the most preva-
lent parasite but "many of the grouse were so heavily infected with
these parasites that part of the lining of the gizzard had been de-
stroyed and there was destruction of the surrounding tissue."

The autopsies of the New York study over the eleven-year period,
1932-1942, show it to have very minor importance in that state.
The average infection rate for adult birds was eight per cent and for
chicks two and six-tenths per cent. The annual extremes were two
per cent (1932) to sixteen per cent (1941) for the adults and zero
( 1933, 38, 39) to four and five-tenths per cent ( 1941 ) for the young
birds. This parasite cannot be considered to be of significance to
grouse in New York on the basis of present evidence, but may be so
locally, and possibly quite generally, in Michigan in some years.

Capillaria annulata (Molin). A haii'-thin worm found beneath the
epithelial lining in the bird's crop and gullet, this parasite is about
two to three inches long. It can do serious damage by causing the
walls of the crop and gullet to thicken. An anemia in a grouse para-
sitized by C. annulata was described by Allen and Gross (1926).

The specimens examined by these authors did not reveal this
species commonly for they say "Fortunately these are apparently
comparatively rare parasites, only five wild birds, all from southern
New England and Columbia County, New York, were found with
them."

The species was not common in the birds examined during the

232 The Ruffed Grouse

New York study in most years, but did reach a high incidence from
1935 to 1937. From an occasional case in the years up to 1934, the
incidence rose to twenty per cent in 1935 and to forty-nine per cent
in 1936. Then the rate declined to thirty-four per cent in 1937 and
to unimportant numbers thereafter, with no cases at all recorded
from 1940 to 1942. In the chicks, it is not common. Only in 1935,
when six cases were found in one himdred and twenty-two speci-
mens, did it make its appearance in more than a single bird.

Except for these New York and New England records, the only
other record is a single bird from Michigan ( Fisher, 1939 ) . It is an-
other one of the parasites having a limited range and is not normally
a serious threat to its host.

Oxyspirura petrowi (Skrjabin, 1929). An eyeworm that is found
mider the nictitating membrane, this parasite has been reported
only from Michigan (Fisher, 1939). Here it seems to be a rather
prevalent and possibly serious pest. In the three years, 1934 to 1936,
the incidence in ruffed grouse examined was twenty-eight, twenty-
two and five-tenths and fifteen per cent respectively. "As many as
seventeen of these worms have been taken from one eye, and a
number of birds developed marked cases of conjunctivitis" ( op. cit. ) .
Another species of eyeworm, Oxyspirura mansoni ( Cobbold, 1879 ) ,
Ransom, 1904, was found in three out of one hundred and fifty-five
grouse examined by Boughton (1937), there being one worm found
in each case. No significance was attached to their presence.

Microfilaria. These blood parasites have been found in a number
of grouse, both old and young, by the New York study ( N. Y. S. Cons.
Dept. Ann. Reports, 1933-39). Their numbers are not significant and
the incidence only reached eleven per cent one year (1936), seven
per cent in 1933 and 1934, and less in other years. Only two cases
were found in the chicks. Clarke (1936) reports one occurrence of
these parasites in a ruffed grouse from Ontario. None of these in-
stances indicated any serious pathological trouble.

Physaloptera larvae (Rudolphi, 1819). These larval nematodes
have been reported by Boughton (1937) from Minnesota and Wis-
consin grouse. They are described as, ". . . two to three millimeters
long . . . encysted near the surface of the muscles of the breast
and in the muscles of the legs. The cysts themselves usually were

The Diseases and Parasites of Wild Ruffed Grouse 233

yellowish in color, measuring about two millimeters long. The num-
ber found in an individual varied from one to many dozens." Only
the larvae were found, and the primary host is not known.

Boughton found them present in three and twenty-five hundredths
per cent of the birds examined from Minnesota. However, their
presence was not considered serious to the birds for, when com-
paring grouse weights to analyze the effects of parasitism, those
having only Physaloptera larvae ". . . were considered uninfected."

These parasites have not been reported in grouse farther east than
Wisconsin.

Tetrameres americana ( Cram, 1927 ) . This parasite is found in the
stomach ( proventriculus ) . It was first reported from ruffed grouse
by the New York study in 1936 (N. Y. S. Cons. Dept. Ann. Rep.,
1936) although common in poultry in the South. In 1937, it had ap-
parently increased somewhat in prevalence, being found in about
nine per cent of the adult birds examined. Thereafter its occurrence
fell off. "Of just what significance this parasite is to game birds is
unknown, but since it is a great destroyer of tissue and causes heavy
losses among poultry in the South and has been found in pheasants
and Hungarian partridge also, it is possible it may become impor-
tant" (N. Y. S. Cons. Dept. Ann. Rep., 1937).

Other Nematodes. Three other species of nematode worms have
been found in ruffed grouse in the northeastern quarter of the United
States and eastern Canada. None is common or of much ecological
significance so far as known.

The gapeworm Syngamus trachealis, Montagu (1811), was re-
ported in six adult grouse out of six hundred and sixty-two examined
during the New York study from 1932 to 1935, none thereafter; and
in seven young grouse out of six hundred and eighty-three specimens
between 1933 and 1939, none thereafter. As the name implies this
parasite infects the bird's windpipe, causing it to "gape" or choke
in an effort to get rid of the cause of irritation.

Allen and Gross ( 1926 ) report a single case of an intestinal round
worm of the genus Contracaecum.

A cecal worm, Subulura strongylina (Rudolphi, 1819) Raillet and
Henry, 1912, was found in a single grouse examined by Boughton
(1937). The specimen was from Minnesota but the parasite is

234 The Ruffed Grouse

known to infect a number of other birds elsewhere in the Western
Hemisphere.

The Cestodes (Tapeworms). At least seven species of tapeworms
have been found in the ruffed grouse. Since many of these found in
autopsies made in the course of the New York study have been listed
in published ' records only by genus or simply as "cecal cestoda" or
"tapeworms," it has not been possible to analyze accurately the oc-
ciurence of the different species in each genus found in New York.

Hyinenolepis microps (Diesing)

Hymenolepis carioca (Magalhaes, 1898) (?)

Hymenolepis sp.

Specimens of this genus of tapeworms have been reported from
New England (New Hampshire), New York, Minnesota, and On-
tario. They are found in the small intestine, primarily in the duo-
denum.

This is the only genus of tapeworms identified in the reports of
the New York study. Only a portion of the tapeworms reported were
determined as Hymenolepis sp. but it seems lil^ely that at least the
majority of those identified as "intestinal cestoda," or merely "tape-
worms," were of this group.

The prevalence of these intestinal tapeworms in adult grouse
varied greatly in different years, the incidence varying from zero
to twenty-three per cent, with no clear-cut trends. The average in-
cidence was seven per cent; but none at all were found in four years
out of eleven.

The record is quite different with the chicks in which these para-
sites were found every year and in consistently higher proportions.
The incidence of infection averaged nineteen per cent over a nine-
year period, 1933-41, with extremes of thirty-seven per cent in 1933
(based on only eight specimens, however), thirt)'-two per cent in
1934, and ten and eleven per cent respectively in 1938 and 1939.
The records indicate a consistency of occurrence not usual with
grouse parasites. It is notable that the greatest incidence occurred
in a period when most parasites of grouse were at a low ebb.

Clarke (1936) found nine cases of H. microps in Ontario grouse,
eight of which were in young birds and one from an adult. All of

1 N. Y. S. Cons. Dept. Ann. Rep., 1932-42.

The Diseases and Parasites of Wild Ruffed Grouse 235

the eight youngsters had minor infections and ". . . no apparent
harm had resulted to the birds." The adult, however, was ". . . cap-
tured alive in a weakened condition . . . There was an enormous
abundance of Hymenolepis in the duodenum and the bird would
probably have perished eventually from this cause." Erickson ( 1944)
found a single ". . . cestode fragment, possibly Hymenolepis sp.
in one bird" from Minnesota.

Mueller (1941) reported the occurrence of H. carioca in New
Hampshire grouse, but gave no details.

These parasites are thus occasionally pathogenic but are nor-
mally not a serious source of mortality as far as can now be deter-
mined. Considerable evidence indicates the group to be more preva-
lent in immature grouse than in adults. This may be because of the
young eating more insects, which undoubtedly act as intermediate
hosts.

Davainea tetraoensis ( Fuhrmann, 1919 ) Staf seth and Kotlan, 1925

Davainea pwglottina (Davaine, 1860) Blanchard, 1891

These are almost-microscopic tapeworms that occur in the intes-
tinal tract, especially in the duodenal region. They have been found
in grouse from New England to Minnesota.

Allen and Gross (1926) mentioned two specimens from New
York that contained unidentified species of Davainea. At about the
same time another species was described from Michigan, D. tetra-
oensis. Clarke ( 1936 ) found this in fifteen out of thirty-one young
grouse in Ontario in three out of five locations. At one site twelve
of thirteen chicks were infected. No serious lesions resulted and
Clarke concluded that this, together with its local distribution, ren-
ders it of no importance in connection with the die-off.

Michigan grouse examined by Fisher ( 1939 ) proved to be com-
monly infected with D. tetraoensis. Of seventy-nine birds examined
in 1934, fourteen contained it. Only a single case in eighty autopsies
in 1935 and four out of eighty-six in 1936 completed this record.
No importance was attached to these cases by the author.

Another species, D. proglottijia,^ was reported from three speci-
mens from Minnesota by Boughton (1937), also in three birds from
Minnesota by Erickson ( 1944 ) , and from New Hampshire by Muel-
ler (1941). It too is found in the small intestine and there is no

^ See Boughton ( 1937) for complete description.

236 The Ruffed Grouse

indication that it is a lethal factor. It is also found in the domestic
chicken, and its intermediate hosts are several species of snails.

All authors in reporting on the species of Davainea note the mi-
nute size of these organisms and the likelihood of overlooking light
infestations. Their prevalence may, therefore, be greater than is
shown by the records.

Raillietina tetragona^ (Molin, 1858) Joyeux, 1927. This species of
tapeworm, from two-fifths to ten inches long, occurs primarily in
the small intestine, occasionally in the ceca. It was identified (as
Davainea t.) in three New England grouse by Dr. Tyzzer and re-
corded by Allen and Gross ( 1926 ) . It was found in five out of four-
teen young Minnesota grouse by Boughton (1937), aged from a
few hours to ten days, and was reported from New Hampshire
grouse by Mueller (1941). The species is also known from the
bobwhite and other game birds, and infects a species of snail and
the domestic fly as secondary hosts. No significance is indicated for
this worm as a grouse parasite.

Choanotaenia infundibulum ( Bloch, 1779 ) Cohn, 1899. Another in-
testinal tapeworm, from two to eight inches in length, was reported
by Boughton (1937) as the most prevalent cestode in Minnesota
grouse. The incidence of infection was about three per cent. It is
also found in several other gallinaceous birds. The domestic fly
serves as a secondary host. In spite of its widespread distribution,
it has not been reported elsewhere in grouse.

The Trematodes (Flukes). At least five species of flukes occur in
the ruffed grouse in the northeast quarter of the United States and
eastern Canada. None are knowm to be of any significance as a
lethal factor. These worms are probably more prevalent than is in-
dicated by the records since they commonly escape notice in routine
autopsies, imbedded as they are in the bird's tissues and assuming
the same color. Mueller ( 1941 ) notes that his examination of forty-
six New Hampshire grouse added two new species of trematodes to
the one previously known from this region and suggests this finding
resulted because his birds were taken largely from low ground
where snails, the intermediate hosts, are abundant.

Harmostomum pellucidum ^ Werby, 1928

^ See Boughton ( 1937 ) for complete description.

The Diseases and Parasites of Wild Ruffed Grouse 237

Harmostomum sp.

The fluke, H. pellucidum, is found in the ceca. It is about one-
twentieth to one-tenth inch in length and one-fiftieth inch wide.
Its life history is unknown. Boughton (1937) found it present in
ten Minnesota grouse, or two and eight-hundredths per cent of the
birds examined. The average number of parasites per bird was five
and two-tenths and the maximum eighteen. This fluke has also been
reported from New Hampshire grouse by Mueller (1941).

Another fluke of the genus Harmostomum, species not identified,
was found in four young giouse in two locations in Ontario by
Clarke ( 1936 ) . The parasites were found in the anterior extremity
of the rectum.

Leucochloridium pricei Mcintosh, 1932. This fluke, first described
from spruce grouse taken in Alaska, was first found in the ruffed
grouse by Mueller ( 1941 ) . It was present in over half the grouse
he examined from New Hampshire (forty-six birds), and charac-
terized one of the most frequently encountered parasitisms. The
worms were located "... in the bursa Fabricii, cloaca, and lower
rectum . . . Heavy infections were commoner in young birds, with
as many as a hundred or more worms present in a single host." The
individuals were about one thirty-sixth of an inch long. The author
does not mention any pathogenic effects.

Prosthogonimus mucrorchis Macy, 1934. This parasite was described
from the lake states and had not been reported either outside that
area or at all in ruffed grouse previous to the single case found by
Mueller ( 1941 ) in New Hampshire. It is about one-fifth of an inch
long and lives in the cloaca. The bird from which it was recovered
was a male weighing two hundred and thirty-three grams, indicat-
ing it to be an immature.

Agamodistomum sp. Larval forms of these flukes were found en-
cysted in the subcutaneous tissue of the bird's breast, imbedded in
the chest muscles, or occasionally on the outer surface of the crop
(Boughton, 1937). Its cysts were pearly white and about one-
fiftieth of an inch in diameter. They were in eighty Minnesota
grouse, or sixteen per cent of the birds examined, and were also
found in one sharp-tafled grouse. The known range is from Minne-
sota east to Michigan.

238 The Ruffed Grouse

BrachtjJaemus fiiscattis. This fluke was reported in one grouse from
Minnesota by Erickson (1944).

In the autopsies of Michigan grouse reported by Fisher (1939),
trematodes were found only one year, 1933. Of seventy-three speci-
mens, two and fifty-nine one hundredths per cent contained cysts of
intestinal flukes.

Of aU the hundreds of grouse examined by the New York study
{N. Y. S. Cons. Dept. Atin. Reports, 1932-42) only one bird was
found to have a fluke infection. In 1938, one bird of 69 had "cecal
trematodes." However, it seems quite likely that in these studies
these obscure worms may have been commonly overlooked. In no
instance, however, has their presence given rise to serious pathogenic
effects so far as is known.

The Protozoa. Protozoans are single-celled, microscopic animals,
the parasitic forms of which are found in several parts of the body.
Some of these parasites cause serious disease in gallinaceous birds.
At least nine species are known to infect the ruffed grouse.

Eimeria angusta E. A. Allen, 1934

Eimeria honasae E. A. AUen, 1934

Eimeria dispersa Tyzzer, 1929

These protozoans are known as coccidia, and the disease they
cause is coccidiosis. Allen and Gross (1926) reported one wild
grouse as having a few coccidia but they found these organisms to
be a serious cause of loss in young grouse in captivity. A few years
later Tyzzer (1929) described E. dispersa from wild ruffed grouse.
This species is found in the small intestine. It ". . . is differentiated
from any other species of coccidia in gallinaceous birds by the ab-
sence of any well-defined polar inclusions in the oocyst" ( Boughton,
1937). This author reports finding it in twenty-four Minnesota
ruffed grouse, six and five-tentlis per cent of those examined.

Two new species of coccidia in ruffed grouse were described by
E. A. Allen (1934), as E. angusta and E. honasae. The former lives
in the ceca and has been recovered from grouse taken in Labrador,
Alaska, and Minnesota. Allen says that "quite a number of the
birds examined had heavy infections . . ." Boughton ( 1937 ) found
it present in four ruffed grouse, or one and one-tenth per cent of
those examined and in one sharp-tailed grouse.

The Diseases and Parasites of Wild Ruffed Grouse 239

E. honasae occurs in the ceca, or rarely in the small intestine. It
was reported by Allen (1934) from birds taken in Massachusetts,
Quebec, Labrador, and Alaska. Few of the birds examined were
found to harbor this species.

Coccidia were found in seventy-nine, or three and eight-tenths per
cent, of the adult grouse examined during the New York study, as
noted in their annual reports, but all were found between 1932 and
1937, none from 1938 to 1942. The highest rate of infection was ten
and seven-tenths per cent in 1936, but this figure may have been
affected by inclusion of some captive birds. Of the young grouse
examined, forty, or four and three-tenths per cent, had coccidia.
Again the record is erratic, varying from zero (three years out of
eight ) to seventeen per cent in 1934 and thirty-seven and five-tenths
per cent in 1932 ( three out of eight birds ) .

Infections of coccidia in wild grouse have not been proved to be
of significance as a mortality factor. However, the possibilities of
this disease assuming importance at times are real, especially among
the young birds. Boughton (1937) concluded that it ". . . is un-
doubtedly a dangerous parasite. It is an intracellular parasite, de-
veloping chiefly in the epithelial cells of the small intestine, and
when present in large numbers may give rise to acute enteritis, caus-
ing denudation of the intestinal epithelium, consequent digestive
derangements and malnutrition, and the bird becomes emaciated
and anemic." Coccidiosis has caused considerable trouble to grouse
in captivity (Allen and Gross, 1926; Bump, 1935) although it does
not necessarily follow that similar infections can occur in wild
populations. The most that can be said is that these organisms are
potentially dangerous to wild grouse, but are not yet proved to
be so.

Leucocytozoon honasae Clarke, 1935. This species is a blood para-
site that causes a malaria-like disease. It is found in both adults
and young but seems to be particularly pathogenic to the young.
So far it has been found and identified only from Ontario, Minne-
sota, and Michigan.

So impressively did its occurrence correlate with the dying-off
of grouse in Ontario in 1933 and 1934 that Clarke concluded it
". . . is most probably the organism responsible for the dying-off
of grouse." It is quite possible that epizootics of this disease might

240 The Ruffed Grouse

have caused local dying-off in the areas studied, but it is certain
that they are not responsible for grouse declines in a broad geo-
graphical sense. As species of this genus are known to be peculiarly
pathogenic to the young of turkeys, ducks, geese, and ostriches
". . . this fact, together with the high incidence of the disease,
especially in the single case of ... (a) sick bird ... (he men-
tions ) . . . and the lack of evidence of any other disease-producing
organism or parasite . . ." ( Clarke, 1936 ) led him to conclude that
it was the organism responsible for sudden grouse declines. Even
the assumption that the cause of grouse dying-off is some pathogenic
organism is not known to be valid, even in restricted areas.

Regardless of any cyclic significance this disease might have,
Clarke did find it in twenty-one out of twenty-four specimens exam-
ined for blood parasites ^ in the areas of dying-off in 1934. The previ-
ous year, the gametocytes of the Leucoctozoon were found in four
of five grouse ". . . in which they might be expected." One of these
was a sick youngster, half underweight, which apparently was a re-
sult of this disease. No other serious cases were noted.

The only other areas from which this organism has been reported
are Michigan and Minnesota. Fisher (1939) records that five
grouse out of seventy-nine examined in 1934 from Michigan showed
its presence in blood samples. In July, 1935, two five-weeks-old
chicks out of seven collected by Dr. O'Roke contained the parasite
(Fisher, 1939). Erickson (1944) found the parasite prevalent in
Minnesota. The incidence among 30 specimens in 1941 was sixty-
three and three-tenths per cent; of 48 specimens examined in 1942,
sixty and four-tenths per cent were infected; and of twenty-four
grouse taken in 1943, eighty-three and three-tenths per cent had
these protozoons present. These birds were collected from Sep-
tember to November.

Subsequent to Clarke's pronouncement on the importance of this
protozoan in 1936, others conducted a diligent search for it in other
areas. None at all was found by the New York study.

The Flagellates

Trypanosoma gallinarum Bruce et al.

Trypanosoma sp.

^ Examination for blood parasites requires the preparation of fresh blood smears in
the field at the time the bird is killed.

The Diseases and Parasites of Wild Ruffed Grouse 241
Ptychostoma bonasae Tyzzer, 1930

Cyathosoina striatum Tyzzer, 1930

These tliree (possibly four) species of flagellates have been de-
scribed from the ruffed grouse. In no instance have they shown any
indication that they might be seriously pathogenic. They apparently
are a normal part of the bird's internal fauna and are not associ-
ated with any disease. They were found in a large proportion of
the birds examined by Dr. Tyzzer and reported upon by Allen and
Gross ( 1926 ) . These flagellates they mentioned were probably the
species later described by Tyzzer ( 1930 ) . Clarke ( 1936 ) reported
four birds from Ontario lightly infected with a flagellate tentatively
identified as Trypanosoma gallinarum. He noted that the organism
is probably more prevalent than would be indicated by the small
number of cases, a suggestion borne out by the prevalence reported
by Tyzzer, yet passing unnoticed in several other reports on grouse
parasites. Fisher (1939) reported one grouse from Michigan con-
taining flagellates of the genus Trypanosoma, identified by Stafseth
and Kotlan.

Tyzzer ( 1930 ) found the two parasites he described as prevailing
in the ceca, over the surface of the mucosa, and occasionally found
in the contents of the large intestine. It appeared that of the two,
Ptychostoma was preponderant near the outlet of the cecum while
Cyathosoma occurred in the dilated portions. He noted that they
may be in ". . . the nature of symbionts essential to the health of
the mature bird . . ." but this is merely a conjecture.

Histomonas m^eleagridis ( Smith, 1895 ) Tyzzer, 1920. This protozoan
is the organism that causes the disease known as "blackhead," a
scourge of the domestic turkey. It is a common cause of death of
grouse in captivity (Allen and Gross, 1926; N. Y. S. Cons. Dept. Ann.
Reports, 1933-42) but has not definitely been identified in wild
grouse. Even though this chapter intends only to cover the diseases
of wild grouse, this organism is included because of the great proba-
bility that it actually does affect the wfld birds when they come into
contact with poultry range. The action of the disease is so rapid that
infected specimens in the wild would be exceedingly difficult to
obtain.

Any poultry range is apt to be infected with blackhead even
though the chickens or turkeys themselves do not show signs of

242 The Ruffed Grouse

the disease. Chickens especially are likely to be carriers. Any grouse
walking over infected range may pick up the organisms from the
ground. Being exceedingly susceptible, the bird would die in a
short time.

The organism first affects the ceca and later the liver, causing
destruction of the liver parenchyma. According to Allen and Gross
(1926) "One or both ceca gradually become filled with cheesy,
necrotic material which is more or less bloodshot. Sometimes the
young grouse and occasionally the old birds die at this stage be-
fore the typical lesions appear on the liver. These lesions on the
liver appear as dark depressed areas with a central light area and an
outer light fringe, at first appearing as watermarks but later the
white areas giow until the dark portion is entirely obliterated. There
is usually also a thick straw-colored fluid in the loose tissue about
the heart and sometimes in the peritoneum covering the viscera."

Again it should be emphasized that this very serious disease has
not yet been proved to occur in wild grouse, but the circumstances
indicate that its occurrence is likely where grouse range borders
the farm poultry yard. Any statement linking this disease with
grouse declines would be purely conjectural.

Trichomonas, species. A protozoan parasite reported by Allen and
Gross ( 1926 ) as "occasionally found in small numbers by Dr. Tyz-
zer." No significance was attached to its presence, nor any details
given concerning its status as a grouse parasite.

Infectious Diseases. The grouse is occasionally subject to infectious
disease of several sorts. The disease of captive grouse called "black-
head" ( and likely to occur in wild birds ) is caused by a protozoan
parasite and has been discussed above. Several others known to
occur in grouse are covered below. All are capable of causing death
and some, as tularemia, blackhead, and aspergillosis, are potentially
agents of serious epizootics.

Aspergillosis. This is a disease of the lungs and air sacs caused by
a fungus, Aspergillus fumigatus. Its spores generally are distributed
in the soff and in such material as moldy straw. The conditions re-
quired for its growth in the bird are not well understood.

It has been most prevalent in captive grouse (Allen and Gross,

The Diseases and Parasites of Wild Ruffed Grouse 243

1926; N. Y. S. Cons. Dept. Ann. Rep., 1933-42), but is also known
to occur in wild individuals. Dr. Tyzzer found it in two such speci-
mens in 1925 (Allen and Gross, 1926). The New York study re-
ported twenty-seven cases in adult wild grouse from 1,816 specimens
examined from 1935 to 1942 {N. Y. S. Cons. Dept. Ann. Rep. 1935-
42). No cases were listed among birds examined from 1932 to 1934,
although one bird found dead by the author on Connecticut Hill,
N. Y., in 1933 after decapitation by a predator, proved to have suf-
fered from aspergillosis.

Tularemia. This disease is caused by a bacterial organism, Pasteurella
tuhrense. It has been found in wild ruffed grouse only in the Mid-
west, primarily in Minnesota. Green and Shillinger (1934) reported
one case among eighty-seven birds (including some sharp-tails)
collected in northern Minnesota in 1932; two instances were found
among seventy-one grouse taken in September 1933 (again the
total included some sharp-tails); in October 1933 three cases of
tularemia were noted in ruffed grouse from among twenty-six grouse
of three species. They concluded ". . . that tularemia is a common
disease of both ruffed and sharp-tailed grouse." They could not
"... however, give any decisive judgment as to the significance
of the occurrence of tularemia in grouse as a mortality factor." These
birds were all collected for the study by game wardens, were all
able to fly, and showed no symptoms of disease, with a single ex-
ception. One of the infected birds was found in a dying condition.
However, it was noted that there was little opportunity to locate
sick or dead birds, hence the disease might well have been more
prevalent than indicated.

The tularemia organism is spread by its alternate host, the tick
( Haemaphy salts cinnaharina ) , and probably by other species of the
same genus. These ticks infest numerous other birds and mammals
and in some are known to spread tularemia, notably in the snow-
shoe hare. The number of ticks per animal varies greatly both by
season and in different years, and may well be correlated with
population fluctuations. As tick numbers rise, the disease increases
until finally an epizootic occurs. This reduces the host population
and with it the disease, and from there the cycle builds again.

This hypothesis is still just that— a theory so far as grouse are con-

244 The Ruffed Grouse

cerned. Despite diligent search, no tularemia has been found in
eastern grouse. The true significance of this disease in ruffed grouse
therefore remains unsolved. At least, it is potentially destructive.

Ulcerative Enteritis. This infection of the posterior part of the small
intestine has been observed in captive ruffed grouse and bobwhite
quail for several decades. Commonly called "grouse disease" or
"quail disease," it caused great mortality in numerous bird-rearing
ventures. Levine (1932) described an epizootic in captive ruffed
grouse and showed that the causative organism was transmitted on
ground infected by the birds' droppings. Morley and Wetmore
(1936) proved that the organism was bacterial, similar to that
which causes human diphtheria, and named it Corynehacterium
perdicium. Bass (1941) later identified the organism in quail as a
". . . gram-negative, anaerobic bacillus . . . ," transmittable by
coprophagy. He was able to immunize quail with killed cultures of
the specific organism.

It was suspected that a disease that would infect grouse so rapidly
as ulcerative enteritis, would likely occur in wild birds, possibly be-
coming a serious mortality factor. Green and Shillinger (1934)
found it among wild grouse in Minnesota, and noted that "Occurring
as a natural infection, it stands out as a disease which may play a
devastating role in the periodic destruction of ruffed grouse at peak
abundance." They reported three cases of the disease, one of the
birds being greatly emaciated and weighing only 210 grams.

For the present, we must list this disease along with tularemia
as a potentially destructive force, but not yet demonstrated to be
effective in wild giouse populations.

Other Diseases. Wild ruffed grouse are occasional victims of vari-
ous other diseases. A number of these are secondary afflictions fol-
lowing wounds from accidents or gunshot. Among these may be
enteritis, peritonitis, pneumonia, hepatitis, ventriculitis, ruptures,
tumors, lead poisoning, septicemia, and so forth; all of these have
been reported by the New York study (iV. Y. S. Cons. Dept. Ann.
Rep., 19S2-41 ) . Other abnormalities reported as a result of that in-
vestigation include tuberculosis ( three cases in 1937-38 ) , cloacal in-
fection, egg-binding, and malnutrition.

Allen and Gross ( 1926 ) recorded two cases of bird pox in grouse
from Massachusetts. They described it as causing warty growths

The Diseases and Parasites of Wild Ruffed Grouse 245

about the head, mouth, and tongue. Recovery probably takes place
unless feeding is so interfered with as to result in starvation.

Green and Shillinger ( 1936 ) have reported the occurrence of the
bacterium, Fasteurella aviseptica, in wild grouse. This organism is
the cause of fowl cholera in chickens. The discoverers note that "if
this organism is as destructive in grouse as it is in domestic fowl, it
may well be an important factor in grouse mortality." No evidence
of its widespread occurrence has as yet been demonstrated.

External Parasites. Ruffed grouse are known to be infested by five
different types of ectoparasites, creatures which live on the body
surface and mostly derive their sustenance from biting or pierc-
ing the bird's skin to take its blood. An exception are the feather
mites which live on the feather substance. The other four types are
ticks, fleas, lice, and flies. It is also likely that grouse are attacked
by "bite-and-run" insects such as mosquitoes that remain on the
body only long enough to take one load of blood.

These external parasites are not normally a serious threat to the
bird's health. However, some carry other diseases which may be
serious (see under tularemia, p. 243), and occasionally the infesta-
tion may be so heavy as to be a serious drain on the vitality. In these
instances it is commonly found that the bird has been weakened by
other afflictions as well as the ectoparasites.

Ticks ( Ixodoidea ) . Four species of ticks of the genus Haemaphy-
salis are known to infest ruffed grouse. They are found on the bird
in aU three stages: larva, nymph, and adult. The degree of preva-
lence decreases with each older stage, the larvae being most abun-
dant, n\Tnphs next, and the adults relatively scarce. The reason for
this is not wholly clear but is probably due to reduction in numbers
of ticks with each stage from mortality and progressively increased
difficulty in locating hosts. Ticks drop off a grouse quickly upon the
bird's death. The adults probably leave most quickly, due to greater
agflity, and this helps to explain the difference in numbers of each
stage of ticks found on birds freshly killed.

They are foimd on the host from spring to fall, and pass the winter
in the ground in the egg stage. Abundance seems to be greatest in
late summer. Further, there is apparently a marked geographic vari-
ation in abundance, the ticks being most prevalent in midwestem
lake states, less in the East.

246 The Ruffed Grouse

Probably the most prevalent species is H. cinnabarina Koch
(1844), the common rabbit tick, found on numerous mammals,
gallinaceous biids, and man. It is a carrier of the dread disease, tula-
remia. Its range is wide and it is known to infest giouse in the east-
em United States and Canada from Minnesota and Alberta to New
England and Labrador.

Another species attacking grouse over a wide range is H. leporis-
palustris. Gross (1930) reported it ". . . present in considerable
numbers on most of the biids collected and examined at Matamek
(in Labrador, eastern Canada) in 1929." Fisher (1939) recorded
1,172 specimens taken from thirty ruffed grouse from Michigan and
examined by Dr. R. G. Green of Minnesota. The average number of
ticks per grouse was one hundred and sixty-seven. This species also
carries tularemia. Fisher also reported one grouse from Michigan
examined by Dr. H. J. Stafseth that had the tick H. punctata punc-
tata, a species found on a number of mammals and birds. The fourth
species, H. chordeilis, was reported from New York in 1941 (IV. Y. S.
Cons. Dept. Ann. Rep., 1941).

The heaviest infestations of ticks have been reported by Green
from Minnesota. The average number on all Minnesota grouse ex-
amined during September was six hundred and forty and in October
it was one hundred and eighty (Green and Shillinger, 1934). The
heaviest infestations were on two birds that yielded 2,985 and 2,468
ticks, respectively. The authors believed that the average figures
were lower than the actual owing to ticks escaping before the birds
were picked up and sealed in bags.

Fisher (1939) quoted Green on the examinations of Michigan
grouse and their ticks: "Nineteen samples of a hundred ticks each
were injected into a corresponding number of guinea pigs. Two of
the nineteen guinea pigs died with lesions typical of tularemia,
showing that the samples of ticks obtained from grouse specimens
were infected with tularemia to the extent of ten per cent. This fig-
ure is probably to be considered a normal finding and does not
represent an unusual situation for the present stage of the game
cycle, as similar findings have been obtained in Minnesota."

Infestation of ticks varies greatly in different years and probably
follows a rather well-marked cycle. However, there is often marked
variation in nearby localities at any given time. Numbers of ticks
present are, of course, correlated with grouse abundance, there be-

The Diseases and Parasites of Wild Ruffed Grouse 247

ing an increase in the rate of infestation and in the degree of infesta-
tion as the birds become more numerous. The correlation is also
connected with the abundance of other tick host animals, notably
hares and rabbits.

Clarke ( 1936 ) said that "Ticks were found on most of the speci-
mens collected ..." in Ontario. Allen and Gross (1926) recorded
heavy infestations on eight of fifteen birds taken in northern Maine,
and stated that ticks ". . . were likewise reported on birds in Al-
berta, Canada."

Ticks have been found more commonly on the chicks than on the
adults in the New York study {N. Y. S. Cons. Dept. Ann. Reps., 1932-
42). With both age-classes tick occurrence has varied greatly in
different years, and is low judging by Midwest standards. The first
instance was that of a single adult bird in 1933. No further cases of
ticks infesting adults were noted until 1939. Sixteen instances in the
three years, 1939-41, a negligibly low incidence, were all. None was
found in 1942. The first ticks on grouse chicks were twenty-five cases
in 1934, representing an incidence of fifteen per cent. They were
identified as H. leporis-palustris. Next year the incidence dropped
to six per cent (seven cases); then for two years the record was
incomplete, although there were numerous cases of tick infestation
in 1937. No chicks were found infested in 1938, but for the next
three years the incidence increased yearly. From a six per cent (ten
cases ) rate of infestation in 1939, it rose to twenty per cent in 1940
(twenty-eight cases) and twenty-one per cent in 1941 (twenty-four
cases). The next year none was reported. From 1935 to 1940 the
species was called H. cinnaharina; in 1941 it was determined as
H. chordeilis.

In addition to the possibility of transmitting tularemia to grouse,
ticks may cause the birds considerable discomfort, and occasionally
ill health. The ticks tend to congregate about the head and neck and
may cause this area to be almost completely denuded of feathers.
With the young birds especially, the drain of blood as the ticks
engorge themselves may reduce the bird's vitality, and sometimes
result in death.

Louse Flies. A single species of the true flies (Diptera) is of
fairly common occurrence on grouse, both young and adult. It is a
hippoboscid, Lynchia americana ( Leach ) . Allen and Gross ( 1926 )

248 The Ruffed Grouse

commented: ". . . reported rather generally distributed but few
specimens sent in." This may have been because the insects left the
birds quickly after death, and were not caught by the collectors.
Clarke ( 1936 ) reported a single case in Ontario.

Nine scattered instances in eleven years were reported by the
New York study on adult grouse, and twenty-two cases in ten years
on chicks (N. Y. S. Cons. Dept. Ann. Reps., 1932-42). These are
blood-sucking flies, but no serious significance is attached to their
presence on grouse.

Lice ( Mallophaga ) . Mallophagous insects of at least two species
are occasional on grouse. Allen and Gross (1926) reported several
on birds sent in during winter and spring. Clarke ( 1936 ) collected
them "... in numbers from many of the specimens taken at the
field stations." They were identified as Degeeriella camerata (Nitz).

Of seventy-one occurrences of lice on adult grouse reported by
the New York study, all but one were in the period 1935 to 1937.
In 1936 the incidence was thirteen per cent. Only one case was re-
ported for chicks, that in 1935. Wliether this erratic occurrence is
actual, or the result of different methods of analysis is not clear.
It hardly seems that an infestation of this type would be so highly
erratic.

Lice are a source of irritation to grouse but are not known to be
of any significance as a mortality factor.

Fleas (Siphonaptera). Two grouse having fleas were reported by
Clarke ( 1936 ) . One of the parasites was identified as Ceratophyllus
species. No importance was attached to its occurrence.

Mites (Acarina). Feather mites of the family Analgesidae, and
belonging to the genus Megninia, ". . . were found in greatly vary-
ing abundance on most of the specimens examined in the field. On
some young birds they were very numerous, but except as a possible
source of irritation, cannot be considered as of pathological impor-
tance." So reported Clarke (1936) concerning Ontario grouse. He
also recorded larvae of the mite Trombicula microti, of the family
Trombidiidae, on grouse, particularly in Algonquin Park, Ontario.
They were not found in great numbers, and were sporadic in occur-
rence.

Air-sac mites {Cytoleichus nudus) were reported in seventeen

The Diseases and Parasites of Wild Ruffed Grouse 249

grouse by the New York study [N. Y. S. Cons. Dept. Ann. Reps.,
1935-37) in the three years 1935 to 1937. None was found in other
years. No indication was given of any importance attached to these
infestations.

TOTAL PARASITE INFESTATION

In discussing each organism that parasitizes the ruffed grouse, the
conclusion was almost always reached that the species was not a
very serious mortality factor. Most have not been demonstrated to
be lethal even when prevalent and widespread in occurrence. None
of the afflictions known to cause death commonly is widely dis-
tributed. The stomach worm {Dispharynx spiralis) is apparently
restricted to parts of New England and the middle Atlantic states,
the blood parasite (Leucoctjtozoon bonasae) primarily to Ontario,
and tularemia to the north-central states. No one disease-producing
organism has been shown to cause major grouse declines over a wide
range.

Possibly of more significance as an indicator of the importance of
disease as a mortality factor than each parasite by itself is the total
prevalence of all parasites and diseases. It is entirely logical that
there would be a cumulative effect when more than one organism
is attacking an individual bird. The effect of several species could
result in death whereas any one of them alone might not have caused
trouble.

The proportion of grouse having one or more parasites or disease
organisms has varied with a clear-cut trend in New York in the
period 1930-42. The autopsy records are not available to corrobo-
rate the belief that parasitism was at a low ebb in 1930 and 1931
but circumstances indicate it to be true. Beginning with 1932 the
incidence of parasitism increased steadily; forty-eight per cent in
1932; sixty-four per cent in 1933 and sixty-one per cent in 1934; then
ninety-five per cent, ninety-two per cent and ninety-four per cent
in 1935, 1936 and 1937, respectively. Then followed a decline in
parasite incidence beginning with a seventy per cent rate in 1938,
followed by fifty-five per cent in 1939 and forty per cent in 1940.
The next two years indicate another period of increase, the 1941
rate being fifty per cent and that of 1942, sixty per cent.

The periods of increase exactly parallel these of growth in the

250 The Ruffed Grouse

grouse population, and the break in the increase phase, in 1938,
came immediately after a significant reduction in grouse numbers
generally in New York. While these are admittedly rough correla-
tions, they almost surely carry considerable significance.

With the grouse chicks, the trend of parasite incidence is not so
pronounced, but rather is more consistent from year to year. Fur-
ther, though there is much evidence that disease is a more efifective
mortality factor among chicks than with adult grouse, the incidence
of parasitism is notably lower in the young birds. This probably re-
flects the fact that many of the chicks were collected when but a
few days old, a time when few parasites are likely to be found. The
nine-year record of nine hundred and thirty-two autopsies of grouse
chicks from 1933 to 1941 ( 1936 excluded ) shows the incidence of
total parasitism as follows: 1933— thirty-eight per cent; 1934— fifty-
eight per cent; 1935— thirty-four per cent; 1937— forty-four per cent;
1938— seventy-five per cent; 1939— thirty-three per cent; 1940— fifty-
one per cent; 1941— fifty-two per cent. The grand average was forty-
three per cent, compared with an average of seventy-two per cent
for all 2,059 adult birds examined {N. Y. S. Cons. Dept. Ann. Reps.,
1932-42).

DISEASE AS A LIMITING FACTOR

Interpretation of the disease factor is among the more diJQficult of
all the facets of grouse ecology. It is manifestly impossible to gather
the vast amount of factual information that would be necessary to
render a well-substantiated judgment. Not only must the dead and
dying grouse be gathered in considerable numbers during a die-oflF
period, an achievement never attained by any grouse investigation,
but the related factors must be placed in their proper perspective
too. Geographical distribution of disease-caused mortality, and its
direction and spread are important. Correlations with grouse popu-
lations must be made, and hence widespread censuses must have
been made, another impossible task. The part played by secondary
hosts of the parasites, their population, condition and distribution,
the relation of weather (and possibly other meteorological condi-
tions ) to grouse health and to the disease organisms, the part played
by predation and hunting ( or lack of them ) , these and other matters
may be fundamental to an adequate understanding of grouse popu-

The Diseases and Parasites of Wild Ruffed Grouse 251

lation changes. So it is no wonder that we cannot yet paint a very
clear picture of the significance of disease among grouse.

Despite all of the defects in our knowledge, certain conclusions
can be made with fair accuracy and other probabilities can be
suggested.

During most years, disease is not a primary mortality agency of
grouse in any given area. A considerable incidence of most of the
parasitic organisms that aJBFect grouse may be considered normal and
not a serious threat.

The incidence of disease organisms and the intensity of the indi-
vidual infections and infestations are directly related to the popula-
tion density of the grouse. When grouse are scarce, they will have
little disease; as the numbers of grouse increase, parasitism and
disease will increase, and when grouse are abundant, disease will
generally become more virulent and an important cause of mortality
in local areas.

When grouse are abundant over a wide range, losses from disease
will probably occur from a combination of several species of para-
sites; and when epizootics do occur from a single species of parasitic
organism, the eflFects are likely to be very local and unevenly dis-
tributed.

It is probable that there is a significant relationship between
disease mortality in grouse and weather conditions. It is well-known
that some parasites reproduce more abundantly under certain me-
teorological conditions than under others, as for example in wet
weather as contrasted to dry. Boughton ( 1937 ) discussed this matter
at some length. He concluded, "... a definite correlation between
the degree of parasitism and the meteorological and topographical
factors appears to occur."

When disease becomes an important mortality factor of grouse,
it probably is primarily effective in killing the young birds during
the summer.

When serious declines in grouse populations occur over consider-
able areas, declines of the type that have been called cyclic, disease
almost certainly plays a part, and just as certainly shares the re-
sponsibility for causing the losses with other major factors as weather
conditions and predation.

252 The Rufied Grouse

REFERENCES AND CITATION SOURCES FOR DISEASES AND
PARASITES OF THE RUFFED GROUSE

Allen, A. A. The Grouse Disease, Bull. Am. Game Protc. Ass'n, Vol. 13, No. 1,
1924.

. The Grouse Disease, Am. Game, Vol. 14, No. 3, 1925.

Allen, A. A., and Gross, A. O. 1926. Report of the Ruffed Grouse Investiga-
tion, Season of 1925-26, Am. Game, Oct., 1926.

Allen, Ena A. 1934. Eimeria angusta sp. NOV. and Eimeria honasae sp.
NOV. from Grouse, with a Key to the Species of Eimeria in Birds, Trans,
of the Am. Microscopical Soc, Vol. LIII, No. 1, Jan., 1934.

Bass, Gharles C. Specific Cause and Nature of Ulcerative Enteritis of Quail,
Proc. Soc. Exp. Biol, and Med., Vol. 46, No. 2, Feb., 1941.

Boughton, Rex V. 1937. Endoparasitic Infestations in Grouse, Their Patho-
genicity and Correlation With Meteoro-Topographical Conditions, Tech.
Bull. 121, Univ. of Minnesota Agr. Exp. Sta., Aug., 1937.

Clarke, C. H. D. The Dying Off of Ruffed Grouse, Trans. 21st Am. Game
Conf., 1935.

Clarke, C. H. Douglas. 1936. Fluctuations in Numbers of Ruffed Grouse,
University of Toronto Studies, Biological Series No. 41, Univ. of Toronto
Press, 1936.

Cram, Eloise B. 1927. Bird Parasites of the Nematode Suborders Strongylata,
Ascaridata, and Spirurata, U. S. Nat. Mus. Bull. No. 140, 1927.

Cram, Eloise B. 1931. Developmental stages of some Nematodes of the
Spiruroidea parasitic in poultry and game birds, U.S.D.A. Tech. Bull. 227,
1931.

Erickson, A. B. 1944. (Wildlife restoration project progress report), Pittman-
Robertson Quarterly, Vol. 4, No. 2, Apr., 1944.

Fisher, Lee William. 1939. Studies of the Eastern Ruffed Grouse in Michi-
gan, Tech. Bull. 166, Michigan State College Agr. Exp. Sta., June, 1939.

Greene, R. G. Disease in Relation to Game Cycles, Trans. 18th Am. Game
Conf., 1931.

Greene, R. G., and Shillinger, J. E. Relation of Disease to Wildlife Cycles,
Trans. 19th Am. Game Conf., 1932.

Greene, R. G., and Shillinger, J. E. Wildlife Cycles and What They Mean to
the Grouse Supply. Trans. 20th Am. Game Conf., 1934.

Greene, R. G., and Shillinger, J. E. 1934. Progress Report of Wildlife Disease
Studies for 1933, Trans. 20th Am. Game Conf.

Greene, R. G., and Shillinger, J. E. 1936. Progress Report of WildMe Disease
Studies for 1935. Proc. 1st No. Am. Wildlffe Conf.

Gross, A. O. 1930. Annual Report of the New England Ruffed Grouse Investi-
gation, Boston, Mass., Sept., 1930.

Gross, A. O. 1931. Annual Report of the New England Ruffed Grouse In-
vestigation, Boston, Mass., Oct., 1931.

The Diseases and Parasites of Wild Ruffed Grouse 253

Gross, A. O. Diseases of the Ruffed Grouse Investigation, Trans. 19th Am.
Game Conf., 1932.

Leslie, A. S., and Shipley, A. E. 1911. The grouse in health and disease. Lon-
don, 1912.

Levine, P. P. 1932. A Report on an Epidemic Disease in Ruffed Grouse,
Trans. 19th Am. Game Conf.

Morley, L. C., and Wetmore, P. W. 1936. Discovery of the Organism of Ul-
cerative Enteritis, Proc. 1st No. Am. Wildlife Conf.

Mueller, Justus F. 1941. Some Parasites Newly Recorded for the Ruffed
Grouse, Bonasa umbellus, in the United States, Proc. Helminthological Soc.
of Wash., Vol. 8, No. 1, Jan., 1941.

N. Y. S. Cons. Dept. Ann. Reports, 1932 to 1942, inclusive.

Stoddart, A. M. 1918. Ruffed Grouse in New York State, Albany, N. Y.

Tyzzer, Ernest E. 1930. Flaggelates from the Ruffed Grouse, Am. Journ.
Hygiene, Vol. XI, Jan., 1930, No. 1.

. 1929. Coccidiosis in gallinaceous birds, Amer. Journ. Hygiene, Vol.

10:269-383.

Wehr, Everett E. 1940. A New Intestinal Roundworm from the Ruffed Grouse
{Bonasa umbellus) in the United States, Journ. Parasitology 26(5): 1940.

Woodruff, E. Seymour. 1908. The Ruffed Grouse, A Study of the Causes of
its Scarcity in 1907. Albany, N. Y., 1908.

Man's Relation to the Grouse

The eflFect of man upon the grouse depends upon the kind of man.
The American aborigines hved their hves almost hterally with these
birds, in their own habitat. The numbers taken by them for food
probably was no more than the excess above the winter carrying
capacity, and a healthy thing for the bii'ds. The clearings and trails
made by the Amerindians were improvements to the grouse range.
And the bird was a fairly important item of food to these men.

The early white settlers also depended upon the "partridge" for
a considerable amount of food. For a time the pioneer's land clear-
ing provided a beneficial interspersion of cover types over the grouse
range. But in most areas it outgrew this effect and more and more
excluded the grouse from the land, restricting it to the regions and
cover that could not be profitably and extensively cleared. Complete
extermination was the fate of this magnificent bird over much of its
original range. More than half of the northeastern states comes
within this category. Such is the march of civilization.

Surely no further elaboration is needed to indicate the importance
of man in gi'ouse ecology. In fact it may be said that the bird's future
rests with him. Man's relations with grouse are many faceted, as
man in the mass is many-sided. As a hunter he kills gi"ouse; as a
trapper he kills the enemies of the grouse, or the food of the grouse's
enemies, thus possibly adding pressure on the grouse itself. As a
woodsman he sometimes improves and at other times destrovs the
bird's home with saw, ax, and fire. As a farmer he protects the woods
with fence or he grazes it with livestock, he clears the land and keeps
it cleared, he keeps livestock and pets which may kill the grouse,
trample its eggs, or transmit disease to it. As a conservationist he
enacts laws, manages the land, and does other things that sometimes
benefit and sometimes injure the grouse. In all these and other ways
man's impact is felt, however unwittingly, by Bonasa umbcllus.

254

Man's Relation to the Grouse 255

AS A HUNTER AND TRAPPER

To the early settler, the grouse was one of several game species
that helped fill the family larder. Its role was purely utilitarian. As
agriculture grew and cities developed, the taking of grouse turned
into a commercial venture. There was little sport in hunting the
"fool hen" grouse of those days. Wilson ( 1812 ) speaks of the advan-
tages of a "good" dog thus ". . . the more noise he keeps up seems
the more to confuse and stupify them, so that they may be shot
down, one by one, till the whole are killed."

It is difficult from the records of the era of commercialization of
grouse to gauge how many were taken, what the populations might
have been, and other relevant information. Audubon ( 1856) records
their sale in the Cincinnati markets for twelve and one-half cents
each in 1820, which might indicate that large numbers were avail-
able. Elliott ( 1864), condemning the slaughter of the young, relates:
"I know of one firm that receives sometimes, on many Saturdays in
succession, five hundred pairs of these birds." Market hunting flour-
ished through most of the nineteenth century, varying in intensity
with the abimdance of the birds, but gradually waning wdth the
more limited range, the more and more restrictive laws, and the
slow change in emphasis toward hunting as a sport Monon (1875)
indicated the resentment toward this business when he says: "The
whole north country, from Amsterdam to Northville (New York),
is infested with pot shooters who hunt (out of season) for the Sara-
toga market." By the turn of the century the shift to grouse hunting
as a sport was about complete, although some illegal market hunting
continues even today.

Forbush (1912) gives one of the few records that indicate former
grouse abundance when he quotes E. F. Staples of Taunton, Mass.,
that in the early 1880's about one thousand birds were taken in a
season on twenty thousand acres that he ranged. He comments that
these were "real good" years, the last good ones that he had observed
up to the time he was interviewed in 1908. With densities only equal
to those of the 1930's (and former peak periods may have been
higher) there could have been on such an area from four thousand
to five thousand birds on the basis of a twenty to twenty-five per
cent kill.

256

The Ruffed Grouse

Effect of Hunting on Grouse Populations. Since hunting has been
attributed by many to be a serious factor in reducing grouse num-
bers ( "Its arch enemy, man, has shot and snared it almost to extinc-
tion," Bent, 1927), an analysis of this problem is of utmost impor-
tance. Among the means of analysis is the reported game take
required of each hunter annually in some states, including New York
and Pennsylvania. While these figures are open to many criticisms,
they are consistently arrived at and are valuable in shov^ing trends.

425.000^°°'°22 580,000

Total Reported Kill -Pennsylvania

" " " -New York

Computed average Bag -New York
" " -Penna.

Fig. 11. Reported Take of RuflFed Grouse by Hunters

Pennsylvania 1915-1943

New York 1923-1940

(The New York figures used for average seasonal bag for the years 1923—1931
obtained by multiplying take-per-licensee by 3.5 to get an estimated take-per-
game-reporting licensee. Those for 1923-1925 are based on hunting licensee
figures; those for 1926-1940 are based on 80 per cent of combined hunting, fish-
ing and trapping licenses. Pennsylvania figures are based on numbers of hunting
licenses. )

These records arc shown in Fig. 11 for the years 1915-43 in Pennsyl-
vania (taken from Ryder, 1944) and 1923-40 in New York (taken
from N. Y. S. Cons. Dept. Ann. Reports).

Any interpretation of these figures must weigh several pertinent
relations. The daily and seasonal bag limits were three per day and
fifteen per season in New York, two and ten generally in Pennsyl-

Man's Relation to the Grouse 257

vania. The number of Hcenses varied from less than three hundred
thousand to nearly seven hundred thousand in each state in different
years. Reports in New York are ordinarily received from only about
three-fifths of licensees and are taken by local clerks when a new
year's license is issued, which is anywhere from one to ten or more
months after the game was taken. From 1923 to 1925, specific hunt-
ing licenses were issued. From 1926 to 1939 the licenses covered
hunting, fishing, and trapping and no separate hunting license was
issued. In Pennsylvania, the figures were based on field estimates
from 1915 through 1936, and on hunters' game-kill reports from 1937
through 1943, received from almost all hunters. Landowners were
not required to be licensed to hunt on their own lands, nor were
licenses required of women. Thus the actual total take is very much
higher than that reported, probably about double. But the trends
should be accurately reflected in these statistics.

It has been thought by some "old timers" that the peaks of grouse
abundance have gradually lessened over each previous height. This
is not borne out by a comparison of the New York kill between 1923
and 1926 with those of 1935 to 1940. The author was about convinced
of this contention in 1936 when the tabulation of the 1934 hunter
reports was completed. It is indicated in the Pennsylvania records.

From all evidence the grouse populations had been generally as
high in New York from 1932 to 1934 as in 1935 to 1938. This is not
reflected in the kill records. Apparently this is a product of social
factors: the change in number of hunters; the unwillingness of many
hunters to shoot grouse for a few years after the closed seasons of
1928-29 and the extermination scare that accompanied the major
decline of that period; the shift of bird hunters to the pheasant, par-
ticularly of the younger men who never knew grouse hunting, etc.
Apparently these factors are gradually overcome as the good grouse
years continue, and thus produce a definite lag in the giouse kill
trend (as indicated by state records during the years of grouse
increase) behind the actual population trend.

When hunting is normal, and grouse are plentiful, hunting nowa-
days in the Northeast (from Virginia northward) results in a legal
bag of about eight hundred thousand birds. In the United States, all
grouse were estimated to have contributed 3,864,000 pounds of
meat in 1942 (S.A.F. Comm. Report, 1944).

Further efforts at evaluating the importance of hunting in grouse

258 The Ruffed Grouse

ecology were undertaken by field studies in 1930, 1931, and 1936.
Several areas of average quality grouse coverts running from about
one hundred to three hundred acres each in size in Tompkins
County, N. Y, (six of them in 1930 and thirteen in 1931), v^ere cen-
sused and the hunting results checked. The kill averaged ten per
cent of the birds in 1930 and sixteen per cent in 1931 (Edminster,
1933), individual units of cover lost from zero to fifty per cent of
their birds to the hunters. The very lov^^ return in the first year
probably reflects the reluctance of the hunters to pursue grouse
during the first open season after two years of closure and grouse
scarcity. This feeling was somewhat abated the next year although
it apparently took several years to disappear entirely.

Another check on the hunting take was made in the fall of 1936
on a portion of the Connecticut Hill area not devoted to the regular
studies. On a total of 1,379 acres of coverts, the hunters took fourteen
per cent of the October grouse population ( N, Y. S. Cons. Dept. Ann.
Rep., 1937). The highest kill on any of the three cover units involved
was twenty-three per cent.

During 1932 to 1935 two studies bearing on this subject were
made in Minnesota and Michigan. While done under different eco-
logical conditions, the results are interesting for comparison. Trip-
pensee (1935) found the hunting take on national forest land in
Minnesota in 1934 to vary widely. On one unit of two sections of
land the kill was only nine and four-tenths per cent; on another unit
of four sections it was thirty-eight and two-tenths per cent. A sum-
mary of his figures for all of the 6,400 acres involved gives an average
take of nineteen and seven-tenths per cent of the fall population.

The study in Michigan in 1932 and in 1935 was summarized in a
letter from Harry Ruhl to me in 1936. The 1932 results on the
Pigeon River State Forest showed a hunter take of six and seven-
tenths per cent of the September population (probably somewhat
higher for the prehunting season population of mid-October ) on the
four square miles censused. The 1935 studies were made on two
areas. Pigeon River State Forest and Escanaba River Tract (Mich.
St. Cons. Dept., Biennial Rep., 1935-36). The hunter bag on the
Pigeon River area was seventeen and three-tenths per cent and on
the Escanaba tract seventeen and four-tenths per cent. The Pigeon
River data when developed on the basis of a September census gave
a kill of only ten per cent, thus indicating that the 1932 records

Man's Relation to the Grouse 259

were much lower than the actual kill. In Pennsylvania in 1940,
Studholme ( 1941 ) found the hunting season loss to be twenty-three
and two-tenths per cent, of which at least seventeen per cent was
attributable to hunting by man.

These Midwest and Pennsylvania records corroborate the New
York results remarkably closely. We may conclude that the normal
hunter kill is in the neighborhood of fifteen to twenty per cent of the
prehunting season population.

We have discussed the total kill. Now let us see what this means
to the individual hunter. What is the average daily and seasonal bag
of giouse? This will be a discouraging revelation to those who are
considering becoming grouse hunters— and reassurance for the ex-
perienced shot who may still be smarting from being unable to get
his eye on 'em the last time out, for there are multitudes who have
fared worse. Before revealing the shockingly low effectiveness of
hunters it may be well to pause and note again that the grouse pre-
sents as hard a wingshot as any American game bird. ( I know quail,
woodcock, and duck hunters will rise up in wrath at that statement! )
That this creature could so develop from a "fool hen" in the space
of a century is a remarkable tribute to its adaptability.

High daily and seasonal bags of individual hunters are plentifully
recorded in the old literature. Forbush (1912) lists a number of
such occurrences from Massachusetts thus: "I can remember when
a market hunter going out from the city of Worcester by train each
day, walking to the covers and returning at night, killed from ten
to fifteen birds daily. . . . Mr. George Howes shot and marketed
three hundred and ninety-eight birds in one shooting season. . . ."
With the disappearance of the market hunter went the stor)' of large
individual kills. The very limit of the present-day laws, as for exam-
ple two per day, ten per season in Pennsylvania, three and fifteen in
New York, makes large bags legally impossible. But the hunter who
attains these figures is rare indeed.

On Fig. 11 is shown the seasonal take per hunter based on the
number of hunters who reported taking some game.^ Of course this
is very conservative since many of these hunters who reported taking
game did not pursue grouse at all. For what it is worth, the average
game-taking hunter in New York has bagged an annual total of from

^ This is based on actual data beginning 1932, on computed data 1923-31 using
the 1932-38 average of 3.5 times as many licenses reporting as reported taking game.

260 The Ruffed Grouse

less than 0.3 grouse to 1.7 grouse, not an alarming record to say the
least. Leopold ( 1933 ) gives 0.3 grouse per hunter for the seasonal
average in Wisconsin, M'hich checks well, since all of the Wisconsin
hunters are not grouse hunters either.

Other grouse-hunting states show similar records. For example,
in 1938, Pennsylvania hunters averaged .3 grouse, Connecticut
hunters .4, each for the season. For honest-to-goodness, dyed-in-the-
wool grouse hunters, the average is very much higher. Many of my
grouse-hunting friends make their daily limit several times each
season, and regularly take their seasonal limit. Van Coevering (1931),
in speaking of the miraculous recovery of the bird, noted that tv\^o
hundred and sixty-five grouse hunters in 1930 put up 11,000 birds,
and took 1,248. The average kill per hunter was 4.7, less than half
the legal seasonal limit.

Data on daily hunters' bags was gathered in the field studies of
grouse hunting, already discussed. In New York in the 1930 and
1931 studies, it took 19.0 and 12.6 hours of hunting, respectively, to
bag a grouse. The average hunting "day" was 5.75 hours in 1930 and
3.4 hours in 1931. Adjusting the take to the basis of an eight-hour
day, the average daily hunter bag was .42 grouse per man in 1930
and .63 in 1931. The actual daily bag on the short hunting days was
.33 and .37 respectively. This daily bag probably increased some in
successive years as both grouse and grouse hunting increased. How-
ever, the tendency is more marked in the increased total take than it
is in the individual take; that is, the increase is mainly in more hunters
taking grouse rather than the same hunters taking more grouse ( see
comparison of graphs in Fig. 11).

The distribution of hunting take is of interest too. Only about one-
quarter of the hunters took any grouse at all. Again we see that
relatively few hunters get grouse, even among those who are hunting
in grouse cover. If we convert our figures to include only the approxi-
mately twenty-five per cent of hunters who can hit grouse, the daily
eight-hour-day bag per hunter would range from 1.6 to 2.8 birds
per day. Half of these figures would give the approximate take on
the basis of the length of day actually hunted.

The records in Michigan in 1932 (Ruhl) indicate an average daily
bag of 0.7 grouse per hunter. It may be noted that this is consider-
ably lower than Van Coevering's (op. cit.) record of 4.7 birds per
average three days' hunting in Michigan in 1930, or 1.6 birds per
man day.

Man's Relation to the Grouse 261

The complete losses in grouse tliat are attributable to hunting b\-
men are larger than the actual take by the amount of the loss from
crippling. This factor is di£Bcult to evaluate and has been the subject
of wide differences of opinions, some observers depreciating it while
others believe it to be as important as the bag itself.

An exaggerated estimate of the crippling loss may be derived b\'
deducting the hunter kill from the difference between the posthunt-
ing season and prehunting season populations as determined by
censuses. This figure will include losses from predation, disease, and
accident, as well as from hunter crippling or losing. This total loss
has been about equal to the hunter-take figure. The corresponding
figure for the 1932 Michigan work ( Ruhl ) , was nine per cent, which
was 136 per cent of the hunter take. Trippensee (1935) indicates a
rather different result, his other losses amounting to only 21.6 per
cent of the hunter kiU for the hunting period.

The proportion of other-than-hunting-take losses of the New York
studies that is actually the crippling loss cannot be accurately deter-
mined, since these areas were not under intensive study. They indi-
cate only the upper limit of such losses. Some information on these
losses may be inferred from experience of expert hunters, aided hv
trained dogs. The crippling loss averaged about one-seventh of the
grouse bagged. These may be considered as minimum figures since
the hunters, being experts, probably left less than the average pro-
portion of cripples.

Some additional light may be shed on this problem by a considera-
tion of the fall losses on census areas that were not hunted. This is
from natural causes but not from gunshot. The loss during hunting
season on no-hunting areas is about equal to the hunter kill on a
hunted area; and the loss from all causes on a hunted area about
double tlie hunter take. Thus the losses other than the hunter take,
whether crippling may be involved or not, are about the same. This
indicates that the crippling loss is probably nearer the minimum
suggested above rather than the maximum. It is my estimate that
the true average loss from crippling will approximate twenty-five
per cent of the hunter take, or about four per cent of the prehunting
season grouse population.

The total hunting kill is the hunter take plus the crippling loss.
By applying the twenty-five per cent of take rule derived above for
tlie New York figures we find the estimated total hunter Idll to be:
1930-twelve per cent; 19Sl~twent\' per cent; 1936- seventeen per

262 The Ruffed Grouse

cent. Since these samples are only indicators of a general condition
we may conveniently use twenty per cent of the prehmiting season
population as the normal anticipated loss from hunting.

An interesting sidelight on grouse hunting is the success ratio—
the proportion of flushed birds brought to bag by the hunters.
About one grouse in a dozen flushed was retrieved. This is con-
siderably less tlian the success ratio of birds actually shot at, which
is a more correct indication of shooting skill. This was from two to
three times the ratio of flushed birds bagged, or about one bird in
four or five shot at was taken.

The effect of hunting on grouse populations varies with the density
of the population, as well as with the local intensity of hunting
pressure. Whether the hunters kill twenty per cent of the grouse, or
more or less, is important mainly as it affects tlie continued abun-
dance of the species. During the years of moderate-to-high abun-
dance, the twenty per cent of kill effected is of little importance in
changing the trend of grouse abundance.

Ordinarily there are about two and a half grouse produced for
each pair of grouse in the spring. Thus two and a half birds from
each four and a half living in the fall may be removed and still
maintain the normal number of spring breeders. If there were no
other sources of loss, the hunter could thus safely take fifty-five per
cent of the fall population. This represents the upper potential safe
limit of kill, but actually it cannot safely be approached owing to
unavoidable losses from other causes. It has already been noted that
winter losses from predation are low when fall populations are low,
other things being equal. Under such circumstances the probability
of predation may be twenty-five per cent or less, even where no
hunting has taken place. Following Errington's theory of vulner-
ability, which seems to hold fairly well with grouse, hunter take
would displace a loss from some other cause for all birds in an inse-
cure position. This becomes effective only in years of high abun-
dance. From both theory and practice we may conclude that a
hunter kill of twenty-five per cent is safe in all but years of scarcity,
and that this figure may be increased somewhat in years of abun-
dance. Surely the record of increased take and increased populations
in New York from 1930 to 1938, along with specific field data, show
that hunting as currently practiced has not been a factor in causing
serious grouse declines.

Tlie rather small part that hunting plays in total grouse mortality

Man's Relation to the Grouse 263

may be gathered by considering the fate of an average hundred
grouse eggs in the spring ( see page 285 ) . Of eighty-two which may
perish the first year, hunters are responsible for only five. Obviously
this five per cent of eggs accounted for by hunting is the same as the
twenty per cent of adults. It should be noted though that the num-
ber of mature birds lost are of far greater importance than an equal
number in the egg and immature stages.

Predator Reduction by Hunters and Trappers. It is a sad commen-
tary on our progress in conservation education that the average
American hunter still considers all hawks as bad hawks, can see no
good in skunks or weasels, continues to believe that if there weren't
so blankety-blank many foxes there would surely be more game for
him to shoot. This attitude results in the actual loss of conservation
values. John Q. Hunter still takes a pot shot at every hawk, owl, fox,
crow, turtle, snake, red squirrel, or whatnot that crosses his path,
and believes himself to be a worker for "the cause" when he does so.
Worse, he is often encouraged by the payment of bounties, or prizes
offered in "veiTnin control" contests. As a fur trapper he also takes
species that affect grouse populations, and some of the legitimate
game animals are also in this class.

The haphazard shooting of predators cannot be evaluated in its
effects on grouse. We only know that by far the greater portion of
this take is of species that do not prey on the grouse at all or are
of no real importance to grouse. Probably the best insight into
man's endeavors with respect to grouse enemies may be gained
by analyzing the organized efforts which have produced tangible
records. Notable among these are bounty systems, and of particu-
lar significance in the Northeast is the story of the Pennsylvania
bounties.

Gerstell (1937) has written a fine analysis and summary of the
results of bounty payments in the Keystone state, from which we will
quote freely. Bounties of one kind or another have been given there
since 1683 but the modem bounties that are aimed at game manage-
ment began in 1915. Most of the species on which bounties have
been paid in recent years are important grouse predators— red and
gray foxes, weasels, goshawks, and homed owls. The bobcat and
mink have also been subject to bounty but owing to their low num-
bers are not of great importance to grouse.

The goshawk's bounty record indicates that: (1) The hunter kill

264 The Rufled Grouse

does not reduce the abundance of goshawks in tlie succeeding win-
ter; (2) the number taken is but a small part of their population;
(3) the kill depends mainly upon the degree of southward winter
migration of the species. Gerstell summarizes: ". . . the payment of
bounties for the destruction of goshawks in Pennsylvania will never
result in the control of the species ... it appears most advisable
to discontinue. . . ."

With respect to the red fox which was removed from the list in
1929, he concludes that the record "seems to indicate that the spe-
cies was not controlled by the bounty," On gray foxes, "an increase
in the bounty rate (from $2 to $4 in 1923) has increased the number
of gray foxes annually presented for bounty, but as yet the species
shows no evidence of being controlled by the bounty." Likewise in
New York, in spite of constant persecution, the gray fox contiimes
to increase its range and its numbers. During the twenty-one years
prior to Gerstell's writing, an average of over fifty-two thousand
weasels had been presented in claim for bounty, yet "the bounty
system has not to any noticeable extent, if at all, controlled the
weasel even though two thirds of the system's cost has been ex-
pended in payments on the species."

The bobcat has reacted differendy. Subject to bounty almost con-
tinually since 1819, "bounty has brought die wildcat under absolute
control in Pennsylvania." In fact, it became so scarce that it was
removed from the bounty list in 1937 in order to afford it some
chance of preservation.

"As a predator control measure, the payment of bounties has
proven generally inefficient as it has placed under control only one
relatively small species population, while its effect on five others
has been negligible. ... It has been impossible to prove that the
operation of the bounty system over a relatively long period of years
has improved game conditions. Furthermore ... the annual
amount of money expended for bounty payments was controlled
not by the abundance of predators, but principally by climatic and
general economic conditions."

Trapping for fur, while undertaken as an economic enterprise,
affects several important grouse predators. Of these, the take of sev-
eral species in New York is reported annually. Most significant in
grouse ecology are the foxes, skunk, and raccoon. Only the weasels
among the more important mammalian grouse enemies are not re-

Man's Relation to the Grouse 265

ported. Analysis of these records, available since 1926, fails to reveal
either: (1) control of the furbearer, or (2) any correlation with
grouse abundance. The take depends considerably on fur prices.
Since the additional economic incentive of a bounty has failed to
show^ game management results, it seems unlikely that trapping or
shooting of predators for fur will have any appreciable effect on
grouse populations.

Organized "vermin hunts" reflect the character of the results of
predator control efforts by hunters not engaged in taking furs. They
are far from being a systematic control over species that may actu-
ally warrant some reduction in numbers. In the first place the aver-
age hunter is not very selective, especially with respect to hawks.
He is unable readily to identify many kinds of predators, particu-
larly the birds. He takes those easiest to get, which are usually the
beneficial rather than the '^bad" kinds. The need for control must be
based on a careful knowledge of the populations of both predators
and game— something which the hunter rarely has.

An analysis of the returns of ten sportsmen's club predator cam-
paigns taken at random from published records in New York and
Pennsylvania showed: (1) Three and six-tenths per cent of the
victims were identified as species which are important grouse preda-
tors (crows not included); (2) seventy-five per cent of the hawks
were either unidentified or were identified as beneficial species, and
this assumes that the Cooper's and sharp-shinned hawks were cor-
rectly named, which is doubtful; (3) of the three and six-tenths per
cent identified as among important grouse enemies, forty-eight per
cent were submitted for bounty; (4) sixty-four per cent of all kinds
taken were crows, seven per cent were starlings, seven per cent water
snakes, four and five-tenths per cent rats, five per cent red squirrels;
(5) of over 7,700 specimens only three were foxes, the most impor-
tant grouse enemy, and thirty-six were horned owls, the nmner-up.
We can only conclude that the greatest benefit from the sportsmen's
nongame hunting is his ovm self-satisfaction, the economics of shot-
gun shell sales and the recreation derived. He would aid conserva-
tion more by shooting clay pigeons.

Use of Guns, Traps, Snares, Dogs, Autos, and Roads. Man's tools
of pursuit have been an interesting sidelight to his relations with the
ruffed grouse, and in some respects have influenced its abundance.

266 The Ruffed Grouse

Considered somewhat in chronological order, we might discuss the
snare, net, deadfall, and various types of traps ahead of guns. But
for the early white man all methods were fair and effective. The
variety of implements utilized to catch the then unwary grouse
were almost endless. Most of these tools would be quite inefficient
on the "smarter" grouse of today but helped to bring untold thou-
sands to the markets of the past two centuries. In the sporting era,
the gun, primarily the shotgun, has been the only weapon of sig-
nificance.

There has been a great improvement in the effectiveness of sport-
ing guns in recent decades. However there have also been changes
in the hunter and in the grouse itself. Phillips (1937) says: "Techni-
cal improvements in the past forty years we can set aside, for the
early hammer guns, in skillful hands, were nearly as effective as the
more dainty modem weapons." He goes on, ". . . ff we compare the
average skill of the present-day brush shooter with the average of
the market shooter of fifty years ago, the result would be highly
amusing and very disastrous to the pride of the modem edition."
Phillips then points out that what tlie modem hunter lacks in skill,
he makes up for in numbers. It has taken increased protection as
well as a constantly growing wariness on the part of the bird itself
to make up for the modem army of hunters that annually takes to
the field.

Dogs have been used to aid man in the pursuit of grouse for a
long time, but in two quite dissimilar forms. We have already noted
(see page 21 ) the early use of a yipping cur dog to hold the attention
of a covey of treed "fool hen" grouse while the hunter picked them
off one at a time, from the lowermost up. Such a mutt is a far cry
from the magnificent, staunch, and silent setters and pointers of the
twentieth century. To many a sportsman the most enjoyable part of
hunting is working with a good dog, and die supreme thrill is in
seeing a perfect point on a closely held bird. In tlie northern states,
bird dogs were mainly trained on grouse up to the last fifteen or
twenty years. With the coming of a completely closed season in a
period of pronounced scarcity ( 1928-29 in New York ) and the great
increase in interest in pheasant hunting, it is rare today to find a
really well-trained grouse dog. Most of the pointing dogs are
"spoiled" on pheasants. So we find that the fai-famed grouse dog,
not long ago the pride and joy of the dyed-in-the-wool grouse hunter,

Man's Relation to the Grouse 267

is but a fond memory in the lore of the old timers, any one of whom
will delight in "pouring it on" in front of a crackling hearth on a cold
winter's night, recreating the exploits of "good old Pooch," known
in more dignified circles as "Royal Master Skylark of Worthingdon."

One of the most significant of recent changes in hunting problems
is the advent of fast transportation for the general public— good
highways and motorcars. Only three decades ago, grouse hunting by
any individual was confined to the few square miles that he could
reach from home on foot, or from the terminus of a short buggy or
train ride. How vastly different it is today. We jump in the car and
in an hour we are at coverts forty miles away. An hour's try at this
locale and we are off again, and in a single day we have taken a
quick course tlu^ough several coverts scattered over a hundred miles
or more. This increased mobihty has aided materially in adding to
the number of hunters, hence to the problem of finding good hunt-
ing for all.

At the same time that so many roads were becoming arterial high-
ways in the Northeast, other dirt roads that proved to be back roads
have become abandoned and inaccessible to modern automobiles.
Thus the hunter's ability to get around to the coverts has increased
in the broad sense while actually decreasing locally in many areas
where there is good grouse range.

The net effect of the changes in transportation are hard to sum
up, but are probably not seriously affecting the grouse. Whereas
formerly each hunter traveled in his own back yard, he's now in
someone else's. And the increase in grouse hmiters has not kept pace
with that of hunters in general, thanks to the pheasant and rabbit.

AS A FARMER AND LUMBERMAN

Land Clearing and Farming. When the white man first began hack-
ing at the forest wilderness in the eastern states, he found game
generally plentiful around the new clearings. This caused the falla-
cious belief in the supposed abundance of game throughout the
wilderness, for it seemed "self-evident that if wild life was abundant
around the settlers' clearings, it must have been more abundant
where man had not intruded" ( Edminster, 1941 ) . It did not occur
to him that his very clearings were aiding in producing this game
supply.

268 The Rulied Grouse

As land clearing progressed, the grouse range was generally im-
proved even diough the quantity of woodland was being reduced.
But in the more fertile and level regions the extent of land clearing
soon passed the optimum for grouse and the birds were tlien gradu-
ally exterminated from these parts of their range, except for occa-
sional islands of wooded swamp or rough land that remained to
support some grouse. In areas of poorer fertihty and steeper slopes,
as in southern New York and northern Pennsylvania, only from one
third to two thirds of the woodland was ever cleared. Here the
remaining grouse range is about the best in the Northeast. It is about
half again as productive of grouse as the Adirondack vdlderness
range even when all the open land is included, and the birds are
produced on only about half the total land area.

It is probable that the optimum amount of open land for grouse
range would be from ten to twenty-five per cent, if it were well dis-
tributed. This would not be fomid ordinarily in a regular farming
setup. Hence man's land-clearing activity ordinarily falls short of
meeting grouse needs in mountainous country and far exceeds it in
agricultural areas.

As a farmer, man maintains his open fields by cultural operations
with plough, harrow, mowing machine, etc. (see Plate 37). This
work serves to prevent grouse range from expanding, and at the
same time it maintains the valuable woodland edges. On intensely
farmed areas, cultivation prevents the grouse from reclaiming its
former range; on the hill farms it maintains the land in a higher-
productive condition for grouse than would ultimately result if the
area were abandoned to continuous forest. Many of the poorer of
the hill farms in the Northeast are being abandoned as unsuited for
profitable agriculture. In New York alone there are five million acres
of such lands— one sixth of the whole state. After abandonment, the
open fields grow into brush and for a period better grouse range
results. As natural afi^orestation progresses the range loses much of
its grouse productivitv and becomes a more extensive forest area.

Relations of Livestock and Pets to Grouse. Wherever hvestock
farming, either for meat or dairy products, is found on areas of
grouse range there is apt to be a conflict of interests. Fortunately,
much of the livestock that does occur in grouse country is fenced
out of the woodland areas, and much of the grazed woods are not

Man's Relation to the Grouse 269

in suitable grouse range anywa}'. When woodlands are pastured it
is detrimental to grouse and when this pasturing is intense enough
to create a visible "cattle line" it renders the woods practically non-
inhabitable to grouse regardless of its other attributes.

It should be pointed out that open-field pasturing is a means of
maintenance of balanced conditions almost as effective as cultural
operations. When accompanied by proper pasture management
measures for the prevention of woody plant encroachment, it is fully
eflFective in excluding grouse and in maintaining woodland edges
outside the pasture fence.

Man's pets, particularly dogs and cats, are of some shght signifi-
cance as grouse predators. His poultry may conceivably play a part
in aiding the spread of disease in grouse but contacts between
grouse and chickens are not common. In many small ways, the ani-
mals that are particularly associated with man affect the lives and
homes of the grouse.

Man's Woodcutting Activities. Beginning after the land-clearing
work has been completed and the extent of potential grouse range
thus delimited, man's most significant consequence to the grouse is
his work in the woods with ax and saw. Potentially he can make the
area most unproductive by clearing the remainder of the woods; or
poorly productive by allowing all the land to return to mature forest
(see Plate 6B). Actually he does neither of these things, but many
of his activities between these extremes determine the types of cover,
hence the character of the grouse range. With respect to man as a
woodcutter even more than man as a hunter may it be said that the
future of the grouse rests with him.

As a lumberman he may clear out extensive forest areas; or he may
cut selectively by species, by size limit, or for certain products as
tanning bark, alcohol wood, mine props, etc. Each of tliese methods
affects the grouse range differently. As a farmer he may take out
fuel wood, fence posts, or barn lumber, or he may contract with an
operator to skin the area. Again he may be interested in the trees
for themselves and not cut anything. Whatever he does fixes the
value of the cover which supports the birds. We have discussed
these shelter and food values in the cover in Chapters IV and V.
We will consider the deliberate use of woodcutting methods as a
tool in managing grouse in the last chapter. But since the ordinan

270 The Ruffed Grouse

farmer or woodland operator does not make his woodland cutting
plans with regard for grouse needs, it is a fact that most grouse range
is the product of the haphazard results of lumbering for various and
smidry individual reasons. For better or for worse, the grouse will
largely depend upon man as a woodcutter for the character of its
cover.

Use of Fire. For various reasons men sometimes set fire to the land's
vegetation. Ofttimes he does so accidentally. It takes no stretching
of the imagination to realize that fire in grouse cover will materially
affect the birds whatever the cause. It may burn up a nest; quite
often does. It may drive the birds out of important units of range
for a considerable period of time. Refening to conditions in parts of
Missouri, Woodruff (1908) says that the annual burning over of
forest floors has removed all tlie suitable cover and caused rapid
diminution of the grouse. Forbush ( 1927 ) includes "prevalence of
forest fires" among the important factors having much to do with the
decrease of this bird. Describing a forest fire, Krieble ( 1941 ) said,
"Grouse were incubating their eggs, and time after time we saw
birds fly out ahead of the flames utterly frantic; then, completely
bewildered they would wheel around and fly headlong back into the
flames and perish. How many grouse, and how many clutches of
grouse eggs were destroyed that day would be difficult to estimate,
but it's certain a severe toll was taken."

Forest fires in the Northeast today are generally so few, so well
controlled and so small that tliey do not greatly affect grouse popu-
lations. In fact small woods fires are not aU on the red side of the
ledger so far as grouse are concerned. After a year or two of barren-
ness, bums usually grow up to briars, cherry, popple, and other
shrubs and trees valuable to the grouse. A change in grouse cover
bringing good summer range and brood cover is effected. If not too
extensive tliis may actually improve the grouse cover conditions.
Phillips ( 1937 ) noted that the largest grouse population he had ever
encountered was on a tract of land in western Quebec, in the autumn
of 1895, which had been burned about seven or eight years pre-
viously.

To sum up, small fires ordinarily do grouse little damage, often
benefit them materially. Large and hot fires are apt to be quite
destructive and the benefits they bring in cover change are rela-

Man's Relation to the Grouse 271

tively unimportant because too extensive for the birds to fully use.
In some areas, such as the Adirondack Forest Park in northern New
York, where fire is rigidly controlled and lumbering of any kind
prohibited on public land, a little forest fire now and then is literally
a godsend to the game of the area.

AS A CONSERVATIONIST

Laws as a Means of Conserving Grouse. Laws, like the proverbial
poor, we always have with us. This is no less true of game laws than
of others. For a time after the first colonists arrived in the North-
east, the social organization was so loose and the supply of game
so plentiful for the few men there to harvest it, that there was no
need for fonnal laws to restrict man's pursuit of wild animals. But
as the settlements expanded and game occasionally became scarce
locally, restrictive laws were quickly enacted. Understanding man's
part in reducing game populations, it was only natural that they
should feel that the losses might be checked by restricting man's
liberties.

There were probably very local, and less formal laws in parts of
the Northeast in the seventeenth century, but the first law on grouse
that became a matter of permanent record was a partial closed
season in New York in 1708. However an earlier law in New York
in 1629 first set up the control of the hunting privilege on all game
in the state. (These and most of the subsequent references to dates
of laws are taken from Palmer, 1912 ) .

The spread and jurisdiction of laws increased rapidly throughout
the nineteenth century. Restricted seasons, curtailed bag limits,
limitations on use of certain types of weapons, and the prevention
of sale followed one after the other. The chronology of some of the
more important of these events affecting grouse in the Northeast
is as follows:

1818: Massachusetts— season closed March 1— September 1.

1820: New Jersey— season closed February 1— September 1.

1837: New York (Kings, Queens, New York, and Westchester
counties)— purchase or sale prohibited out of season.

1838: New Jersey— closed season extended to January 10— No-
vember 1.

1846: Rhode Island— season closed February 1— October 1.

272 The Ruffed Grouse

1851: Delaware— season closed March 1— July 1.

1858; Maine— season closed March 1— July 1.

1862; New York— sale prohibited during closed season, whole
state.

1867; New York— possession permit required to keep them alive
over winter.

1869: Pennsylvania— prohibited baiting, trapping, snaring; West
Virginia— season closed February 14— September 1; Maine
—closed season extended to February 1— September 1.

1878; District of Columbia— season closed and sale prohibited
February 1— August 1.

1887; Delaware— no hunting when snow on groimd.

1897; Pennsylvania— sale prohibited.

1911; New York— sale prohibited.

1928: New York— first completely closed season (effective for
two years).

This list is far from a complete record of the attention given to
grouse by the legislatures, but it gives the trend of events by pe-
riods from complete freedom to complete closure.

For a long time it was felt that game abundance could be ac-
complished by the simple procedure of legislating it. As more and
more restrictions successively failed to accomplish this end, various
supplementary efforts were one by one brought into play. But it
should be clearly recognized that even though legislation and law
enforcement are not synonymous with game management, they are,
ff adequate, the essential foundation of any soimd system of game
management. To say that our present laws are not adequate is to
state the obvious. Their immense variation from state to state is
ample evidence of this.

Probably the greatest weakness in the grouse laws ( as well as all
other game laws ) of most of our states today is the lack of authority
in the executive conservation officer to adjust seasons and bag
limits without resorting to specific legislation. Legislative machinery
is notoriously slow, and grouse do not wait on the good senators to
undergo marked increases and decreases in numbers. If lack of con-
fidence in the executive setup deters the legislature from delegating
this authority, then they should readjust the system so that there are
adequate safeguards. Most important, the authority is direly needed
somewhere ff good game management is to result.

Man's Relation to the Grouse 273

Except for the ability to adjust the annual take to the population
conditions each year, the laws relating to the grouse are generally
just and adequate. Restriction of weapons to modern guns and long
bows, prohibition of sale and elimination of commercialization are
about complete. The major field for future improvement lies with
regulation of land use.

Land Management for Wildlife Conservation. Within the frame-
work of his laws, and apart from his control of the hunting harvest,
man's primary eflForts as a conservationist are applied on the land
itself: plantings or seedings to add durable elements to the habitat;
cuttings to improve the existing cover and induce the growth of
other desirable plants; and supplementary practices such as pro-
tection from grazing, establishment of posted refuges, control of
other animal populations as deer and fur-bearers (see Plate 38).
For the most part we have aheady discussed the effects of these
practices on the grouse. Plantings, cuttings, and protection result
in various changes in cover conditions. These we have discussed at
some length. The relations of grouse to predators and other animals
has also been covered. We may now discuss the value of refuges and
sanctuaries. The application of all of the principles involved in these
relations will be treated in the chapter on management of the ruffed
grouse. But before we consider the practice of establishing refuges
and sanctuaries, we may pause to say that man's highest accomplish-
ment as a conservationist is his care of the land. The degree to which
he does this well is the measure of his success and of the endurance
of his customs, his pleasures, and of his very civilization.

Refuges and Sanctuaries for Grouse. In the history of man's efforts
to protect and increase game for hunting, the idea of leaving cer-
tain areas unhunted in order to assure the survival of at least some
individuals came early. But it has become a public policy in the
United States only in recent years. Pioneering in this work have
been the Audubon Societies and the U. S. Biological Survey (now
Fish and Wildlife Service ) in establishing sanctuaries for persecuted
birds, notably those threatened with extinction by the feather trade.
Of recent years the Fish and Wildlife Service has undertaken a big
program of refuges for migratory waterfowl. These have been im-
proved and managed (in contiast to a sanctuary which is merely
protected) both for resting areas and breeding giounds. The state

274 The Ruffed Grouse

of Pennsylvania has pioneered in estabhshment of refuges for up-
land game, notably deer, on lands acquired for game purposes. The
fever for acquiring public game lands for refuges hit New York
in the mid-twenties (see Plate 40A). The slogan was "A game
refuge for every county." While not the panacea it was hoped to
be, the refuge still is a valuable tool of game management.

The idea of the use and value of game refuges has been con-
siderably confused, particularly in the minds of sportsmen. It was
conceived that a game species, such as grouse, within the confines
of a refuge would increase greatly and overflow the refuge bound-
aries. This excess would be shot in the surrounding public hunting
grounds while the breeding stock would be preserved. It was also
contended that game in open-shooting cover near a refuge would
seek protection there when hunted. As it turns out, these theories
do not apply very well to grouse.

In the first place, a clear distinction is needed between the breed-
ing refuge and the protection, or "seed-stock" refuge. Breeding
refuges are needed only when the breeding facilities of a specie^
are hampered or endangered. Such a case is the well-known de-
struction of waterfowl breeding grounds by drainage activities. This
is not the case with grouse. The change of land ownership and plac-
ing of signs and a boundary wire do not help the birds' breeding
one iota. Furthermore, the principle of protection refuges assumes
that hunting is a vital factor in the survival of the species. Under
existing practice this is not true with the ruffed grouse in most
areas.

The questionability of the value of refuges for ruffed grouse in-
duced an evaluation of the principle in New York by censusing a
2,120-acre refuge area and a comparable public shooting area from
1935 to 1937 (Edminster, 1937). ". . . instead of producing a
greater supply of grouse, (the refuge) actually had fewer birds
than the check area in two of the three years . . . while analysis
of the effect of the small differences in the areas may serve to ex-
plain some of the variations in grouse numbers, the fact still re-
mains that the protection afforded by the refuge . . . did not serve
to enlarge the crop of grouse." The study concluded that refuges
were not of value in increasing grouse in normally hunted areas in
years when grouse are at least fairly plentiful. It indicated that
refuges might be more useful in times of scarcity, and that areas of

Man's Relation to the Grouse 275

abnormally heavy hunting might well have some refuges as insur-
ance against total local extermination. However, it may be desirable
to establish game refuges for other species in areas inhabited by
grouse.

MAN AS A LIMITING FACTOR

In a direct sense, as a destroyer of grouse, man is seldom the
limiting factor under modem conditions of hunting. Locally this
may occur, and potentially he might be generally over-destructive
if the legal restrictions and the restraints of hunting ethics were
lifted. In an indirect sense he is the dominant limiting factor in all
but wilderness areas. More vital to the grouse than man's hunting
are his use of the ax, saw, plow, fence, fire, and livestock which in
various combinations determine what shall constitute the grouse's
"home on the range."

o

REFERENCES AND CITATION SOURCES ON RELATIONS
OF MAN TO THE RUFFED GROUSE

Audubon, J. J. The Birds of America, 1856.

Bent, A. C. Life Histories of North American Gallinaceous Birds, U. S. Nat.

Mus. Bull. 162, 1927.
Dillin, J. G. The Effect of Forest Fires on Grouse, Bull. Am. Game Prot.

Ass'n, Vol. 9, No. 3, 1920.
Edminster, F, C. Man as a Predator on Ruffed Grouse Areas, Game Breeder,

Vol. XXXVII, No. 4, 1933.
Edminster, F. C. An Analysis of the Value of Refuges for Cychc Game Species,

Joum. Wildlife Mgt., Vol. 1, No. 1-2, 1937.
Edminster, F. C. Wildlife Management through Soil Conservation on Farms

in the Northeast, U.S.D.A. Farmers Bull. No. 1868, 1941.
Elliott, D. G. The Game Birds of the United States, U.S.D.A. Ann. Report,

1864.
Forbush, E. H. Game Birds, Wild Fowl and Shore Birds of Massachusetts

and Adjacent States, 1912.
Forbush, E. H. Birds of Massachusetts and Other New England States, Vol. 2,

1929.
Gerstell, Richard. The Pennsylvania Bounty System, Res. Bull. No. 1, Pa.

Game Commission, 1937.
Krieble, C. G. From Behind the Camera, Pa. Game News, Vol. 12, No. 5,

August, 1941.
"Monon." Shooting Out of Season, Forest & Stream, Vol. 5, No. 2, Aug. 19,

1875.

276

The Ruffed Grouse

Palmer, T. S. Chronology and Index of the More Important Events in Ameri-
can Game Protection, 1776-1911, U.S.D.A. Biol. Survey BuU. No. 41, 1912.

Phillips, J. C. Man's Influence on Ruffed Grouse Populations, 1937.

Ryder, W. C. A Review of the Ruffed Grouse, Penna. Game News, Vol. XV,
No. 3, June, 1944.

Studhobne, A. T. Ruffed Grouse Environment in the Scrub Oak-Pitch Pine
Forest Type, M.S. tliesis, Penn. State, 1941.

Trippensee, R. E. How Many Ruffed Grouse Can We Shoot? National Wal-
tonian. Vol. 3, No. 4, October, 1935.

Van Coevering, Jack. A Ruffed Grouse Miracle, Outdoor America, Vol. X,
No. 4, Nov., 1931.

Wilson, Alexander. The Natural History of the Birds of the United States,
1812.

Woodruff, E. S. A Preliminary List of the Birds of Shannon and Carter Coun-
ties, Missouri, Auk, Vol. 25, No. 2, April, 1908.

. New York State Conservation Department Aimual Reports for the years

1931 to 1941.
-. What Can Forest Wildlife Contribute to the War Program, S. A. F.

Comm. Report, Joum. Forestry 42(5), May, 1944.

■fer;\T^vA^^

^wm"^

10

Productivity and Populations

POTENTIAL AND ACTUAL PRODUCTIVITY^

Productivity is defined by Leopold ( 1933 ) as "the rate at which
mature breeding stock produces other mature stock," Before an
appraisal is made of this, the actual productivity, let us consider tlie
potential productivity— that constant property of the species which
is the theoretical rate at which the species could increase if no mor-
tality occurred. From this base line, we may examine the factors that
affect this theoretical possibility and derive the actual result. Being
promiscuous, breeding the first year after birth and laying a large
clutch of eggs (averaging over eleven eggs per clutch), the ruflFed
grouse has a high-breeding potential. Like the plant aphid that we
learned about in elementary biology, it would overpopulate the
earth in relatively few years if nothing checked its increase. Assum-
ing a balanced sex ratio, which is normal, a pair of perfectly health)'
grouse producing twelve eggs would result in fourteen birds the
first season; continuing with equally successful progeny, this orig-
inal pair of birds becomes 110 the second year, 782 the third, 5,486
the fourth, and so on.

That such productivity is never even remotely approached is due
to the perpetual interplay of numerous delimiting conditions, en-
vironmental influences, and decimating factors. To begin with, uni-
versal breeding is rarely attained; the occurrence of a few non-
nesting females, nonbreeding males, or infertile eggs makes an
initial reduction in the potential. Then losses in the egg, immature
and adult stages result from a never-ending gamut of hazards.

Subtracting these losses from the potential productivity, we have
remaining the actual. When the fall and winter losses bring the
population back to the original number of breeders, we have a

^ The first portion of tliis chapter is taken largely from Productivity of the Ruffed
Grouse in New York, Edminster ( 1938).

277

278 The Ruffed Grouse

stable population. When the annual increment is higher we have
an increasing population; and when the losses exceed the increment,
the population declines. In the long run, it cannot continue to in-
crease, nor can it continue to decrease without risk of extermina-
tion.

The wild-life manager strives to manipulate controlling factors so
that annual losses equal the annual increment in an environment
populated to its carrying capacity. Further, he aims to have the
greatest possible part of these inevitable losses occur from hunt-
ing by man rather than from other causes.

This objective is not regularly attainable in the grouse for it does
not have a stable productivity. Its numbers vary so widely that it
has been termed a cyclic species. But even apart from its general
fluctuations, the variation in the effect of many of the decimating
agencies year by year cause other variations in the productivity.
Marked irregularity was noted in areas and on individual coverts
within areas. But it is noteworthy that the early attributes of pro-
ductivity—sex ratio, breeding, number of eggs, fertility, viability
of eggs, nest survival— have remained remarkably constant, while
great variation occurred in both brood and adult survival.

ANNUAL FLUCTUATIONS IN GROUSE NUMBERS

The most complete and continuous productivity record of the
New York grouse investigation was obtained on the Connecticut
Hill Survey Area. The regular census units totalled two thousand
three hundred four acres of coverts (other portions of the survey
area which were utilized for special studies are not included here
due to interruptions in the census records ) . The population changes
taking place on this area for the nine-year period 1929 into 1938
are shown in Table 9.

For five years out of nine the productivity resulted in a larger
breeding population than the previous year and for four years in
an increased crop. At the outset, shortly after the low ebb of num-
bers in 1928, the population was far below carrying capacity. After
three years of very low populations, conditions favored a big in-
crease: Predator pressure was relatively low, disease incidence was
near zero and hunting pressure was eliminated for purposes of this
study. Weather conditions proved favorable too.

Productivity and Populations 279

An increase of ninety-three per cent from breeding stock took
place in 1930 with Httle reduction the following winter, leaving a
net increase of fifty-seven per cent in the spring of 1931. The seasons
of 1931 and 1932 also had high productivity, one hundred nine per
cent and one hundred twenty per cent respectively, but owing to
mounting winter losses the net yearly gains in breeding population
dropped to fifty-three per cent in 1932 and to only twenty-six per
cent in 1933. The forces of decimation were catching up, both preda-
tion and disease having increased notably. Then productivity in
1933 fell way off owing to a very high brood mortality of unknown
cause, resulting in a summer increase of only eighteen per cent of
the spring population. This was followed by a thirty-seven per cent
winter loss which, while somewhat lower both in number of birds
and percentage of September population than that of the previous
year, resulted for the first time in a reduction in breeding stock,
amounting to a twenty-six per cent net loss for the year.

Table 9

Population Changes on 2,304 Acres of Grouse Cover,
Connecticut Hill Area- 1929- 1938 ^

No.

No.

Loss of

Grouse

Fall <Lr Winter

Adult

Young

Adults in

Population

Loss of

Year

Breeders

Maturing

Summer

(September)

Adults

1929

123

40

(32%)

1930

83

4-

77

—

160

30

(19%)

1931

130

+

146

4

= 272 -

73

(23%)

1932

199

+

253

- 15

437

187

(43%)

1933 2

250

4-

65

- 20

295

110

(37%)

1934

185

+

265

33

417

176

(42%)

1935

241

+

99

- 53

287

132

(46%)

1936

155

+

153

- 23

285

175

(61%)

1937

no

+

143

5

248

108

(44%)

1938

140

'^ Figures taken from Edminster (1938) and derived from data in N. Y. S. Cons.
Dept. Ann. Reports, 1930-38.

2 1933 figures calculated from sample data.

Recovery in 1934 reached a high rate again with a productivity
of one hundred twenty-five per cent, the highest noted in any year
except 1938. This brought the fall population nearly up to the high
level of 1932. A normal winter loss (42 per cent) for a dense popu-

280 The Ruffed Grouse

lation left a breeding stock for 1935 which was thirty per cent
higher than that of the year before. In the ensuing summer, brood
survival again fell disastrously as it had in 1933, giving a recovery
of only nineteen per cent for 1935. This time the abnormal loss was
apparently largely caused by extreme precipitation, ten inches of
rain in forty-eight hours, which also provided the worst floods in
the history of the region. With the fall population thus lowered to
two hundred eighty-seven birds, we would expect a reduction in
winter loss. The number of birds lost was lower than in the pre-
vious winter, but the percentage lost remained high. In fact the
forty-six per cent reduction was the highest in the first seven years.
So also was the net loss in breeding stock— thirty-six per cent from

1935. Again we find the immediate explanation in local environ-
mental conditions— adverse winter weather that resulted in abnor-
mally high vulnerability to predation. And I pause to note that any
excellence of habitat would not prevent an increase of predation
under those extreme conditions of snow and low temperatures.

From the relatively low population carrying into the spring of

1936, recovery was efi^ected for the third time in the period of the
survey. Productivity was good although not as high as in some years.

Survival through the winter of 1937 proved to be the lowest yet
observed, sixty-one per cent of the September, 1936, population
being lost by the next April. This reduced the breeding population
to the lowest level since 1930. Once more, however, a productivity
of one hundred twenty-five per cent brought a quick recovery which
continued with increasing fall and spring populations until the
summer of 1940. Then, as had occurred twice before, the produc-
tivity fell to a low level, only fifty-two per cent, this time the result
of an abnormally high nesting loss.

From this record, we may conclude that during periods of grouse
increase, the net summer productivity is normally about one hun-
dred twenty per cent of the breeding population. At the peak, occa-
sional disasters, affecting brood mortality primarily, may reduce this
expectation practically to the vanishing point. The summary of the
actual productivity rates on the Connecticut Hill area, together
with the subsequent changes in breeding populations as compared
with the preceding year, are given in Table 10.

There is a direct relationship between yearly recovery, or pro-
ductivity rate, and population density. This same relationship car-

93

—

109

+ 57

120

+ 53

18

+ 26

125

- 26

19

+ 30

84

- 36

125

- 29

(High)

+ 27

114

52

122

Productivity and Populations 281

Table 10

PRODUCnVTTY RaTES AND CHANGES IN BREEDING POPULATIONS—

Connecticut Hill Area— 1930-1941 ^

Percentage Increase of Percentage Change in

September Population Breeding Population

Year over That in April ' from Previous Spring

1930
1931
1932
1933
1934
1935
1936
1937
1938
1939
1940
1941

^ Figures taken from Edminster ( 1938) and from data in N. Y. S. Cons. Dept. Ann.
Reports, 1930-41.

' Also referred to as the "recovery rate."

ries through to the changes in the subsequent breeding population.
Productivity increased annually for three years until the popula-
tion reached a total of four hundred thirty-seven birds (5.3 acres
per bird) in the fall of 1932 and held two hundred fifty birds (9.2
acres per bird ) into the 1933 breedmg season. With the population
at this high level, recovery in 1933 fell to the exceedingly low rate
of eighteen per cent. This change in trend resulted in the first de-
crease in breeding stock, in 1934. However, productivity then re-
turned to a high rate and another irruptive peak was reached. The
1933 situation was repeated in the spring of 1935, and the summer
productivity also repeated the pattern of two years before. The
reduction in both breeding and fall populations reached their low
for this decline in 1937, which was the largest decline witnessed.
Both figures fell to a point below any year except 1930. However,
this decline cannot be compared in intensity with that of 1926-27.
The full decline in breeding population from 1935 to 1937 was
fifty-four per cent; the decline in the early fall population for the
same period was forty-one per cent. From 1937 to 1940 the breed-
ing population increased to another peak in 1939^0, dropping again

282 The Ruffed Grouse

in 1941. Productivity was normal from 1937 to 1939 for increasing
populations. The very low productivity in the summer of 1940 pre-
ceded a moderate decline in 1941.

Reviewing the trends of the whole twelve-year period, the cor-
relation between breeding density and subsequent recovery seems
to be of basic importance. They appear to fall into three groups:
( 1 ) low breeding populations that have a low recovery rate; (2) low
breeding populations that are followed by a high recovery rate, or
moderately high populations having a high recovery; and ( 3 ) high
populations (or occasionally only moderately high ones) that are
followed by poor recoveiy rates (see Fig. 12).

Generalizing these correlations, we may regroup them into two
series in order to illustrate the inverse ratio between breeding den-
sity and productivity in its two opposite phases. Combining the
first two groups as those having low-to-moderate breeding densities
( 1931 almost falls with the second group anyway and 1930 had so
low a population that it was not yet able to attain a high reproduc-
tive rate, thus making it an exception to the principle involved),
we see the contrast with the other group of years having high breed-
ing populations that lead to low recoveries.

Errington's records on marginal giouse populations at Prairie du
Sac, Wis., suggest the same three phases, but here too the details
are somewhat obscured by inadequate data (letter from Errington,
January 21, 1942).

The inverse ratio between breeding density and productivity is
also shown for gallinaceous species other than ruffed grouse. Er-
rington's data on bobwhite quail ( a noncyclic species ) in southern
Wisconsin, R. E. Yeatter's work on prairie chickens and bobwhite
in Illinois, and that of Arthur Hawkins and others on mixed popu-
lations of bobwhite, Hungarian partridge, ring-necked pheasant,
and prairie chicken in Wisconsin, all illustrate the principle (Aldo
Leopold and Paul L. Errington, MS).

From the consistent evidence we may conclude that the losses
of birds that succumb following seasons of high-breeding density
are destined to happen from some cause as a result of the population
condition regardless of what the actual and immediate cause of
death may be. They are vulnerable as a result of their own num-
bers and the adjustment in population level is in the cards, irrespec-
tive of the operation of the various mortality factors. However, one

Productivity and Populations

283

should be cautioned against looking upon these inverse ratios as
ends in themselves. They are merely the manifestation of popula-
tion mechanics.

If density of population should prove to be instrumental in setting
the stage for catastrophic losses, prevention of such densities might

150

^100

Low breeding populations
foUowed by high recovery rate

i

S

50

Low breeding populations
with low recovery rate

High breeding populations
followed by low recovery rate

100 200

NUMBER OF GROUSE IN BREEDING SEASON

300

Fig. 12. Correlation of Grouse Recovery Rate with Breeding Density
Connecticut Hill, N.Y., 1930-1937
(Based on data from New York State Cons. Deft Ann. Reports and Edminster,
1938.)

conceivably prevent these losses. Then the recovery rate might be
maintained at a consistently high level. Because of these possibilities
this subject received particular attention in the New York studies.
(Edminster, 1938; N. Y. S. Cons. Dept. Ann. Reports, 1935-37).
Two coverts (apart from the regular work area on Connecticut

284 The Ruffed Grouse

Hill) totaling 777 acres were subjected to artificial control of the
grouse population beginning the winter of 1933-34. All but four
grouse were collected after the public hunting season. With this
beginning, the population was deliberately reduced each successive
winter to about half normal survival in order to maintain a breed-
ing population well below carrying capacity.

The productivity rates for 1935, 1936, and 1937, from moderately
sparse breeding populations were one hundred per cent, one hun-
dred sixteen per cent and one hundred sixty-one per cent compared
vdth nineteen per cent, eighty-four per cent and one hundred
twenty-five per cent, respectively, on the uncontrolled portion of
the survey area in the same periods. Thus, additional evidence sup-
ports the contention that as populations approach the range-carry-
ing capacity, their productivity declines, and that productivity is
higher in breeding populations well below carrying capacity.

LIFE EQUATION

In terms of populations, rather than the individual, the life story
of tlie grouse tends toward an equation— the population at any given
period of the year resulting from a balance of the losses on the one
side of the scale and the gains on the other during the preceding
year. Actually this yearly circle of events on any given unit of range
almost invariably leads to an inequation— with the gains sometimes
exceeding the losses and other times the reverse. This equation
might be expressed as: BP + MY — AL = BT' where BP — breed-
ing population, MY = maturing young, AL = adult losses and BT'
= breeding population of the succeeding year. If the yearly changes
actually result in a stable population, the equation may be sim-
plified to BP = BT' in which case MY = AL. When the population
increases from one year to the next, B'P' exceeds BP and hence MY
exceeds AL. This condition may be expressed graphically as fol-
lows (Fig. 13) for eighteen grouse breeders (nine pairs) producing
one hundred eggs.

Such an inequation cannot long continue. The trend must be
toward an actual equation, but oscillating first to one side and
then to the other of a true balance. From 1932 through 1941 in
New York, the balance has been nearly true. This theoretically true
equation is illustrated in Fig. 14, again for nine pairs of breederb
producing one hundred eggs.

Fig. 13. Life Equation of Grouse for an Increasing Population

Fig. 14. Life Equation of Grouse for a Stable Population

285

286 The Ruffed Grouse

VARIATIONS IN DIFFERENT HABITS

We have considered the actual productivity of the grouse popu-
lation on one area, which is typical of the disconnected coverts in
the submarginal farm region of southern New York and northern
Pennsylvania. Productivity in wilderness range of connected coverts
was studied in New York's investigation on an area in the eastern
Adirondack mountain region. (N. Y. S. Cons. Dept. Ann. Reports,
1932-37; Edminster, 1938).

There is an outstanding difference in the Adirondack record as
compared to that of Connecticut Hill-a seemingly greater insta-
bility of population. The first year of the Adirondack area survey,
1932, was apparently a peak year, and winter mortality data in-
dicate that 1931 also was a year of high population for this range.
Then in 1933 a marked drop occurred as a result of low brood sur-
vival. It will be noted that this same condition prevailed on Con-
necticut Hill that year. Then followed a recovery in 1934 to the
same level as 1932. This also took place on both areas. In 1935 we
again note a very low survival of young which brought a drop in
population in both areas. Recovery began here in 1936 and was con-
tinued on the Adirondack area in 1937 while Connecticut Hill re-
ceded. Thus the trends in productivity on the connected covert area
were somewhat synchronous with those on the disconnected coverts
during this period, but fail to show clearly the inverse relationship
between recoveiy and breeding density.

As an average, the productivity on the Adirondack area was
markedly lower, about thirty per cent, than on Connecticut Hill.
Thus we have an indication that the productivity as well as the
carrying capacity of connected coverts may be lower than that of
disconnected coverts in the northeastern range.

CHARACTERISTICS INHERENT IN THE SPECIES
THAT CONDITION PRODUCTIVITY

There are many innate characteristics of the ruffed grouse, as
wdth all birds, that directly affect its productivity. For the most part
these are consistent factors of the bird's physiological complex. The
more important are discussed in the following paragraphs:

Productivity and Populations 287

Sexual Relations. The ruflFed grouse breeds at the age of about ten
months. Practically all birds breed each year/ This matter has not
been checked perfectly since it is impossible to locate all grouse
nests on a sizable area, or to determine that every grouse has bred.
However, all the evidence indicates that a failure in the breeding
of grouse, at least in good grouse range, is rare.

The species is promiscuous in its sexual relations, which fact helps
prevent mimated birds. Audubon (1856) recognized the species as
being polygamous while Baird, Brewer, and Ridgeway ( 1905 ) sug-
gested that both sexes were promiscuous. There is no doubt a
fairly high proportion of grouse that mate with only a single bird
in a breeding season but this is apparently more a matter of expe-
diency than character.

A factor that tends to limit the breeding efficiency of the species
is the sexual-readiness cycle which occurs in both male and female,
and called "sexual rhythm" by Allen ( 1934 ) ( see page 30 ) . It ap-
parently works out very satisfactorily in nature since infertility is
very low, although it gives rise to serious problems with birds in
captivity.

Size of Egg Clutches. The size of the egg clutch has been gener-
ally known for the ruffed grouse for well over a century. Wilson
(1812) recorded tlie range as from nine to fifteen, while Audubon
(1856) gave it as from five to twelve. Forbush (1929) gives the
general range as from seven to fourteen eggs with occasionally up
to sixteen or seventeen, and he notes one case of a grouse nest con-
taining twenty-three eggs. Smith (1921) also records a nest of
twenty-three eggs and notes that it was probably the product of
two hens.

The average number of eggs per clutch for six hundred thirty-four
first nests ( that is, not including renests following a destroyed first
clutch) in New York from 1935 to 1941 was eleven and four tenths.
The commonest size was twelve eggs; the next commonest was
eleven eggs, and these two sizes made up over half the total. Nests
with nine, ten, thirteen or fourteen eggs made up most of the bal-
ance, although there were cases from three to twenty-four. Only
in one year was any number of eggs other than eleven or twelve
most prevalent; in that year there were more nests with thirteen

^ King did not find this to be true in his Minnesota studies.

288 The Ruffed Grouse

eggs than any other size. The yearly average varied from ten and
nine-tenths per nest to eleven and nine-tenths, with no regular
trends. There is probably no significance in these moderate varia-
tions.

It is likely that most egg complements below seven are the result
of physiological deficiencies in the female, and those exceeding six-
teen are probably the product of two separate females. A nest that
contained nineteen eggs was definitely known to result from two
birds, as was also one sixteen-egg nest.

The size of the clutch in renestings following the destruction of
the first nest is much smaller than in the first attempt. The earlier
in the season a nest is destroyed, the larger is likely to be any second
nest chitch that is laid. In sixty second nests throughout New York
the number of eggs ranged from three to twelve and averaged seven
and nine-tenths. One case observed was apparently a third attempt
at nesting, as judged by the time the eggs hatched (about July
25) (data taken from N. Y. S. Cons. Dept Ann. Reports).

Fertility of the Eggs; Hatchability. The fertihty records of several
hundred first nests were obtained in New York State from 1931 to
1941. The average proportion of infertile eggs was about two per
cent, the maximum in a single year three and five-tenths per cent
and the lowest one and one-tenth per cent. There was no regularity
in the variations and apparently no significance in them. The de-
gree of infertility did not reach a high enough level any year to
indicate any serious trouble. Records were obtained for a number
of renests too. These averaged a little higher in the proportion of
infertile eggs (for example, 5.4 per cent in 1937), thus confirming
the well-established fact that fertihty is lower with late breeding.
This factor is of little significance, however.

These records may indicate higher rates of infertility than ac-
tually took place. Any egg not hatching that showed no embryonic
development was considered to be infertile. A fertilized egg that
dies within a few days of the start of incubation will present the
same appearance; hence any cases of this nature that may have oc-
curred were placed in the wrong category. This only further em-
phasizes the consistently low infertility rate.

When we consider the rather complex nature of the breeding of
grouse, the near-perfect results are remarkable. Certainly we do not

Producti\aty and Populations 289

see in this record any indication of a mishap hkely to cause an ab-
normally low productivity. However, even a twelve-year record
may not bring out such a relation, if it exists, particularly since no
catastrophic declines occurred in the grouse populations in New
York during that period.

Eggs that die during incubation are identified by the presence
of the embryo when examined after hatching time. Most of these are
birds that fail to hatch properly, or in time, and die at full develop-
ment. However, as already noted, any cases of dead germs in the
very early stages of incubation would be misidentified as infertile.
Therefore, the figures given for losses by embryo death are possibly
a little lower than actual. The average proportion of embryos that
die during incubation, as determined from the New York records,
is about two and one-half per cent and the annual figures varied
from two-tenths per cent to five and nine-tenths per cent. Here, as
in the case of the infertile egg record, there is no regular trend in-
dicated, and the losses do not constitute a serious threat.

Dead embryos may result from either external influences, as low
temperatures or too long exposure when the bird is off the nest, or
an innate weakness of the germ. Since the latter cannot be iden-
tified as a cause of death, the tendency is to attribute the losses to
exposure. The fact is we do not know the answer. Shifts in per-
centage of embryo losses do not correlate with recorded weather
data, tending to indicate that some of these losses may well be due
to inherently weak germs ( data derived from IV. Y. S. Cons. Dept.
Ann. Reports).

Longevity and Breeding Years; Incubation Period. The full po-
tential span of life for grouse is rarely, if ever, attained in the wild
and never has been determined. In captivity, grouse have been
known to live for many years, but here too they rarely live long
enough to die from natural body degeneration. In the face of all
the hazards facing a grouse in its life, the potential life span is un-
important. Judging from the character of the bird, it probably can
live a decade or more.

The actual life span of grouse depends upon when its time is be-
gun. If we count from the time of hatching, the average length of
life is a very few months; if we begin counting only with birds that
reach maturity, the span is about twelve months; if we count only

290 The Ruffed Grouse

birds that reach a breeding season, the average lifetime extends to
about eighteen months.

The number of breeding years for either sex is not known, but all
indications are that it is the full life span. Surely, it is the full actual
life of all wild birds.

The normal incubation period is twenty-four days. This may some-
times vary slightly, usually by being lengthened. This abnormality
may result from excessive periods of time off the nest, which cumula-
tively result in a measurable extension of the full incubation period.

Ratio of Females Nesting. For the hen grouse, taken as an individ-
ual, it is normal to breed, nest, and lay a complement of eggs each
year. When we consider all female grouse, however, we find that
there are occasional exceptions. We might expect a few grouse to be
miable to locate males in marginal grouse range or in poorly popu-
lated range; but this apparently does not constitute the full story
of failure in breeding of wild grouse. Why some grouse do not breed
when there is an adequate number and distribution of cock birds we
do not know. We can only surmise that there are occasional indi-
viduals that are physiologically or socially out of balance, as among
humans.

Determination of the nesting ratio is di£Bcult, even as an ap-
proximation. Theoretically, the number of successfully nesting fe-
males {i.e. the number of broods), plus the number of broken-up
nests, less the number of renest broods equals the total number of
females when breeding and nesting are perfect. The nesting ratio
can, therefore, be derived from the formula:

Per cent nesting females =

(no. broods -f- no. broken-up nests — renest broods) X 100

no. females

With a good census, the number of females and the brood data
may be determined with accuracy. The di£Bculty arises in obtain-
ing the number of destroyed nests. Even the most diligent search
with trained observers will not bring complete success on a large
area. With our figure for destroyed nests too low, it is clear that
the nesting ratio will be lower than is actually the case. We can
estimate this discrepancy ff we may assume that the broken-up
nests not found are in the same ratio to those found as are the un-

Productivity and Populations 291

found hatched nests to the hatched nests actually located. Know-
ing these items we calculate the fourth. However, this assumption
is not always vahd.

Applying the above formula, and applying the corrective factor
for unfound destroyed nests, the nesting ratio ranged from well
below one hundred per cent to well over one hundred per cent.
Without the corrective factor, the nesting ratio was somewhat
below one hundred per cent. These latter figures are, of course,
conservative. At least some of the first set are excessive.

From this wide variation we may conclude: (1) There was a
marked variation in nest-location success year by year; (2) the
proportion of hatched nests located exceeds that for broken-up
nests; (3) some grouse fail to nest, and this proportion seems to
vary in different years; (4) the nesting ratio is high, approaches
perfection (if not actually reaching it) in some, and probably in
most, years.

Nimiber of Broods. The ruffed grouse raises one brood of young a
year. There are no exceptions to this rule in nature despite occa-
sional \ATJtten statements to the contrary. For example, Davie
(1898) concluded that grouse not infrequendy rear two broods
between the first of April and the middle of October, He was prob-
ably mistaking a late brood coming from a renest as a second brood.
When one considers that the mother bird cares for her young ones
for three months or more, and consumes more than another month
in egg-laying and incubating, it is obviously impossible to produce
more than one family in a season.

Sex Ratio. The proportions of the sexes is most accurately obtained
in the spring during the breeding season when many types of data
that bear on it are availal^le. It is at this season that the sex ratio
assumes its greatest importance too, affecting as it does the breed-
ing and productivity of the species.

All our evidence indicates that an approximate balance of the
sexes is normal,^ all seasons considered, but tliat there tends to be
a slight preponderance of males in the fall, and a small but con-
sistent excess of females in the spring. This adjustment of the ratio
from fall to spring results from the greater mortality of the male

^ King advises (letter of October 5, 1943): "My work in Minnesota leads me to
believe that at some phases of the cycle there is far from a balanced sex ratio."

292 The Ruffed Grouse

birds in the early spring because of their courtship behavior.

The spring records from 1931 through 1941 on the Connecticut
Hill area showed an extreme variation in percentage of males, rang-
ing from forty-two per cent to fifty per cent in different years, with
the corresponding proportion of females being from fifty-eight per
cent to fifty per cent. Sex ratio records on the Adirondack, Pharsalia
and Catskill areas of the New York investigation varied from forty-
five to fifty per cent of males in different years.

The sex ratio has been watched with great care as it has been
suspected of being an indicator of abnormal mortality, possibly
having a correlation with cyclic losses. Forbush (1912) recognized
a differential mortality among the sexes of grouse during popula-
tion declines as early as 1906. Leopold (1933) attributes this dif-
ferential mortality, which results in an abnormally high loss of
females and correspondingly unbalanced sex ratio, to disease.

The evidence I have observed has not shown this phenomenon.
However, the populations have not dropped precipitously in this
period either, and hence have not produced the conditions where
the unbalanced sex ratio is alleged to occur ( New York data derived
from N. Y. S. Cons. Dept. Ann. Reports).

POPULATIONS IN RELATION TO PRODUCTIVITY

We have discussed the fluctuations in the population of grouse
over periods of several years. These changes took place in specific
sets of conditions, but do not correlate the grouse populations to
the cover, or enable us to compare the populations with those on
other areas. To reduce population figures to a common basis, we
may obtain the ratio of birds and cover, either on a units-per-acre
or an acres-per-unit basis. Since grouse densities are relatively low
(compared with many rodents and small birds for example), the
figures will be most easily handled if derived as an acres-per-grouse
unit.

Grouse Nest Densities, If all females nest, and the sex ratio is fifty-
fifty, the density of nests would be half that of the adult birds in
the spring season. Due to the difficulties in locating the nests, the
entire number on a sizable area is never formd, hence the actual
density of nests is never determined with full accuracy. Evidence

Productivity and Populations 293

does indicate, however, tliat the nest density is very close to a
one-to-two correlation with breeding bird density.

Individual distances between nests has varied greatly. In one
instance, two nests were only twenty-five feet apart, but only one
nest was in use when observed, and hence both may have been made
by the one bird. In another instance, two females nested seventy-
five feet apart. In still another instance, two females were incubat-
ing concurrently in nests just fifty feet apart. In other years the
closest nests were from one hundred feet to over three hundred
feet apart. As an average, of course, nests are much farther apart,
even though they are relatively concentrated in the outer fringe
of the woodlands and brush areas. Most nests are more than six
hundred feet from the next nearest grouse nest even when the
density is high.

Densities of Grouse Broods. The density of grouse broods is lower
than the nest density by the proportion of nest failure that is not
adjusted by successful renesting. In comparison with the rather
incomplete determination of nest density, the brood statistics can
be very accurate, based as they are on observations taken over a
whole summer period.

In most years, the density will range between forty and seventy
acres of cover per brood in good range.

The number of broods seems to vary less by a notable degree than
either the density of grown grouse or the survival of the young birds
in these broods. There appears to be somewhat of a correlation be-
tween nesting success, i.e., number of broods, and breeding density:
the greater the breeding density, the lower the nesting success ratio;
and the lower the breeding density (within optimum bounds) the
higher is likely to be the success ratio. Thus an equalizing efi^ect
tends to stabihze the number of broods on an area.

Somewhat correlated with brood densities is the area needed by
broods. Since grouse brood territories overlap, in contrast to breed-
ing territories, the territorial requirement does not in itself limit the
brood density. The area actually used by broods, as indicated by
location of a series of observations, varies greatly. Of considerable
significance is the minimum area of cover that may support a grouse
brood. Evidence indicates this to be in the neighborhood of nine
acres, or slighdy less. I know one isolated covert of nine acres that

294 The Ruffed Grouse

raised a single brood year after year, aiid another of nineteen acres
raised two broods in some years. No smaller isolated coverts I have
known have maintained a grouse brood tliroughout the summer.

The average brood travels over a much more extensive area than
this. An average territory diameter of one-quarter mile is about nor-
mal, while extremes of travel range from possibly a very few acres
(a brood flushed every time from June through August in a single
briar patch ) , to several hundreds of acres over a diameter of at least
one and five-tenths miles.

Densities of Adult Populations. Densities of game bird populations
are usually taken during the spring breeding season and at the peak
population period in late summer or early autumn just as the young
birds have matured. For the grouse these times are April and Sep-
tember. Because the grouse uses only a part of the total range in
partly farmed areas like Connecticut Hill, all figures are based on
the ratio of area of coverts, exclusive of open land, to numbers of
grouse.

For this area as a whole, the greatest breeding season density
was about a grouse per nine acres. The lowest was thirty-six acres
per grouse, although lower densities had occurred before 1930.
Most years, the density was nearer the maximum.

There is less variability in the population density during the
peak period than in the breeding season. The greatest density was
five and two-tenths acres per grouse and the least nineteen acres
per grouse. Whatever the breeding density of grouse may be, within
the limit of a grouse per eighteen acres, the summer's productivity
will normally bring tie population up to a fairly high and fairly
constant level in September. Stating this another way, a satisfactory
fall population normally results from any breeding population that
has not fallen below a density of a bird per eighteen acres. The
effect of high breeding populations in reducing the recovery rate
tends to prevent irruptive peaks, thus somewhat stabilizing the
population.

The October hunting season densities are from about five to
twenty-five per cent below the early September figures, the average
drop being about twelve to fifteen per cent.

When we consider individual coverts (units of cover that are
more or less isolated from other cover by open fields, and ranging

Productivity and Populations 295

usually from one hundred to three hundred acres each ) , we do find
greater variation in grouse densities from year to year. Some units
increase their group populations from one year to the next while
others decline. Summation of these records for the whole area tends
to average out tliese irregularities. These records for individual
coverts are of interest primarily in showing the potentialities of the
different coverts for producing and holding grouse. Some show a
particular abilitv to produce giouse, i.e., high summer productivity,
others to hold them over fall and winter until the breeding season.

Extremes of breeding season densities of single cover units range
from less than four to almost fifty acres per grouse. Areas that reach
the greatest densities in their highest years, have relatively high den-
sities in their lowest years; those with low densities in poor years
also have a low density even in the best years. The primary key to
the spring density of levels of cover imits lies in the winter shelter
(primarily evergreen cover) conditions.

The cover units that provide the extremes of peak population
densities are not usually the same as give the corresponding ex-
tremes in breeding population densities. The greatest density of
grouse at about September 1 in a single cover unit was a grouse for
every one and eight-tenths acres (N. Y. S. Cons. Dept. Ann. Report,
1935), while the lowest peak density was less than one to thirty
acres.

These extremes, together with all the other year-to-year records
of individual coverts, show clearly that animal populations are very
dynamic, ever changing, and, barring occasional losses that prevail
rather uniformly over large areas as a result of some climatic factor,
are normally little related to other populations even a short distance
away. General increases and decreases in population, such as we
have discussed for a whole area, are merely the summation of these
individual unit trends. When the majority of units are increasing in
population, the whole area will probably show an increase; when
the majority are declining, the net total is likely to decline too. And,
as already noted, the population fluctuations for a whole area are
less violent than those of the individual component coverts, for, as
we make the summations, the individual gains and losses will cancel
each other to some extent.

Population densities on other areas have shown as much varia-
tion from Connecticut Hill as have the individual cover units among

296 The Ruffed Grouse

themselves in each of the areas. This is as would be expected, and
may indicate a different productivity or carrying capacity; or It
may simply mean the records were obtained at a different phase
of the population trend.

An area of eight hundred sixteen acres of continuous forest in
the eastern Adirondacks that was censused from 1932 to 1936
showed consistently lower densities than Connecticut Hill. The
breeding season densities ranged from a grouse per twenty acres at
the high ( although there was evidence of a somewhat higher popu-
lation the previous year) to forty-three acres per grouse. Produc-
tivity was also consistently lower here than on Connecticut Hill.
The peak densities ranged from thirteen and six-tenths acres per
grouse to forty-one acres per grouse. (N. Y. S. Cons. Dept. Ann. Re-
ports, 1932-36; Edminster, 1938). This record supports the general
principle that areas of continuous forest are not as productive and
do not have as high a carrying capacity as broken-up cover.

Records were obtained on an area in the Catskill Moimtain region
of New York of one thousand one hundred twenty-eight acres of
almost continuous woods in the western part of the mountains. It
was censused in March, 1936 and 1937 by biologists of the Resettle-
ment Administration. Here the densities were considerably lower
than on the broken-up covert Connecticut Hill area, being thirty-
seven acres per grouse in 1936 and thirty-one acres per grouse in
1937.

From all these records we may conclude that in the better North-
east grouse range we may anticipate a breeding season density of
eight to ten acres per grouse and a peak-season density of a bird
per five acres in the good years. In poorer cover and poorer years
the density falls to half these densities or even poorer, even on an
ordinarily good range. Shooting season populations are usually
around fifteen per cent below those of the peak season.

Numerous other censuses of ruffed grouse have been made in
recent years, notably in Pennsylvania, Michigan, and Minnesota.
While comparisons of these data with New York records are diffi-
cult to interpret, they do indicate the adaptabihties of the species
in various regions and the great variations that occur in grouse
abundance.

Fisher (1939) gives data on grouse densities on five areas in
Michigan censused for two or more years each between 1932 and

Productivity and Populations 297

1936. All were continuous forest. They are consistent in showing
the highest spring densities in 1933 with a marked decline in 1934.
In only one of three of the areas observed in 1936 had the popula-
tion regained its 1933 level. Maximum spring densities recorded by
Fisher for various areas were six and five-tenths, nine and two-tenths,
fifteen and two-tenths and fifty-three and three-tenths acres per
grouse; minimum spring densities on these same areas, in order, were
thirty-three and six-tenths, twenty-two and one-tenth, two hundred
thirteen, and one hundred twenty-eight. Here it appears that grouse
densities were higher tlian those of the Adirondack New York area,
but lower than those of Connecticut Hill. The September peak
season densities for these areas indicates a greater maximum sum-
mer productivity than for any New York areas. These maxima were:
two and seven-tenths, three and five-tenths, four and seven-tenths,
four and eight-tenths and seven and three-tenths acres per grouse;
minimum peak season densities on these same areas, in order, were
nine and seven-tenths, eight and nine-tenths, twenty-one and three-
tenths, fifty-eight and two-tenths, and eight and seven-tenths acres
per grouse. These high densities were never repeated two years in
succession; in fact, on all but one of Fisher's areas, a very high den-
sity level was reached only once in the period involved. This in-
dicates greater variability in peak populations than observed in
New York.

King ( 1937 ) reported on April and October populations of grouse
on an area of forest in Minnesota for the years 1927-36. The maxi-
mum spring breeding densities, four acres per grouse, were notably
higher than any New York areas achieved, although comparable
with many individual cover unit records. The high density period
extended three years from 1932 to 1934, increasing gradually from
a spring density of forty-two and eight-tenths acres per grouse in
1927 to the maximum and dropping back to twenty-six and one-tenth
acres per grouse in 1936.

The October grouse densities in King's area show much greater
range of variation than I have experienced. The maximum density
reached two and six-tenths acres per grouse in 1933. A decline to a
low of thirty-three and six-tenths acres per grouse took place by
1936. These records support the conclusion that grouse populations
fluctuate more widely in continuous cover regions than in discon-
nected cover.

298 The Ruffed Grouse

In the scrub oak-pitch pine forest type of Pennsylvania, Stud-
holme (1941) studied the grouse populations on a 1,440-acre area
from October, 1939, to April, 1941. The breeding season densities
were very uniform, about sixteen acres per bird, while the fall shoot-
ing season densities were from nine to ten acres per bird.

Population Densities of All Game Birds in Grouse Range. There are
several other species of game birds that occupy the same range as
ruffed grouse in the Northeast. The one of greatest importance, and
which occupies substantially the same range as the grouse, is the
woodcock ( Scolopax rusticola ) . An occasional ring-necked pheasant
(Pliasianus colchictis), or bobwhite quail {Colinus virginianus)
utilizes the margins of some ruffed grouse coverts. In broken-up
grouse range the mourning dove (Zenaidura macroura) is quite
common, while an occasional black duck {Anas ohscura) will be
found in territories of partridges. All of these species are found on
the Connecticut Hill area, the woodcock and mourning dove ^ being
most prevalent.

Woodcock ordinarily are from one-seventh to one-fourth as abun-
dant as grouse, the latter usually occur in years of rather low grouse
abundance.

Ring-necked pheasants occurred in small numbers except in win-
ter. The numbers of quail and black duck were not significant.
Mourning doves were about as abundant as woodcock.

The density of the mixed stands of game birds is only an approxi-
mation. The average spring density was about six acres per bird.
This was composed roughly of seven-tenths ruffed grouse, one-
eighth woodcock, one-twentieth ring-necked pheasant, and one-
eighth mourning dove. September populations averaged about four
and one-half acres per game bird. Here the proportions of species
were approximately three-quarters grouse, one-sixth woodcock, one-
thirtieth pheasant and one-twelfth doves.

CARRYING CAPACITY AND SATURATION POINT

Carrying capacity plays a key part in the determination of pop-
ulation density, hence in productivity. Leopold (1933) states that

^ The mourning do%e is not legal game in New York.

Productivity and Populations 299

"carrying capacity is a property of a unit of range" rather than a
property of a species. Errington and Hamerstrom (1936) define
carrying capacity for bobwhite quail as "the upper limit of survival
possible in a given covey territory as it exists under the most favor-
able conditions." They then restate it "as the level beyond which
simple predation upon adult birds, their o\\ti territorial intolerances,
and their tendencies to depart from coverts overcrowded with their
own or some otlier species do not permit continued maintenance of
population." In a later paper, Errington (1941) again summarizes
it thus: ". . . carrying capacity functions as a threshold of security
below which the numbers of wintering birds can rarely be forced
very far through attacks of wild predators alone." While we repeat^
edly refer to the carrying capacity of the environment or range,
these definitions clearly imply that it may not be a property of the
range alone but also partly a species property. This is commonly
the case with the rulfed grouse.

Any unit of grouse range unquestionably has a fixed upper limit
of winter survival, but this limit is conditioned primarily by environ-
mental attributes, such as the strategic location of cover components,
predation, and weather plus innate species characteristics, rather
than by limitations of food and shelter. Just as carrying capacity for
bobwhite quail is more a function of species properties than is the
case of the hoofed mammals, so carrying capacity for ruffed grouse
is even less dependent upon the food and shelter conditions of the
range than in the case of the quail.

The food and shelter conditions in normal grouse range have a
potential carrying capacity far in excess of the actual. Predation,
either direct or conditioned, usually reduces the winter population
well below the ability of the range to sustain and shelter. Even more
important, the species itself will not tolerate crowding beyond a
definite point. This brings in the concept of "saturation point," a
purely species characteristic. When the potential carrying capacity
of the range is a density that exceeds the saturation point, then satu-
ration point becomes a component in determining the actual carry-
ing capacity. This seems to be the case on good grouse range. On
poorer or marginal range, the potential carrying capacity of the en-
vironment is more likely to be lower than the saturation point, thus
not allowing the latter to be a factor. In this case, the carrying capac-
ity may actually be an environmental attribute exclusively. But even

300 The RiifFed Grouse

here, intraspecific intolerance sometimes plays an important role in
determining the nimiher of birds accommodated.

Using saturation point as the maximum density that the species
itself will tolerate ( a somewhat broader conception than that given
by Leopold ) , evidence shows this to be a bird to four acres. In every
case where tliis density was surpassed in the early fall, immediate
reaction set in in the fonn of dispersion and accelerated decimation.
On these well-populated coverts, carrying capacity was conditioned
by the action of this species characteristic. The maximum actual
carrying capacity exhibited by any covert observed (as measured
by winter survival) was four acres per bird of coverts exclusive of
open land. Areas like Connecticut Hill, taken as a whole, have a
carrying capacity of about eight to ten acres per bird. On a continu-
ous forest range like the Adirondack Mountains, it is about twenty
acres per bird.

ENVIRONMENT AFFECTS PRODUCTIVITY

We have observed that the actual productivity of a wild animal
like the ruffed grouse is far below the potential productivity. It was
noted that some of the inherent characteristics of the species are im-
portant in holding back the attainment of this potential; that its own
populations provide a check to its continued increase. We now con-
sider the other major limitation on productivity— the environment.
The environment is the physical world, organic and inorganic, in
which the grouse lives. It includes shelter, food, climate, and other
animals, including man— all the plants, animals, soils, and physiog-
raphy that make up its home and associates.

It takes no imagination to appreciate the profound effects that the
environment exerts in reducing the numbers of any animal species.
They succumb from exposure and starvation, are eaten by preda-
ceous enemies, and waste away as a result of the actions of para-
sites or disease organisms. But just how important are these various
factors? How are they interrelated?

Decimating Agencies. Most obvious factors are those agencies that
actually cause the death of grouse, either in the egg, immature, or
adult stages. The following discussion of grouse mortality is based
primarily upon the studies of the New York State Conservation De-

Productivity and Populations 301

partment on the Connecticut Hill area in New York, supplemented
by reports of other workers where available.

Mortality of Grouse Nests. My first year or two of experience in
observing the fate of numbers of grouse nests left me feeling quite
concerned over the future of the species. It was quite a shock to
learn that such a large proportion of the nests were destroyed. Grad-
ually it became apparent that what had seemed to be a high rate of
loss was actually normal and the birds increased in number at a rapid
rate in spite of these losses. Further analysis clearly reveals that this
loss is a natural part of the ecology of this species, as are similar
losses with many other birds.

Nest losses are generally high, ranging roughly from one-quarter
to three-quarters of all, which indicates a wide variation. The vari-
ation occurs on any area in difi^erent years, and also appears to be
consistently higher in some areas than in others. On Connecticut
Hill, nest losses ranged from thirty-two per cent to seventy-seven
per cent over eleven years while nests observed over the state at
large showed annual losses ranging from twenty-seven and five-
tenths per cent to fifty-six and one-tenth per cent.^ The general av-
erage annual loss on Connecticut Hill was slightly under fifty per
cent, compared with an average of only about forty per cent over
the whole state. This indicates that the Connecticut Hill nest mor-
tality has been somewhat higher than the general average (N. Y. S.
Cons. Dept. Ann. Reports).

The nests examined from the Adirondack Mountains had a notably
lower mortality rate than for the rest of New York, ranging from
seven and one-tenth per cent loss to twenty-four per cent. This is
quite logical since the predator pressure in these continuous cover
areas is generally lower than elsewhere in the state (data derived
from N. Y. S. Cons. Dept. Ann. Reports).

In west-central Pennsylvania, Studholme ( 1941 ) found the mor-
tality of eggs he observed to be twenty-two and five-tenths per cent.

Almost all nest loss is due to predation. Over the state as a whole,
over three-quarters of the nest loss was ascribed to predators, with
some cases being unidentified as to cause of destruction. No doubt
some of these were the result of predation too. On Connecticut Hill,

^ This figure is unduly iiffected by the 77 per cent loss on Connecticut Hill and
a high loss on tlie Pharsalia area. Excluding the data from these two areas, the figure
drops to 31.9 per cent.

302 The Ruffed Grouse

practically all of the nests that failed were destroyed by predators.

Most of the other nest losses are due to man (farmers, lumber-
men, etc., in their various work operations), with some losses result-
ing from fire (which may assume great importance locally), deser-
tion, flooding, etc.

The time of destruction is of considerable importance since it has
a direct bearing on the probability of renesting, hence on the ulti-
mate success in nesting. Very few nests are destroyed during the egg-
laying period— the last half of April and first week or ten days of
May in southern New York— probably not over five per cent of the
total nest losses occurring at this time in normal years. As soon as in-
cubation begins the rate of destruction increases, slowly at first, then
quite rapidly after the first week. Most of the nests are broken up
during the last ten days of incubation.

Now there is an inverse relationship between tlie length of time
that incubation has progressed and the likelihood of the bird's re-
nesting. The longer the incubation has proceeded, that is, the nearer
the nest is to hatching, the smaller is the probability of renesting.
If a nest is destroyed before incubation has begun, the hen will
probably make another nest. It is even possible that some of these
second nests may occasionally be mistaken for a late first nest. Up
until about two weeks after setting there is some chance, though a
constantly reduced one, of the bird renesting if broken up. Hens
whose nests are taken during the last ten days of incubation rarely
make another nest that year. And since most of the nest losses occur
during this last ten days, most broken-up birds do not make another
attempt.

Ratio of Females Unsuccessful in Nesting and in Rearing Broods.

We have seen that the mortality of grouse nests averages about forty
per cent. This means that about sixty birds out of every hundred
succeed in hatching their first clutch. We also noted that a rather
small but significant proportion of birds that lose their first nest will
try again, and some of these will succeed in hatching. Thus the ratio
of unsuccessful nesting is somewhat lower than forty per cent.

The exact ratio of renesting is difficult to gauge on the relatively
sparse data available. It is likely below twenty-five per cent. Using
this figure, about ten of each forty birds that fail in their first nest-
ing attempt (out of an average hundred birds), will nest again. The

Productivity and Populations 303

mortality among these second nests will be about half. Hence, we
may anticipate that five of them may hatch. This brings average suc-
cess in nesting to sixty-five per cent, and the failure ratio to thirty-
five per cent. Fluctuations will, of course, follow the same course as
occurs in rate of first nest mortality.

Another way of stating this fact is that on an average sixty-five
per cent of female grouse will hatch a brood. However, not all of
these hens will succeed in bringing their brood, even a single chick,
to maturity. Some will lose their entire family before the summer is
over. The facts on this matter may be derived by obtaining the num-
ber of female grouse with and without a brood at summer's end ( i.e.,
when the chicks are mature). Generally only about half the hen
grouse succeed in bringing at least one youngster to adulthood.
About one female in sLx that hatches a brood fails to raise any of the
chicks, in an average year. The ratio of hens succeeding in rearing
broods varies widely in different years from about three-quarters
to only one-quarter of the females.

Losses Among the Young Birds. Most baffling of all grouse mortal-
ity problems is that of the young in the first two or three weeks after
hatching. Our experience indicates that a loss of from one-quarter to
one-half of the chicks may be expected in the first month of life, and
that most of this loss usually occurs in the first three weeks. This
large initial loss was estimated by Sandys (1902) as likely being at
least one-third of the young before they attained the size of quail;
Forbush (1913) gave one half as the normal loss of young before
reaching maturity, while Roberts (1932) stated that young grouse
have so many enemies that even under normal conditions only a
small remnant of a brood escapes. The mean of twelve years' rec-
ords on Connecticut Hill for brood mortality up to September is
sixty per cent. Extremes have been fifty per cent and seventy-seven
per cent. Excluding two years of unusually high losses, 1933 and
1935, the annual loss has ranged from fifty to sixty-three per cent.
On the Adirondack area the mean loss over a ten-year period was
about SLxty-three per cent with extremes of forty-three per cent and
eighty-eight per cent (N. Y. S. Cons. Dept. Ann. Reports). Stud-
holme ( 1941 ) gave the brood loss in Pennsylvania as sixty-two and
six-tenths per cent. Thus the losses of young grouse are fairly con-
stant, with an occasional bad year when the results are apt to be

304 The Ruffed Grouse

disastrous. While the immediate causes of death of these chicks re-
main obscure, the relationship of high population density to poor
recovery, and vice versa, is probably the guiding consideration.

The normal curve of infant loss is steep during the first month
of life, levels off considerably during the second month, and then
drops a little more abruptly for a time in the third month (see
Fig. 15 ) . The causes of the early season losses are somewhat obscure.
Certain facts are quite clear, and these help to delimit the prob-

GROUSE CHICKS LOST

80

- 20

June 1 June 15 July 1 July 15 Aug. 1 Aug. 15 Sept. 1

SUMMER GROWINGUP PERIOD

Fig. 15. Normal Grouse Infant Mortality Curve June to September

lem. Predation is of little significance at this season. Diseases and
parasitic infestations apparently are not of great significance in nor-
mal years. And man has little direct effect on brood survival.

When a mother grouse brings off an average brood of about ten
chicks she has a big problem of family control on her hands. Ten
active little birds are no simple group to keep well fed, brooded, and
husbanded together. In the first place, there will be considerable
variability in the vitality of the individual youngsters. Even as with
us mortals, some grouse will be born weaker than others. These

Productivity and Populations 305

weaknesses may arise from unequal incubation, excessive chilling
during embryonic development, through some accident of develop-
ment in the egg, or they may be carried through the egg from some
weakness in the mother. We have no proof for any of these hy-
potheses other than the inference that may be drawn by the elimina-
tion of other factors and by parallels with the known facts of do-
mestic animals. There is little question but that a considerable por-
tion of the early deaths of grouse chicks arises from these weeklings'
dropping by the wayside when diey fail to keep up with the travels
of the family.

An interesting sidelight on grouse psychology is the reaction of
mother giouse to the size of their broods. Evidence gathered time
and again indicates that they have little cognizance of numbers.
Following a disturbance when the brood is flushed and dispersed,
the chicks reassemble at the call of the mother when the coast is
clear. If some of the birds fail to arrive with the majority, the brood
will go its way with no apparent concern on the part of the parent.
She seems to be quite satisfied as long as she still has some chicks
with her. It is apparent that this characteristic will result in the
abandonment and consequent loss of quite a few chicks. Accidents
of other types will occasionally result in losses: chicks that stray and
become lost, little ones that tumble into some cavity from which they
cannot escape, and otlier similar chances sometimes cause the death
of young grouse.

Probably the most vital immediate cause of early brood losses is
the weather. Young grouse are not always able to withstand the
drenchings of a cold June thunderstorm or the chilling of an ab-
normally cold night. If well brooded during these emergencies little
trouble is likely to occur. But with many chicks to care for, the like-
lihood of some of them getting chilled or soaked is considerable.
As the size of the brood decreases the mother's ability adequately to
brood them in adverse weather improves and the chicks themselves
are more hardy as they grow older. Thus the losses from exposure
are usually over in the first few weeks.

It must be recognized that these heavy losses to baby grouse soon
after hatching are a normal expectancy if they do not exceed about
thirty-five per cent in the first three weeks. But occasionally an ab-
normal combination of adverse weather in June or early July will
result in far greater losses. In 1935 a period of excessive rains in early

306 The Ruffed Grouse

July, covering a three-day period, clearly caused a large loss of young
grouse, and led to the highest brood mortality (77 per cent) ob-
served in the eleven-year study. Since these excessive losses occurred
following high breeding populations, it is likely that they were pre-
conditioned by population density relations, whatever may have
been the immediate decimating agencies.

As the young grouse grow older, predation gradually becomes a
more important cause of losses. Along about the time the birds
reach eight weeks of age, the loss from predation notably rises. This
August increase in deaths from predators is largely attributable to
the accipitrine hawks, mostly shai-p-shinned hawks, as they are more
common. The susceptibility of the young grouse to these attacks in-
creases at this time for a short period. The chicks assume more inde-
pendence of their mother and take more excursions on their own as
they begin to feel their oats. Those that escape are soon educated
to cope with this danger and the rat6 of loss drops off again. After
the moult in August and September, the young birds assume adult
plumage and are hardly distinguishable from their elders. Their
losses then are considered to be a part of the adult mortality.

Adult Mortality. As a matter of convenience, we calculate the an-
nual losses of adults from September to September rather than on a
calendar year basis, since the young grouse become adults in Sep-
tember. The yearly losses of adult birds on New York's Connecticut
Hill study area over a twelve-year period have a mean of forty-eight
per cent. This mean was exceeded in eight years and in four years
the losses were below this figure. The extremes were twenty-three
per cent and sixty-four per cent. In five of these years the loss was
between fifty-one per cent and fifty-eight per cent, and only once
did it exceed the top of this range. On the Adirondack area the ex-
tremes were similar, twenty-two per cent minimum and seventy-
three per cent maximum witli a mean of about forty-six per cent
over a ten-year period {N. Y. S. Cons. Dept. Ann. Reports).

The spacing of losses of grouse through the year is fairly consist-
ent. The high mortality period is from January through April, with
the hunting season period added in heavily hunted regions. Summer
losses are ordinarily negligible, but rise considerably in September
as the young of the year join the ranks of the grown-ups. There is a
generally steady increase in rate of loss through fall and winter.

Productivity and Populations 307

culminating in a peak in the breeding season in March and April.
From then on to summer, the losses drop off rapidly.

The increased mortality rate in winter is due in a large measure
to the greater vulnerability of the birds to predation resulting from
winter weather and poor cover conditions, and from the increased
intensity of hunting by predators. The continuance of the high mor-
tality into the breeding season is to a considerable extent the result
of increased losses of male birds resulting from exposure during the
courtship period. This differential mortality among the sexes seems
to be balanced at other times of the year by greater losses among
females than males. This is particularly true in the summer and fall.

In most years the greatest immediate source of adult loss is preda-
tion. However, it is highly probable, in fact quite certain, that
predation takes more than its fair share of the blame. When there
is evidence of predator work on the remains of a grouse, the cause
of death is usually credited to the predator. Many times the predator
plays a secondary, and often unimportant part in causing the death
of the grouse. Grouse that are weakened by wounds from hunting
or from physical accidents are often taken by predators before suc-
cumbing from their injuries. I recall an occasion when a man winged
a bird and was unable to retrieve it. Over a month later my setter
dog pointed a grouse within a hundred yards of this same location
and, finally breaking point, caught the bird. It proved to be the one
winged some time before— still in good health but unable to fly. Or-
dinarily this bird would have been caught by a wild predator— and
its loss erroneously attributed to that animal. Likewise, some grouse
that die from disease or accident are eaten as carrion. These cases
too would have evidence of predator work, although many times
they can be identified as carrion from other evidence.

Notwithstanding these facts, most adult grouse in the Northeast
wind up their lives as food for some other wild creature, as the Lord
intended they should. The exact proportion that die primarily at the
'Tiands" of claws or talons cannot be determined because of the
obscuring evidence already noted. And even among those that actu-
ally die directly and essentially from predation are many whose fate
was influenced by weather conditions, overpopulation, scarcity of
other prey species, etc. After an evaluation is made of all of the com-
plicating factors, we conclude that predators cause about fifty per
cent of the kflling of grown birds in normally hunted coverts and

308 The Ruffed Grouse

eighty per cent in coverts protected from hunting by man. The dif-
ference, thiity per cent of the mortahty, is the nonnal hunting-season
loss from gunning. In coverts not well hunted tliis figure may be
lower, in overhunted areas sometimes higher. The other one-fifth of
the mortality accrues from disease, accidents, and other causes.

Disease as a decimating agency is difficult to evaluate. Often its
effects are obscured by predator action. It is inconsistent in that
its importance varies greatly year to year and from one area to an-
other. Generally it apparently is not a high or widespread cause of
death in grouse. In ordinary years of good grouse abundance, that
is years when grouse populations are not rapidly and sharply declin-
ing, we estimate that the various diseases and parasite infestations
cause from five to ten per cent of the mortality. It is probably
greater in imshot range than elsewhere.

There is evidence that diseases sometimes play a much bigger
part in grouse mortality, these usually being years of great decline
from peak populations. Allen found that unusually large numbers
of grouse died from attack of the gizzard worm on an area near
Ithaca, N. Y., in 1926-27 and apparently contributed largely to the
cyclic decline that took place.

In spite of the many organisms that afflict the grouse, many of
which can be lethal, we are hesitant in gauging the importance of
disease as a factor in grouse mortality. During the period of my
observations, no epidemics in grouse were observed. True, there
were great variations in the incidence of infestations, the incidence
generally rising with the increase of grouse populations. There was
also variation in the observed losses from disease in different years.
But in no year did we find evidence to show that disease was a mor-
tality factor of prime importance.

Death resulting from mechanical injuries sustained in physical
accidents is not uncommon with grouse. Here too, as with disease,
the primary cause of death may be obscured by the handling of the
bird by predators. Accidents are more common with young adults
than with older birds, many occurring during the first autumn after
reaching maturity. These early autumn losses are often associated
with the "crazy flight" phenomenon (see page 43), and are a
function of the fall shuffle of grouse that takes place with the
break-up of the family groups.

As a general rule, accidents usually cause about five per cent of

Productivity and Populations 309

the losses of adults, but this figure will vary widely in different
localities and years.

I have already indicated that hunting by man in normally well-
hunted areas results in losses approximating fifteen per cent of the
birds. This is the equivalent of about thirty per cent of the yearly
mortality. It is self-evident that this figure will vary widely in dif-
ferent times and places. Adding up the losses from predation, hunt-
ing, disease, and physical accidents, we practically account for the
whole mortality. An occasional bird may be killed by fire; very rarely
it is possible that one succumbs from starvation or over-exposure to
bad weather. These factors are negUgible, however, as direct deci-
mating agencies.

Limiting Effects of Environment. There are many environmental
influences that condition the decimating agencies in one way or an-
other. We have just noted that starvation is of no importance as a
killer of adult grouse. But when a grouse wanders out into an apple
orchard to get at a desired food, it may thereby expose itself to an
easy attack by some predator, or a hunter. Thus the food problem is
interrelated with predation and hunting losses. The pursuit of food
may likewise be a factor in bringing about disease. Concentration
of a food source may mean concenti^ation of grouse, and thereby fa-
cilitate the spread of disease organisms. Some items of food carry
disease organisms, as for example the sow bug that is the alternate
liost for the parasite, Dispharynx spiralis. If the parasitic infection
weakens the bird so that it is killed the more easily by a predator,
then the food relation, the sow bug, is two steps removed from the
actual cause of death.

Climatic factors are of immense importance in conditioning the
decimating agencies. When conditions of snow, temperature, and
wind cause grouse to roost in the snow, the birds become more vul-
nerable to predator attack. Likewise the quality and distribution of
food and shelter play a part in the complex of weather-cover-preda-
tion. Then man further complicates these relationships by affecting
the area in various ways— by cutting, burning, cultivating, draining,
or by adding still another in the chain of factors, livestock, that graze
and browse the vegetation. Insects and the other lower forms of
animals play many parts in the interrelationships of the grouse en-
vironment. Physical factors, as slope and light, add to these complexi-

310 The Ruffed Grouse

ties. And finally the grouse themselves affect each other's survival
by their fighting, courting, and other relations.

We cannot unravel all of these many involved relations of grouse
ecology. Even if we could, we would surely find that all must be
considered in evaluating tlie needs of the species, and in translating
them into management. No one aspect of the maze can get far out
of balance without seriously affecting the whole.

Pervading all grouse environmental needs as of first importance
is habitat: food, and shelter. The cover definitely delimits the possi-
bility of grouse inhabiting any portion of the range. Within the
framework of acceptable cover, all of the other factors and influ-
ences combine to determine the level of the grouse population.

Cycles of Abundance. A cycle, according to the dictionary, is a
round of operations or events— a series which returns upon itself.
A recent adaptation of this usage in connection with animal popula-
tions refers to changes in numbers of a species that take place regu-
larly according to uniform time intervals, and usually include a pre-
cipitous decline phase. During this decline, the species may lose up
to ninety per cent or more of its members in a relatively short time.
Species that are thus regular and violent in their population changes
are said to be cyclic species. The ruffed grouse has been among the
group so classed.

The general concept of an animal population cycle is that an
abrupt drop in numbers occurs some time after the species has
built its population up to carrying capacity— i.e., all tliat the environ-
ment can support. In some cases this carrying capacity level is tem-
porarily exceeded, thus assuring a quick and large drop in num-
bers. The size of the population itself may bring about its down-
fall through the medium of some mass-decimating agency; in this
case the regularity of the cycle would be due to the constancy of the
time required to bring the population to the critical level. It may be
that the periodicity is involved in a factor that is independent of
the animal population in question. Such might be the case ff years
of sunspot maxima, which themselves occur in cycles, brought about
some environmental influence which caused catastrophic decima-
tion regardless of the position of the population level at the time.

That these two types of cycles exist in nature, no careful student
of the subject can deny. Whether or not ruffed grouse populations
adhere to any such biological or physical laws is another matter.

Productivity and Populations 311

Before we examine the evidence relating to grouse and cyclic be-
havior, let us pause a moment to cite clear-cut examples of the two
types of cycHc effects on living things.

The case of the apple tent caterpillar {Clisiocampa americana) is
a good example of the cyclic eJBFect brought about by a population
increase to an untenable density. The caterpillars find an abundance
of food on wild and cultivated apples, cherries, and other food plants
and, with their inherently high reproductive ability, increase their
numbers rapidly over a period of years. The parasites that live on
these larvae have a progressively easier task of locating their hosts
as the population of the host grows. The curve of parasitic infesta-
tion heightens even faster than the curve of tent caterpillar abun-
dance and finally, in one opportune year, catches up and causes
the abrupt loss of most of the worms. The next year the population
of the caterpillars is at the low of the cycle. Likewise the parasitic
infestation is lowest, since the parasites have nearly ruined their
own food supply. The upward trend of the cycle begins, slowly at
first, but gathering momentum each year. And once more the curve
of infestation sets out to catch the curve of host abundance. Thus
we have a simple cause-and-effect cycle of abundance in two ani-
mal species brought about by the specific relationship of parasite and
liost. True, other factors such as the weather conditions will affect
this picture so that it never appears in actuality quite as smooth in
action as we have pictured it. Nevertheless, these populations do
follow this basic cyclic pattern.

Cycles in some forms of plant and animal life caused by physical
phenomena that themselves occur in cycles are fairly well estab-
lished. Best knov^m are the cycles in "sunspots" that repeat at about
ten- to eleven-year intervals (see writings of R. E. DeLury, Harlan
Stetson, and others ) . These solar changes apparently are related to
the well-established weather cycles ^ that occur in periods bearing
multiples of about eleven years (Abbot, 19-35). It has been shown

^ Authorities do not yet agree as to the effects of sunspots on our weather. In
Science (Vol. 95, No. 2473, May 22, 1942) it was reported: "Sunspots have their own
effect on the earth's weather. They give off vast streams of electrically charged parti-
cles that shoot through space. Some of them, entering the earth's atmosphere, serve as
nuclei for tlie condensation of water vapor in the upper atmosphere and thus lead to
the increase of cloudiness and of rainfall." Nine months later, in Science News Letter
(February 27, 1943, p. 137), Dr. Seth B. Nicholson of the Mt. Wilson Observatory
was reported to have concluded that "Vague correlations between sunspots and the
weather probably exist, but the weather is affected so much by other factors that tlie
influence of sunspots is insignificant."

312 The Ruffed Grouse

that these long-term variations in temperatures, precipitation, and
other weather factors affect tree growth (Douglass, 1941), as they
logically would. There is considerable evidence that sunspot cycles
are related to changes in population of some rodents, of migratory
salmon, and other animals. The relationship may be indirect, through
the effects on environment or climate; or it is conceivable that it
may bear some direct relationship, as yet not imderstood. It is now
well-known that simspot activity affects our radio waves. Who are
we to say that these changes in solar radiation may not directly af-
fect forms of Iffe on the earth, with animals dependent on vitamins,
hormones, and what not? One author has even gone so far as to cor-
relate wars and business cycles with sunspots (Stetson, 1937).

Now what is the evidence supporting and refuting the existence
of giouse cycles, and what conclusions may we draw?

That there are fluctuations— and wide too— in grouse numbers
is well-known. Clear and fairly complete records of years of abun-
dance and scarcity go back to 1904 for some areas, notably the
northeastern states. Scattered records indicate certain years of
scarcity or abundance for some areas back as far as the late 1860's,
and inferences may be drawn from a few writings prior to that
time. But until the studies of the last two decades, little reliable in-
formation was available on the population mechanics of tiie bird.

Mere fluctuations in population of a species does not in itself
indicate any tendency to a cyclic behavior. All species are constantly
changing their numbers, not only in detail but often in general
sweeping surges. On the mathematical law of probabflity, when a
species has a high population level (for its kind), it is most likely
to decline in numbers; and conversely, when it is scarce (but not
below the tlireshold of satisfactory mating), it is most likely to in-
crease. This is one more case of the natural law of biological equi-
librium commonly referred to as "the balance of nature."

In order to evaluate the evidence on grouse cycles, let us now
consider the many causes, features, and characteristics of cycles
and see how they fit into our knowledge of Bonasa umbellus.

1. Peaks and Troughs of the Cycle are Regularly Spaced in Years
in any One Portion of the Continent.

Years of scarcity of grouse in the Northeast are well documented
for 1927-29, 1924, 1915-17 (Stoddart, 1918), 1907 (Forbush, 1912),
1904 (Forbush, 1912), 1896-97 (Dillin, 1920). There is some evi-

Productivity and Populations 318

dence that grouse were also relatively scarce about 1877, and shortly
after the end of the Civil War. Years of abundance have been re-
corded as preceding these years in each case since 1895. There is
enough regularity to these dates, especially when the declines of
1904 and 1924 are segregated as being markedly less violent than
the others, to lead one to the conclusion that cyclic spacing is in-
volved. However, no such declines have occurred in the Northeast
from 1927 to 1944, although many relatively local scarcities have
occurred. A fairly general reduction of grouse in 1937 might be
classed in the category of the 1904 and 1924 declines but hardly
with the others. A general decline in grouse numbers has taken place
from 1944 to 1946, how serious we are not certain owing to war-time
cessation of grouse studies. It was bad enough to cause the Penn-
sylvania Game Commission to close the season in 1946. It may be
particularly significant that the failure of the expected major cyclic
decline of the 1930's to materialize occurred when, for the first time,
scientists were waiting ready to delve into its every manifestation.
Had no such critical group attended the grouse situation during
these years, a cyclic decline might have seemed more apparent in
many places. In fairness to the evidence supporting the concept of
a quick, immense decline, it should be pointed out that such a loss
of grouse did occur in the Minnesota-Michigan region in 1933-34
with grouse studies in progress there at the time.

2. Years of Decline in a Single Cycle will Vary with the Portion of
the Continent, the Progress being from Northwest to South and East.

This progressive, temporal sweep of progress of the cyclic die-
oflF, apparently a marked feature of rodent cycles, is not at all con-
sistent in ruffed grouse. In comparing the western and northern
cyclic years with those of the east, some are ahead in phase, some
behind, and some in harmony. Thus Criddle (1930), records low
population years in Manitoba as 1898, 1907, 1918 and 1928. Here
the 1907 record coincides wdth the eastern records, while the others
are one to two years behind. This is just the opposite of the general
cycle characteristic. Michigan cycles (Tubbs, 1940) have produced
low ebbs in grouse in 1901-02, 1915-16, 1923-25 and 1933-35.
With the exception of 1915-16, which is synonymous vdth the north-
eastern low, these troughs occinred ahead of the more eastern area.
This is the opposite of the Manitoba case. Wisconsin and Minnesota
grouse declined to a low ebb in the 1933-34 period, along with

3l4 The Ruffed Grouse

Michigan, but a corresponding die-off did not occur in the North-
east. Leopold and Ball ( 1931 ) generalized the cycle curve for the
whole continent but noted several exceptions, among them: the
Wisconsin low in 1919 is later than average; in 1905-06 a little ahead
of most areas; in 1899 again it is late, this time about two years.
Clarke (1936) gives cyclic low periods for Ontario as: 1933-35;
1924-25; 1914-16; 1904-06; 1894-95; 1883-85; 1874. These are con-
sistently a few years ahead of the dates for the northeastern states.
From all the information at hand we may conclude that the geo-
graphic lag from Northwest to South and East that is a characteristic
of true cyclic species does not hold consistently for ruffed grouse.

3. The Occurrence of Cyclic Trends in Different Animal Species
is Synchronized.

There is apparently a correlation in cyclic declines of many, if not
all, of cychc vertebrate species. When one species of rodent de-
clines in an area, others may disappear at the same time. Carnivorous
species may decline a year or two after the herbivores decline, after
the loss of their food supply takes full effect. Synchronism in the
latter case is the result of simple cause and effect: loss of food,
loss of life (see Elton and Nicholson, 1942; Chitty and Chitty,
1941). Elton (1942), in discussing wild-life cycles in Labrador and
Ungava, summarized it thus: "The lemming crashes; the predators
crash and migiate; the lemming begins to recover before the slower-
breeding predators can catch it up. This is a rough description of
a part of a phenomenon that is now known to be generally found in
nature . . ." In the case of unfform die-off, it must occur as a re-
sult of a common cause acting on more than one species.

This correlation in population trend between species was well ex-
hibited on the Connecticut Hill study area in 19-35-36. Red squirrels,
gray squirrels, cottontail rabbits and mice (mainly Microtus) were
from seventy-five to over ninety per cent lower in numbers the sec-
ond winter compared to the first. Predation was not primarily re-
sponsible, nor could significant disease conditions be correlated
(although our abihty to diagnose pathological troubles is admit-
tedly very limited). No answer was apparent in connection with
food, cover, weather, man, or any other matter that could be dis-
cerned. The cause, or causes, of this mass decimation remained
unsolved. It did, however, show up the simultaneous occurrence
of a precipitous decline in several species. The general thought

Productivity and Populations 315

has been that grouse follow the rodents by about two years in their
own die-ojff. This did not occur on this area in anything like a cyclic
decline although the grouse population did show a marked decline
in 1937.

The cause and effect synchronism in one group of animals is set
up by the loss of lemmings, white hares, and ptarmigan in the far
north. This dislocates the food supply of such predators as the snowy
owl, lynx, Arctic fox, and goshawk. A shortage in their food supply
causes them to migrate southward in search of better hunting, or
die where they are. Those that migrate come into the grouse range
and exert abnormal pressure on that species, which results in their
marked decline. There is no doubt that this chain of events occurs
to some degree. Southward winter incursions of snowy owls and
goshawks have often been recorded. A few of each species were
seen on Connecticut Hill in some years. And there is no doubt that
the goshawk is a very efficient grouse predator (see page 206).
Griddle ( 1930 ) suggests that a single goshawk may take as many as
fifty grouse in a winter. An "invasion" of these birds into a grouse
range already adequately populated with indigenous predators may
therefore have a significant effect on grouse numbers. Criddle rec-
ords that such an invasion occurred in Manitoba in the winter of
1907-08. This period was immediately following the drop in grouse
and could not have caused it, although, as Criddle says, "we . . .
need to know more of the influence of goshawks on a grouse popula-
tion already depleted by other agencies."

The evidence seems clear that the recorded declines in grouse
are not consistently synchronous with die-off periods in other
species. This adds weight to the evidence against the grouse being
typically cyclic in this respect.

4. Uniformity of the Die-off.

When a cyclic decline occurs it strikes aU coverts within the
broad area of effect, possibly continent-wide for some species some
times. King (1937), referring to grouse says: "Cyclic declines are
not the result of overpopulation . . . when the crash comes it in-
cludes all areas, those on which there are peak populations and those
on which there are very sparse populations." There is considerable
evidence to the contrary.

When grouse disappeared in Wisconsin in September 1933, it was
noted as "spotty" (Wisconsin Conservation Department, News Re-

316 The RufiFed Grouse

lease, October 18, 1933), and 'up to the present time it has affected
ruffed grouse principally, and on areas where the birds were most
plentfful ... In many instances (it) is affecting only townships
... In other counties practically the whole grouse population seems
to be affected . . ." Out of thirty-four counties submitting reports,
grouse were seriously affected in eleven, but were normally plentfful
or better in twenty-three.

Experience in New York indicates great local variation in popu-
lation trends. One portion of Connecticut HiU had a high popula-
tion in 1927 when most coverts in the region were at a low ebb.
Throughout the years of the detailed studies in several areas in
New York, some cover units would be at a peak while others were
low. Broad effects result from the cumulative effects of local condi-
tions. When the majority of coverts aie up, there is a pronounced
peak; when the greater proportion are down, there is a trough in
the general curve of population. But at any time there will be some
coverts, and some areas or regions, that are contrary to the general
condition.

Clarke (1936) concluded: "Periodically, ruffed grouse popula-
tions in Canada suffer serious diminutions in numbers. They do
not occur simultaneously throughout the whole country. Even in
Ontario there are local differences of at least three years in the
time at which diminution begins." Fisher (1939) says: "The fluctua-
tion in abundance of ruffed grouse is not uniform throughout its
range in Michigan. The birds may be increasing in some localities
while decreasing in others."

This evidence is supported by recent writings on the irregular
nature of mammalian cycles too. MacLulich (1937) found cycles in
varying hares to be regional phenomena arising from changing
local conditions. Cross (1940) concluded that ". . . periods of
maximum abundance in the nmnbers of (red) foxes in Ontario are
regional phenomena and not province-wide . . ."

5. Severity Varies with Location and Species.

The typical cyclic surges are most severe in the North and West,
less severe to the South and East, with any given species. Likewise,
the severity, that is the degree of loss in the die-off periods, is con-
sistently greater with some species than others, as for example with
the voles and lemmings in the north. It is most severe on range ex-
tended from its indigenous range.

Productivity aiid Populations 317

Testing these hypotheses with ruffed grouse is difficult owing
to the lack of adequate quantitative data. There is some indication
that grouse de<:imations may be more severe in the North and West
than in the East. Both King (1937) and Fisher (1939) have re-
corded losses exceeding ninety per cent for early fall populations
in Minnesota and Michigan respectively. The New York State hunt-
ing statistics between 1923 and 1935 indicate a decline of about
seventy-five per cent although this may be lower than the actual
drop, since no hunting was permitted in 1928 and 1929, years of
lowest abundance. No such severe declines have occurred in the
Northeast in the 1930's to correspond with the records from the
North-central states.

How the ruffed grouse die-offs compare with those of other species
as to severity is not wholly clear. There is little doubt but that they
are less violent than the decimations of some of the rodents, such as
the lemmings, the red and gray squirrels, and the snowshoe hare.
There is some evidence (Criddle, 19^30) that sharp-tailed grouse
fluctuations are more \iolent than those of the ruffed grouse. Other
species, as the bobwhite quail (in its northern range), the red fox,
and some other predatory species exhibit fluctuations alleged to be
cyclic in nature that are less severely marked than those of ruffed
grouse.

The contention that a cyclic species exhibits its most violent
changes in population at the periphery of its range, a doubtful
attribute at best, does not hold with grouse. The species is notably
stable in its southern Appalachian range. Chapman (1939) says:
"There is no evidence of a grouse cycle in Southern Ohio."

6. Cyclic Mortahty Primarily Affects Young Birds.

It is believed that all, or practically all, the losses of grouse in a
cyclic dechne are birds of the year. King indicates this to be true of
ruffed grouse and Green and Evans (1940) come to essentially
the same conclusion for snowshoe hares. This is likely to be true
whether the losses are cyclic or not. There are normally more young
birds than old, and the resistance to all decimating agencies is lower
with the young ones. This is truer when the birds are mere chicks
in the brood period. Abnormally high losses on the Connecticut
Hill area in 1933 and 1935 were both the result of losses in young
birds. This may not always be the case, however. The 1937 spring
population fell to the lowest level since 1931 as the result of winter

318 The Ruffed Grouse

losses. There was no evidence of an undue preponderance of birds
of the year in these losses. Since these cases appear to deal with
noncyclic losses, they do not disprove the point. Since the evidence
that has been recorded supports this characteristic, we may accept
it. Certainly it checks with the breeding density-recovery inverse
ratio principle.

7. Weather Conditions have no Correlation with Cyclic Grouse
Declines.

Griddle ( 1930 ) says "that rainfall seems to be a far less influence
in accounting for grouse fluctuation than is generally supposed."
Braestrip (1940) concludes that "there is probably general agree-
ment today that it is impossible to explain the phenomenon by direct
climatic influence on the animals, as was at first supposed." Clarke
(1936), Middleton (1934) for British grouse cycles, and King like-
wise come to the conclusion that climatic factors do not affect the
cyclic die-off.

Certain it is that some climatic factors, notably temperature and
precipitation, profoundly affect grouse survival. As we note in the
chapter devoted to weatlier in its effect on grouse, abnormalities
of weather conditions can bring about abnormally high losses of
grouse. These have been observed on Connecticut Hill both during
early summer ( on young grouse ) and in late winter.

It has been suggested that adverse weather just after hatching
time may cause starvation of the young chicks by killing off their
insect food supply. Counts made on Connecticut Hill showed a
normal average insect density in the zone available to grouse chicks
of around 300,000 per acre in typical grouse brood cover types.
Since a large share of these insects live in the duff on the ground and
are types (such as ants) that are not greatly affected by weather
changes, this suggestion does not seem to offer much likelihood of
importance.

8. Parasitic Infections and Diseases are not Correlated with
the Cycle.

Following an early enthusiasm for disease as an answer to the
cycle problem resulting from the work of Allen and Gross (Allen,
1928; Gross, 1925) in the mid-twenties, most workers have dis-
counted the importance of epidemic sickness in giouse. However,
Clarke (1936) attributes to a particular blood parasite, Leuco-
cytozoon bonasae, primary association with grouse cyclic losses in

Productivity and Populations 319

Ontario. Green (1935 et ante) has long held that disease plays a
dominant part in wild-life cycles, particularly documenting evi-
dence in tlie cases of snowshoe hares and ruffed grouse. Elton
(1942) pointed out the occurrence of disease in foxes and sledge
dogs in the North associated with cycles. He stated that ". . . the
cycle rests on mice or lemmings, . . . and is also marked by out-
breaks of epidemic nervous disease that devastate fox populations
and appear also among sledge dog teams . . ."

If cycles are a function of population density, then disease is
likely to play a large part. There is no question but that the in-
cidence of infestations increases with increasing grouse abundance.
This was clearly observed in the specimens taken annually on a por-
tion of the Connecticut Hill area. Not only does the incidence of
infestations of all sorts increase with growing populations but the
severity of the average case increases. For four years, from 1929
to 1933, tlie density of grouse increased on Connecticut Hill, par-
alleled by increasing incidence of infestation, before a case of
death from disease was found. That year several were noted. This
is significant considering the difficulty of finding grouse bodies be-
fore the scavengers get them. Likewise, there is no doubt but that
disease (lesions caused by Disphartjnx spiralis) was an important
cause, if not tire most important cause, of an abrupt loss of grouse
observed by Allen near Ithaca, N. Y., in the fall of 1927. The evi-
dence surely does not permit us to discard the disease factor in
major grouse declines.

9. Predator-Buffer Species Relations are not Primarily Correlated
with Grouse Die-off s.

Practically all authors of recent years have supported this con-
clusion.' Nevertheless, when an agency normally accounts for a
large proportion of a population, as predators do with grouse, a
delicate balance exists that may be seriously upset if conditions
temporarily favor an increase in predation. This occurs most com-
monly in winter and spring and is usually induced by climatic con-
ditions. The possibihties of increased predation on grouse result-
ing from losses in rodent prey species may be of considerable im-

1 It may be inferred from Elton ( 1942 ) that grouse may well fall victim to buffer-
predator population changes. He notes: "After the lemming crash, the foxes were seen
to have turned in the emergency to other supplies." Then: 'The vole cycle profoundly
influences some predatory birds. . . . These changes are reflected in the southv^'ard
migrations of several species. . . ."

■<£-

%:

320 The Ruffed Grouse

portance. Predators may not cause major grouse declines but in
most areas they play a part in implementing them.

10. Sunspot Cycles are Correlated with the Die-off in Grouse.

Among the first to suggest that sunspots might be correlated with
wdld life population fluctuations was DeLury (1930). The concept
of some direct action from the sun affecting animal hfe so fitted the
need for an all-pervading, mysterious something to explain the
newly recognized animal population cycles that it was grasped
quickly by many workers in the field. To make the solution complete
it was even felt ( or possibly hoped ) that in some unexplained man-
ner the recurring intensities of sunspots, with their effect upon the
ultraviolet and possibly other vitally important rays from the sun,
directly resulted in the decimation of vast miJtitudes of susceptible
animals. Such a theory is generally discounted today.

As the meteorological and biological specialists working in this
field progressed with their studies, it was evident that sunspot
cycles actually do affect Iffe on the earth but little agreement was
reached as to their relation to animal cycles. Effects upon the
weather, which in turn greatly affects animal survival, seem to be
clearly demonstrated. It is conceivable that the changes In the sun's
rays accompanying the sunspot phenomenon may alter the quality
in vegetable foods developed through photosynthesis. Braestrup
( 1940 ) gives evidence that mineral deficiencies may thus be caused,
deficiencies that may be vital to life. Wing ( 1935 ) , in studying wild-
Iffe cycles, came to the conclusion that "the sun is the dominating
factor in wild life," logical enough but not unveiling the modus
operandi of cyclic behavior. Clarke ( 1936 ) is among those who dis-
count any connection between the sunspot and wild-life numbers
cycles, as are also MacLulich (1936) and Cross (1940), Canadian
associates all. MacLulich, with convincing evidence, thoroughly lays
the ghost and concludes: ". . . the fluctuation in numbers of neither
lynx (Lynx canadensis) nor varying hare (Lepus americanus) are
correlated with sunspots. . . ." Elton (1942) likewise discounts
this theory, concluding ". . . There can be httle doubt that it is
wrong. . . ."

The shortcomings of the sunspot theory are many. In the first
place, the cycles do not always check by any means. The grouse
fluctuations are too erratic geographically to be guided by such an
all-pervading force. The geographical lag does not fit into the

Productivity and Populations 321

character of sunspot action. The fact that some grouse populations
are high when the general trend in an area is low does not fit the
pattern. And, except for its effect on animals through weather con-
ditions, the mechanics of sunspot effect on animal numbers leaves
a lot of explaining to be done. Nevertheless, there is enough meat
to the theory so that it cannot be entirely disregarded. Affecting
weather as they do, they must exert some effect on animal Iffe. The
possibility that the retarding effect of ultraviolet rays on plant
growth may affect the food supply of certain herbivores, thus in
effect causing a type of starvation, has yet to be proved or disproved.
It may have merit.

11. Numerous Other Suggested Causes of Cycles Warrant Less
Consideration.

The possibility that inbreeding may result in degeneration of the
stock is adequately taken care of by the fall shuffle of populations
which prevent inbreeding. Man's hunting is easily ruled out since
the cycle is effective in wilderness areas where no hunting takes
place. In fact, there is some indication that the periodic decimations
are more pronounced in extensive wilderness range than in broken-
up range in man-inhabited country.

Lack of food can hardly be considered seriously,^ except as al-
ready noted: (1) quality of food may conceivably be affected by
sunspot action; ( 2 ) adverse weather may affect insect food supply
of yoimg chicks. Criddle (1930) indicates a correlation between
cycles in grasshoppers and cycles in sharp-tail grouse, and suggests
there may be a correlation also with ruffed grouse cycles. However,
this seems remote when we consider the minor importance of grass-
hoppers in the ruffed grouse diet. Shelter likewise cannot be corre-
lated with abnormal die-off since these decimations seem to take
place in all types of cover and range. There is no evidence from the

^ Elton (1942) makes a good case for food scarcity being connected with tlie pre-
cipitation of cyclic decline of lemmings and voles in the north. "Lemmings must make
very great inroads upon the vegetation, which sets definite limits to the amount of
their increase and to their local distribution on the timdra. ... As the cycle in
population mounts steadily for several years, it reaches a breaking point and crashes.
The danger signals are not long delayed in a community dependent upon the ex-
iguous supplies of a northern land. Vegetation is denuded of the reserves built up in
^e previous time of lemming scarcity." Kenneth Doutt of the Carnegie Musetmi
says lemmings were abundant in Labrador when he arri\'ed there in Winter 1945.
By June most had died. Their remains were everywhere, and many were found dying.
There was plenty of food available. He believes lack of food could not have caused
the losses.

322 The Ruffed Grouse

eastern states that the bird's ability to reproduce is periodically
lessened. King advises (letter, October 5, 1943) that he did find
evidence of this condition in Minnesota.

SUMMARY OF ANALYSIS OF PERIODIC FLUCTUATIONS

From all of the information available and from the field studies I
made, the following conclusions seem to be warranted:

The ruffed grouse undergoes marked periodic changes in popula-
tion over its optimum range. These fluctuations are not sufficiently
regular to be caused directly by some all-pervading common cause.
They are inconsistent geographically, both in time of occurrence and
severity of action. The supposed Northwest to Southeast lag is not
consistent. The decimations that annually occur, whether normal or
abnormal, are the product of the interaction of environmental fac-
tors, primarily weather conditions, parasite and disease infestations,
and predation. These factors are conditioned by the habitat, by
variations in climate ( including sunspot cycles ) and, in some areas,
to a minor extent by man. These factors may individually or col-
lectively go berserk on occasion with resulting catastrophic losses of
grouse. The effect on grouse is sometimes felt only after a chain of
connected events is set up with other animals, such as the rodent
loss = predator's food scarcity == shffted predator pressure chain.
The rodent declines may be set up by food scarcity, may be con-
nected with chmatic factors, and are probably not directly correlated
with sunspots.

Grouse naturally have a high potential productivity which results
in high populations in favored years. High breeding densities then
lead to poor recoveries through high losses in young birds. This
mechanism, whatever may be the immediate decimating agencies,
seems basic to the periodic losses in this species. Further, this phe-
nomenon is not assumed from individual year's records, but is shown
to follow in series of years. It seems to have three phases, one of high
recoveries from moderate breeding populations, a second of poor
recovery from a high breeding population, and third, in occasional
periods, a continued poor recovery from low breeding populations.
Irregularity is a normal characteristic of grouse population changes.

Cause of grouse die-offs cannot be traced to a single source.
There is no simple answer. It is the result of a complicated interplay
between a multitude of natural factors and many kinds of organ-

Productivity and Populations 323

isms. It is probably not the result of any unknown agents but it is
brought about by little understood biological mechanics of well-
recognized factors. To say that there is much still to learn about this
subject is to state the obvious.

REFERENCES AND CITATION SOURCES ON PRODUCTIVITY
OF THE RUFFED GROUSE, AND ON CYCLES

Abbot, C. G. Solar Radiation and V^^eather Studies, Smithsonian Misc. Ck)ll.,

Vol. 94, No. 10, Aug. 15, 1935.
Allee, W. C. Animal Population Cycles, Scientific Monthly, Vol. 56, No. 6,

June, 1943.
Allen, A. A. Diseases of the Ruffed Grouse, Am. Game, March, 1928.
Allen, A. A. Sex Rhythm in the Ruffed Grouse and Other Birds, Auk, Vol.

LI, No. 2, April, 1934.
Anon. Grouse Cycle is Reached in 1933, mimeo. News Release, Wise. Cons.

Dept., Oct. 18, 1933.
Audubon, J. J. The Birds of America, 1856.
Baird, S. F., Brewer, T. M., and Ridgeway, R. liistory of North American

Birds, Vol. Ill, 1905.
Braestrup, F. W. The Periodic Die-off in Certain Herbivorous Mammals and

Birds, Science (New Series), Vol. 92, 2390, Oct. 18, 1940.
Bump, Gardiner. The New York Grouse Investigation, Proc. New England

Game Conference, 1933.
Bump, Gardiner. Some Characteristics of the Periodic Fluctuations in Abun-
dance of Ruffed Grouse, Trans. 4th, N. A. Wildhfe Conference, 1939.
Chapman, F. B. The Development and Utilization of the Wildlife Resources

of Unglaciated Ohio, Release No. 110, Ohio Wildlife Res. Sta., May 1, 1939.
Chitty, Dennis, and Chitty, Helen. Summary of "Canadian Arctic Wildlife

Enquiry, 1939-40" Journal Animal Ecology, 10 (2), Nov., 1941.
Clarke, C. H. D. Fluctuations in Numbers of Ruffed Grouse, Univ. of Toronto

Studies, No. 41, 1936.
Clayton, Henry H. Solar Relations to Weather, 1943.
Griddle, Norman. Some Natural Factors Governing the Fluctuations of Grouse

in Manitoba, Canadian Field-Naturahst, April, 1930.
Cross, E. C. Periodic Fluctuations in Numbers of the Red Fox in Ontario,

Joum. Mammalogy, Vol. 21, pp. 294-306, 1940.
Davie, O. Nests and Eggs of North American Birds, 1898.
DeLury, R. E. Sunspots and Living Things, Trans. 17th Am. Game Conf.,

1930.
DeLury, R. E., and O'Connor, J. L. Regional Types of Response of Wildlife

to the Sunspot Cycle, Trans. 1st N. A. Wildlffe Conf., 1936.
Dillin, J. G. The Effect of Forest Fires on Grouse, Bull. Am. Game Prot.
Ass'n, Vol. 9, No. 3, 1920.

324 The Ruffed Grouse

Douglass, A. E. Dendrochronology and Studies in "Cyclics," Univ. of Penna.

Bicentennial Conference, Phila., 1941.
Edminster, F. C. Man as a Predator on RuJffed Grouse Areas, Game Breeder,

Vol. XXXVII, No. 4, 1933.
Edminster, F. C. Productivity of the RufiFed Grouse in New York, Trans. 3rd

N. A. Wildlife Conf., 1938.
Elton, Charles, Voles, Mice and Lemmings, 1942.
Elton, Charles, and Nicholson, Mary. The Ten-year Cycle in Numbers of the

Lynx in Canada, Joum. Animal Ecology, 11 (2), Nov., 1942.
Errington, P. L. Winter-Carrying Capacity of Marginal RufiFed Grouse En-
vironment in North-central United States, Can. Field Nat., Vol. LI, No. 3,

March, 1937.
Errington, P. L. An Eight-winter Study of Central Iowa Bobwhites, Wilson

BuU., Vol. 53, No. 2, June, 1941.
Errington, P. L., and Hamerstrom, Jr., F. N. The Northern Bobwhite Winter

Territory, 1936.
Fisher, L. W. Studies of the Eastern RufiFed Grouse in Michigan, Tech. Bui.

166, Mich. State College, 1939.
Forbush, E. H. Game Birds, Wild-Fowl and Shore Birds of Massachusetts

and Adjacent States, 1912.
Forbush, E. H. Useful Birds and Their Protection, 1913.
Forbush, E. H. Birds of Massachusetts and Other New England States, Vol. II,

1929.
Green, R. G. Disease in Relation to Game Cycles, Trans. 18th Am. Game

Conf., 1931.
Green, R. G. Relation of Disease to Wildlife Cycles, Trans. 19th Am. Game

Conf., 1932.
Green, R. G. The Periodic Disappearance of Game Birds, Minn. Conservation-
ist, Oct., 1935.
Green, R. G., and Evans, C. A. Studies on a Population Cycle of Snowshoe

Hares on the Lake Alexander Area, Joum. Wildlife Mgt., Vol. 4, pp. 220-

238, 267-278, 347-358, 1940.
Green, R. G., and Shillinger, J. E. Wildlife Cycles and What They Mean to

the Grouse Supply, Trans. 20th Am. Game Conf., 1934.
Gross, A. O. Disease of the Ruffed Grouse, Auk, Vol. 42, No. 3, July, 1925.
Hatfield, D. M. Animal Populations and Sunspot Cycles, Chicago Naturalist,

Vol. 3, No. 4, 1940.
King, R. T. RufiFed Grouse Management, Joum. Forestry, Vol. XXXV, No. 6,

June, 1937.
Leopold, Aldo. Game Management, 1933.
Leopold, Aldo, and Ball, J. N. British and American Grouse Cycles, Can. Field

Nat., Oct., 1931.
MacLulich, D. A. Sunspots and the Abundance of Animals, Joum. Royal

Astronom. Soc. Canada, July-August, 1936.
MacLuhch, D. A. Fluctuations in the Numbers of the Varying Hare, Univ.

of Toronto Studies, No. 43. 1937.

Productivity and Populations

325

Middleton, A. D. Periodic Flunctuations in British Game Populations, Joum.

Animal Ecology, Vol. 3, No. 2, Nov., 1934.
Roberts, T. S. The Birds of Minnesota, 1932.
Sandys, E. Upland Game Birds, The Am. Sportsmans Library, 1902.
Smith, L. W. Records of Interest from Meriden, Gonn,, Auk, Vol. 38, No. 3,

July, 1921.
Stetson, H. T. Sunspots and Their Effects, 1937.
Studholme, A. J. Ruffed Grouse Environment in the Scrub Oak-Pitch Pine

Forest Type, MS. thesis, Pennsylvania State College, June, 1941.
Tavemer, P. A. The Ruffed Grouse and Island Populations, Gan. Field Nat.,

Vol. 54, Sept. 9, 1940.
Tubbs, F. F. Grouse Gycles, Mich. Gons., Vol. IX, No. 12, October, 1940.
Wilson, Alexander. The Natural History of the Birds of the United States,

Vol. VI, 1812.
Wing, L. W. Wildlife Gvcles in Relation to the Sun, Trans. 21st Am. Game

Conf., 1935.
. New York State Gonservation Department, Aimual Report, 1930 to

1940 inclusive, annually.

11

Management of the Ruffed Grouse

The grouse is essentially a bird of tlie woods. Hence its manage-
ment, so far as it concerns its habitat, must be woodland manage-
ment. To be intelligent this management should be derived from a
broader base of interest than tlie production of wood products alone.
It must be based as well on the whole ecology of the grouse; on its
cover preferences, food requirements, principles of interspersion;
the need for drumming logs, dusting sites, and escape shelter; and
the relations of the bird to other animals and to man. Merely the
considerations of timber growing will not suffice. The assurance of
a stand of the most valuable lumber species, well-formed trees prop-
erly spaced, handled by "sustained yield" methods will not auto-
matically care for the development or maintenance of good grouse
habitat. It can operate to depreciate the grouse cover value.

We have heard much in recent years of the conflict between game
and forestry interests. Some have held them to be irreconcilable.
This feeling was well summarized by Davenport ( 1941 ) : "By now,
I presume it is quite generally understood and taken for granted
. . . that stands of timber trees such as are in condition to produce
the most high-quality timber per acre per year are automatically
low in their game carrying capacity. In general, therefore, the prac-
tice of intensive forestry on any sizable area is due to eliminate
game management as a major consideration on that area."

I believe that this clashing is unnecessary and that we cannot
aflFord to continue it. Fundamentally it has resulted from a conflict
of personalities, one which will gradually disappear with greater
understanding. When we can have the wisdom to look at our forests
and woodlands for all that they are— sources of a great variety of
vegetable products from ferns to ship masts; sources of many valu-
able animal products; modifiers of local climate, watershed pro-

326

Management of the Ruffed Grouse 327

tectors, and soil savers; and finally, and sometimes most important
of all, a medium for escape and recreation where we humans may
build our health, our character, and keep from going insane— then
we may see the natural and logical integration of other interests,
including the management of grouse, with forestry.

Since we must begin woodland management with forestry as a
nucleus, we may say that grouse are a by-product of woodland man-
agement attained by modifying forestry practices in manners favor-
ing grouse to the fullest extent practicable without interference with
the major use. There will be some circumstances under which grouse,
or more likely game in general, will be the primary crop, with wood
products secondary, but these will not be general.

EXTENSIVE vs. INTENSIVE MANAGEMENT

There has been a great advance in recent years in our knowledge
of practices that can bring about a greater game supply. The game-
management methods advocated have been derived through "re-
search"—I put this word in quotation marks owing to the widespread
abuse it has received in the wild-life field— experiments directed
toward learning the needs of a particular animal or group of animals.
Having a restricted point of view, it is to be expected that recom-
mendations founded upon this type of investigations would conflict
with other interests. However, where game is the primary crop, the
major interest, the game management recommendations should gen-
erally prevail. This is intensive game management, where the ob-
jective is to produce the greatest possible number of the desired
game species almost regardless of other values, as on game preserves.

Extensive game management may be practiced where one or more
game species are desired in reasonable numbers along with other
products and values that are of primary interest. This type of opera-
tion implies merely the modification in favor of game of the prac-
tices used to produce the paramount crops, and possibly the addition
of some simple practices especially for the benefit of game. By far
the most of grouse management will be in this category.

It must be recognized that the distinction between intensive and
extensive management is not always great. There is a complete series
of intermediate degrees, meeting at the point where game and one
or more other products are of equal value as primary crops.

328 The Ruffed Grouse

Ruffed grouse management, like most practical game manage-
ment, must be largely of the extensive type for economic reasons.
Intensive management for grouse will largely be demonstrational on
public lands ( as Connecticut Hill is ) or on estates v^^here the value
attached to each grouse is very high.

To emphasize this point let us consider for a moment the eco-
nomics of grouse production. To paint the best possible picture, let
us assume optimum conditions. The highest shooting season popu-
lation possible is about a grouse to each two acres of cover ( we will
assume that only woodland and brush areas are considered). The
maximum possible safe kill is fifty per cent (and this only on the
assumption that winter losses are negligible ) , or the bagging of one
grouse for every four acres. If we assign a value of $2 for a grouse
brought to bag, we have a return of 50^ an acre. The investment
value of these lands may average around $10 per acre (from $4 or
less per acre for "skinned" and worn-out areas to very high values
for mature timber stands). The gross return from grouse would,
therefore, be about five per cent. This just about covers the interest
on the investment, but with no margin for development, mainte-
nance, taxes, or management costs. And no profit.

Moreover, such a return never would be attained. Even in the best
range the birds do not attain the production quoted, year after year.
There will always be considerable winter loss, hence the kill in fall
should be kept below fifty per cent. The desired kill will not be
returned from some coverts as it is not possible to hunt an area uni-
formly. Counterbalancing some of these factors, it may be contended
that $2 is too low for the value of a grouse. Well then, how much is
a bird worth? If one values a bagged grouse much over $2 he is
placing a high money value on the recreation enjoyed in pursuing
it. This is the condition I have already referred to as justifying in-
tensive management. But this will hardly be of interest to the run-
of-the-mill, "one-gallus" hunter, or to the average landowner.

Thus, most grouse management must of necessity be of the ex-
tensive type. This will be true on public lands if for no other reason
than limitations of funds. It will also be true of private lands with
the exception of some estates. The economics of grouse production
on private lands is further complicated through most lands in grouse
range being open to free public hunting, so that the landowner does
not get the full return that our discussion of economics assumed.

Management of the Ruffed Grouse 329

DEVELOPMENT OF HABITAT

The foundation of grouse production is the habitat. Hence ma-
nipiJation of the habitat is the core of management. In discussing
the various methods of developing and handUng grouse habitat, the
recommendations will be directed specifically at producing grouse,
but correlations with other interests will be indicated. The degree
of application will be suggested with all land management objectives
in mind so that it may be thoroughly practical.

In considering measures that may be used for the improvement
of grouse range let us assume that we are talking about a certain
type of range. The principles that are found to be good for this set
of conditions may be modified for other situations as needed.

Let us assume the land to be privately owned, in a hilly farming
area where the woodlands are on the back of the farms, overlying
property lines in masses of several hundred acres. The owner accepts
the concept of "multiple use," as to game and forest crops and he de-
sires to handle his woodlands so as to give the fullest measinre of
economic returns and social benefits. Being interested in birds as
well as trees, he is perfectly willing to concede the use of certain
small areas for brush cover, to keep his woods roads brushed out a
bit more than is ordinarily done, or to plant some shrubs— whatever
reasonable things the job requires.

WOODLAND PROTECTION

The first essential of woodland management is the protection of
existing values. Four types of protection are most commonly needed;
two of these being cared for by proper silviculture. Most important
and most commonlv lacking in many parts of the range are the ex-
clusion of domestic livestock and prevention of uncontrolled fire
( see Plate 41 ) . The other destructive elements are insects and dis-
ease organisms. The control of these infestations is a part of the
silvicultural handling of improvement and harvest cuttings.

We have already noted that livestock are incompatible with wood-
land preservation. The obvious remedy is to confine the animals to
their pastures by adequate fencing. To some farmers this seems to
be economically unfeasible, especially if the stock are sheep, which

330 The Ruffed Grouse

require a rather expensive woven-wire type of fence to hold them.
Surely, most farmers would not consider this fencing justifiable only
for improvement of the woodland for grouse. Fortimately, under
almost all conditions in the northeastern country, many other values
combine to make fencing between woods and pasture feasible. The
continued production of wood products of all kinds definitely re-
quires that glazing be kept out in order that ground cover and
reproduction of trees remain normal and healthy. Watershed pro-
tection is greatly enhanced by ungrazed woodlands in which the
forest floor is much more absorptive than in those trampled by stock.
Lastly, and most telling from the point of view of the farmer, it is
usually true that domestic livestock lose more than they gain from
woodland grazing. Poor forage and excessive exercise, combined
with increased diflBculties of control, make woodland pasturing a
losing proposition.

Occasionally wild herbivores, usually white-tailed deer in the
Northeast, may overbrowse the range with essentially the same
results as with domestic stock. The solution, however, is vastly dif-
ferent. Adequate seasons, bag limits, and hunters to reduce the herd
below the carrying capacity level can solve the problem in a single
shooting season, provided the public is not so oversold on the buck-
law principle as to refuse to allow the killing of does. After all, cor-
relating the kill to the production is merely a form of sustained yield
management.

The status of fire in woodlands in the Northeast is clear. All pre-
cautions should be taken to prevent the occurrence of accidentally
man-set fires, and adequate provision should be provided for the
efificient control of fires set by lightning, or made by man for legiti-
mate purposes. This principle does not in any way invalidate the
proper use of fire.

In extensive forest areas, often containing a high proportion of
public land, the setting of fires is most often by recreationists. Con-
trol is nowadays well organized in these areas and large areas are
seldom burned. Adequate trails for transportation, lookouts, tele-
phone and radio equipment, and other fire-spotting and fighting
facilities are generally well organized. In the noncontinuous range
the problem is different. Fires are most commonly the result of field
and hedge-row burning in connection with farm or road-maintenance
work. Control facilities are few and not so well organized. But the

Management of the Rulied Grouse 331

human population is denser and ti-ansportation facilities are good,
so that in tliese instances, too, fires do not ordinarily damage very
large areas. Farmers should be particularly careful with fires to
prevent such escapes. Burning of fields and hedge-rows is misdi-
rected effort and should be discouraged. And, of course, any use of
fire in the woods itself should be attended by the greatest care.

In some parts of the country where fires are most prevalent in
farming lands, especially in the Appalachian oak and oak-pine areas,
the use of cleared lanes for fire control is justified. The greatest need
in that section is for a change of habit on the part of the residents
with respect to burning the land. They should be taught and shown
that it does not pay.

IMPROVEMENT OF WOODLAND COVER

In forestry, the systematic tending of woodland is known as silvi-
culture. If the objective, or one of the objectives, is a crop of grouse,
and improvement work is directed partially toward that end, it is
still essentially silviculture, although some wild-life men, fearing a
confusion with timber culture, prefer to use other terms, such as
fericulture, game management, etc. The attempt to substitute other
terms implies separation from woods culture for wood products, and
such separation is not only undesirable but impossible.

"The ideal woodland for wild life . . . will be an uneven aged
stand, predominately second growth and immature ... a mixed
woods of both hardwood and conifers. ... It is well to maintain
a scattering of all species native of the type. ... All shelter and
food types needed . . . must exist within the area covered by an
individual of the species" (Edminster, 1941). This is in general con-
sistent with sound timber management, in that we need stands of
all ages to insure a continuing harvest and the inclusion of many
species for diversity of products. The degree to which the needs of
trees for wood products agree with those of grouse will, however,
depend considerably upon local markets, for markets are vital in
deciding what the crop plants should be. But market and other
problems merely modify the general forestry principle that a good
woodland is well-balanced with respect to both age classes and
species composition.

Determination of silvicultiiral practices to be followed in any

332 The Ruffed Grouse

given woodland first requires an analysis of the existing situation.
From the standpoint of grouse needs, the two primary considera-
tions are shelter and food. Other requirements must be fitted into
the framework of these basic facilities. These must be evaluated
with respect to quantity {i.e., extent), quality, and arrangement
( i.e., interspersion ) . Varying needs of the different seasons must all
be cared for. To illustrate, let us plan the development of a unit of
grouse cover; a hypothetical cover entity which provides all the
needs for a single grouse population group-a breeding pair in
spring and its subsequent production through the summer, fall, and
winter seasons, or any multiple of such a group occurring on the
unit. Of course this hypothetical area is useful only as a standard;
it never actually exists, nor is it feasible to attempt to reproduce it
in existing cover. It merely serves as a pattern by which we may
evaluate deficiencies in actual habitat and plan for improvement
measures.

A imit of grouse cover is shown diagrammatically in Figure 16.
Two basic variations are shown: the square unit, here measured
as twenty-five acres, which is most useful in analyzing cover that
has many and irregularly-shaped cover type subdivisions; and the
parallel strip unit of indefinite length, for application to extensive
coverts.

These cover unit diagrams indicate several basic needs: (1) All
seasonal cover requirements are included (conifers for shelter
needs; hardwoods for nesting and winter food; brushy areas of cut-
over or overgrown land for summer and fall food; open areas to
make edges); (2) proportions of basic seasonal cover needs are
indicated; (3) interspersion of types is idealized. Cover composition,
controlling quality of shelter and food conditions, is not included;
quantity and arrangement are.

In analyzing an actual area of grouse range we may consider the
quantity and arrangement elements first, the quality problem later.
First we must determine what cover types exist, and where. This
can well be done by making a map.' If aerial photographs are avail-
able (as they are for most of the Northeast today— inquire of the
nearest office of the Agricultural Adjustment Administration or Soil
Conservation Service ) , these are very useful for this purpose. Other-

^ For a detailed method of mapping forest areas for wild-life management, see
Wehh (1942 V

J

L

:•:•:•:• BRUSHY LAND 16"^ \::;l:'

HARDWOODS 40%
•;K\\^ 930'x 135' (10 Acres)

•;:.V: :i030'x 50' (4 Acres) .•::--:Tfrf^

A. I pOPEN LAND - 4% 1045' x 30' (1 Acre)* | f"

1% 6% 21.5%

43%

21.59& 6^0 Ifo*

Fig. 16. Hypothetical Unit of Grouse Cover. A. Square Unit
of 25 Acres. B. Linear Unit of Parallel Strips, 1400 Feet Wide.

" Only half the open land indicated is counted in a single
"unit," since each serves two adjacent units. This "open land"
may sometimes be part uf a slasfjing wfiich Is often rocut.

333

334 The Ruffed Grouse

wise a sketch map may be made by compass and pace measurement,
or by plane-table, according to the facilities and abilities of the
individual.

Upon measuring the amounts of the five basic cover types (see
page 63):— predominantly conifers; mixed w^oods; predominantly
hardwoods; bmshy areas; and open fields or lanes— the excesses and
deficiencies may be gauged. A balance between conifers and hard-
wood woods with from thiity to fifty per cent of each element is
very desirable. A good distribution of brushy edges and openings
covering about ten to twenty per cent of the area is very important.
Open land is not so important in quantity but is most valuable when
available in great length of edge in proportion to cover area. Some-
times cuttings, woods roads, streams, and lakes must substitute for
open fields in providing these edges.

In Fig. 17 we have taken an actual area of woodland and have
cover mapped it to evaluate the material with which we must work
(A), and have applied the principles of Fig. 16 to indicate the
changes in cover type arrangement that may expediently be carried
out to improve the area for grouse (B). Our area in this case consists
of one hundred and ninety-six acres of cover with eleven thousand
feet of open field edge, and continues on the south with more wood-
land cover across the property line. It is twenty-nine per cent over-
grown land, located in three large fields; fifty-seven per cent even-
aged hardwood stands that are deficient in shelter; and fourteen per
cent conifers, mostly in one large piece. There is no slashing since
the hardwoods were mostly lumbered thirty to fifty years ago, and
there are no mixed woods.

Our plan for adjustment of the cover arrangement and propor-
tions requires several changes. The objectives are an increase in the
coniferous element, and a better distribution of shelter and brushy
cover. The ratio of open edge to area ( feet to acres ) of fifty-six is
already good. The quantity of overgrown land is too large to main-
tain, hence a portion will be planned for maturing woodland.

It is not our purpose now to go into the details of the silviculture
involved in this particular job, except it may be said that it served
both wood production and game. Interplanting of conifers was made
in the overgrovm fields. A portion of the interior of the big hardwood
stand was spot-lumbered for selected wood products and the open-
ings planted to white pine and spruce. Improvement in distribution

[:::::::::1 Brushy Cover

[S\\\Ki Hardwoods
Mixed Woods
Coniferous Woods

Brushed out Property Line"*

Fig. 17, Plan for Cover Type Changes in a Woodland. A. The Woods before
Treatment. B. After Changes are Complete.

335

336 The Ruffed Grouse

of openings and brushy cover was accompHshed mainly by main-
taining favorably located portions of the overgrown fields, opening
up a woods road system, and devoting a narrow strip along each to
the production of wood products requiring frequent cutting. Along
the south boundary the property line was cleared as a part of the
road system.

As a result of these changes we find that the proportions as well
as the arrangement of the cover types have changed materially.
Brushy cover has dropped to nineteen per cent but is much better
scattered; hardwood stands have decreased to forty-two per cent,
while conifers are finally only thirteen per cent. The great improve-
ment in shelter came by the change to the mixed woods type, now
composing twenty-six per cent of the area. This, with the stands
of evergreens, makes a total of thirty-nine per cent of adequate
shelter.

One of the best indexes of improved conditions for grouse is the
increase in interior edge between cover types (only one side of
woods road openings are counted). In this case we had fifty-eight
thousand feet of interior type edge at the start, and at the comple-
tion of the change two hundred and one thousand feet, an increase
of three hundred and forty-six per cent.

INTERPLANTING OF CONIFERS

Changes in the woodland are ordinarily brought about by remov-
ing trees and planting trees. The former is generally the more im-
portant. Planting within a woodland is often unsatisfactory, and is
definitely limited in application. When extensive hardwood areas
exist that are seriously deficient in shelter, the underplanting or in-
terplanting of tolerant conifers may be advisable. Areas of open land
abandoned and overgrown with natural reproduction of hardwoods
may often be interplanted to advantage with conifers (see Plate
42). The need for that type of improvement must be judged in the
light of the probability of natural regeneration of these species. If
seed trees of desirable species are lacking, there is little likelihood
of natural establishment in the predictable future. In such areas
planting is necessary if coniferous shelter is to be had.

So far as practicable, plantings should be confined to open areas,
i.e., fields in herbaceous cover, or sparse, young, woody reproduc-

Management of the Ruffed Grouse 337

tion. But cases are common where there is httle opportunity for open
land plantings, or even if there is such opportunity the range is still
largely deficient in shelter. In these cases interplanting is the only
solution. Both game shelter and the diversity of wood products are
thereby improved.

Full advantage should be taken of existing crown openings in
which to plant. Here the young seedlings will have the most sun-
hght. Additional planting areas should be prepared by making
openings in the crown through removal of large-crowned "wolf"
trees, group selection cutting, or small clear cuttings. So far as pos-
sible this should be accomplished by making cuttings of marketable
materials in places that suit the needs of the wood harvest plan.
Planting should be made in the spring immediately following the
cutting. Plantings are not recommended under a full-canopied
woods.

Species of conifers for this use should be quite shade-tolerant.
Best in the Northeast for this purpose are the spruces (Norway,
white, or red), white cedar, and white pine. If the openings are
fairly large and the anticipated competition from hardwoods not
severe, balsam fir, red pine. Banks pine or other adaptable species
may be used. Spacing of plants should be the standard plantation
spacing, five to eight feet apart. The stock should be the best qual-
ity available, with an even balance of roots and top. This is gener-
ally a four-year-old transplant (known to foresters as a 2-2 plant,
indicating two years in a seedling bed, followed by two years in a
transplant line ) ff white pine, white cedar or spruce are used, or a
three-year seedling of the other species. Most state, federal, and
commercial nurseries handle these grades.

IMPROVEMENT OF SHELTER BY CUTTINGS

The improvement of shelter in pure hardwood stands as suggested
above requires considerable time. Even when the best of planting
stock is used and the soil is adapted to conifers it will be five to ten
years before their shelter will be of much value. In the interim,
temporary improvement may be gained through cuttings of the type
which foresters call "cleanings." Two methods may be used. One is
to half -cut small trees that are not needed for wood products. By
cutting each tree enough to cause it to fall to the ground, yet leaving

338 The Ruffed Grouse

some continuous bark and sapwood between top and stump, the top
will remain alive and produce a leaf crop. Being close to the ground,
these tops improve shelter for birds. Certain species are better than
others— as oaks (particularly pin oaks) and beech— since they hold
a part of their dead leaves on the tree through the winter. The sec-
ond method is simply clear-cutting units big enough to open the
crown. Species that coppice abundantly are best. Beech, oaks, and
the maples are all good. In about two years the sprouts will have
created considerable thick cover. Brambles, pin cherry or aspen
may come in quickly to add to the cover. The waste tops and
branches may be thrown over a few high-cut stumps in large loose
piles (see Plate 43). The result is a thicket of the slashing type
which has considerably more shelter value than the open hardwoods.
By the time these temporary values are lost, the conifer plantings
are beginning to be effective.

In stands that possess a scattering of conffers, improvement in
shelter value may often be brought about by careful cutting. If the
few conifers are old trees, a heavy seed crop may often be induced
by girdling a few of them in the spring. Hemlocks seem to be par-
ticularly adept at producing a swan-song crop of seed, the final
contribution of a dying matriarch. When the conifers are very small
their growth may be speeded by a light release cutting— just enough
of the surrounding plants removed to give the seedlings a little
direct sunlight. This practice is most effective with the native pines
and spruces. Caution should be exercised in this type of release cut-
ting for shelter, as it must be remembered that we wish the ever-
green branches to remain alive close to the ground. We do not wish
them to grow high and less useful as shelter too quickly. Hemlock
and spruce in particular should not be released once they are big
enough to furnish cover above the winter snows. They should be
allowed to remain under a canopy in order to retain their shelter
values as long as possible.

We must continually remember that for grouse the best winter
shelter is furnished by the evergreen woody plants, primarily the
conifers in the Northeast. From Pennsylvania and southeastern New
York southward the mountain laurel and rhododendron often make
a fair substitute when the pines and hemlock are absent. Even
among the conifers some are better than others. By species, the hem-
lock is clearly best, the spruces and white pine next, then balsam fir,

Management of the Ruffed Grouse 339

red pine, arborvitae, red cedar, and the hard pines follow in about
that order. In respect to age and size, the best winter cover is made
by an all-age stand in which young trees with low-hanging branches
are protected by other trees with sturdy boles. When the younger
trees are absent, the shelter is entirely up in tlie air and much less
useful. This condition often results from an overstocked stand in
which the lower branches soon die. Some of the pines, in particular,
are apt to develop this way. The solution is to thin the stand as it
ages in order to allow some light to reach through to the ground.
This may be accomplished by the orderly development of the crop
trees and the marketing of the small material for Christmas trees,
pulp wood, etc. The trees along tlie edges of the conifer clumps or
stands often are limby and of poor form for timber use because of the
extra light on the open side. They may well be allowed to remain
until mature for their game shelter value.

IMPROVEMENT OF FOOD BY CUTTINGS

While it is generally true that any grouse range meeting the
shelter needs of grouse will probably have an adequate food supply
for a fairly good population, the food problem must not be over-
looked. Most woods possess considerable grouse food at certain
seasons, but many are seriously deficient in the food requirements
for all the seasons.

Protection from domestic livestock is fundamental to a good food
supply. Without it the succulent herbaceous vegetation in the
ground cover will be destroyed. The second need in providing for
this vitally important phase of the woodland association is a moder-
ate crown density over much of the area— sixty to eighty per cent of
the ground shaded at midday. This condition is easily maintained
by selective cutting of crop trees and by needed thinnings or weed-
in gs. Thirdly, a well-balanced association as to species aids in pro-
ducing variety in the ground cover. Contrarily, single species stands
often have a poor understory.

The importance of the ground cover plants to the grouse is often
overlooked in considering woodland management, particularly since
direct control of these plants often seems impossible. Actually, they
may be controlled by adjustments in the tree cover. In our discus-
sion of the grouse foods it was noted that eight of the first twenty-

340 The Ruffed Grouse

five year-round important foods belong to the woodland ground
cover association, namely: sedges (Carex); bunchberry (Cornus
canadensis); strawberry {Fragaria); wintergreen (Gaultheria);
Canada mayflower (Maianthemum); partridgeberry (Mitchella);
ferns ( Polypodiaciae ) ; and sheep sorrel ( Rumex ) . Among the forty-
one genera of secondary food plants, twenty-one are in the ground
cover association, mostly in the woodland understory. Protection
from fire and grazing plus a controlled crown density are the man-
agement measures needed to maintain a supply of these important
foods.

A considerable part of the grouse's food comes from shrubs. Like
the ground floor herbaceous plants, shrubs in the understory of a
woodland must be managed by controlling the light conditions at
the canopy. Protection from grazing is quite essential for a satisfac-
tory shrub stand, since many of these plants are eaten by catde.
The same sixty to eighty per cent crown canopy recommended for
stimulating the ground cover is suggested for shrub development.
But, while shrubs are an important element in the whole woodland
plant association, they serve best as food when along the woodland
edges and in openings in the woods (see Plate 43B). It is here that
their fruiting is best and the production of buds most abundant.

Among the twenty-five primary food-producing genera, eleven
are predominantly shrubs or vines. The shrubs include most of the
dogw^oods ( Cornus ) , the hazelnuts ( Corylus ) , hawthorns ( Cratae-
gus), laurels (Kalmia), sumacs (Rhus), roses (Rosa), brambles
( Rubus ) , blueberries ( Vaccinium ) , and the viburnums ( Viburnum ) .
The vines include poison ivy {Rhus toxicodendron), greenbrier
(Smilax), and grape (Vitis). Of the forty-one genera of plants of
secondary value, thirteen are shrubs or vines.

The mountain laurel, flowering dogwood, and some of the blue-
berries and viburnums are prevalent in the woodland understory.
The others are predominantly found along the edges and in open-
ings. All fruit best where they receive plenty of sunlight. It is advis-
able then to develop shrubs mainly along the woodland borders and
in woodland clearings or lanes.

ESTABLISHING WOODLAND SHRUB BORDERS

A shrub border between the woodland proper and an open field is
a logical conservation development on farms. In addition to its wild-

Management of the Ruffed Grouse 341

life value the border of shrubs affords protection to the woods from
drying and damaging winds. It benefits the field crops by keeping
the trees far enough away to avoid root competition for moisture
and plant food, and the shading of tall trees. The width of such a
shrub border must necessarily depend upon the desires and needs
of the individual owner. A minimum of about twenty-five feet is
needed to maintain the association and to provide a gradual increase
in height growth from the low outside shrubs next to the open field
to the trees of the woodland proper. On large units of land managed
for timber and game these shrub borders may well be fifty or more
feet in width.

Two methods may be used to develop such a border: planting the
desired species in the open field edges; or cutting out the trees from
the forest border ( see Plate 44 ) . The planting problem will be taken
up later when we discuss development of open lands.

Ordinarily there are enough shrub seeds and seedlings available
in a woodland edge to take over the strip when the trees are re-
moved. If this is not true, they may be established by planting. It is
suggested that no planting be done for at least two years after
cutting in order to give full opportunity for germination of dormant
seeds.

Elimination of the trees is accomplished by cutting. All classes
should be removed or girdled. Many will sprout abundantly and
these will have to be cut back each summer until their vitality is lost.
This may take several cuttings, but in the interim the shrubs are
assuming dominance of the stand.

USE OF POISON IN ESTABLISHING SHRUB BORDERS

Those who may wish to avoid the repeated cutting necessary to
ehminate the hardwood trees from the borders where shrubs are
wanted may wish to kill them completely the first year by use of
poison. Sodium arsenite or white arsenite has proved satisfactory for
this purpose. The solution is made by mixing the chemical powder
with water. From one and a half pounds to three pounds of poison,
or even more, for each gallon of water may be used. The stronger
the solution the more quick and sure will be its killing effect. Since
some species are more resistant than others, and larger trees more
resistant than smaller ones, it is best to begin with a weak solution
and strengthen it as the results indicate the need. It is also advisable

342 The Ruffed Grouse

to add caustic soda (lye) to the solution in the ratio of one pound
to each four pounds of the poison. The solution may be kept in a
corked container until used up. Full respect should be paid to its
properties as a poison to animals as well as plants.

The poison solution is applied to cuts made around the base of
the tree at six-inch intervals. The cut should go well into the sap-
wood, and about a thimbleful or teaspoonful of poison solution
applied in each. The best season for this work is late summer.

An ordinary ax and oil can may be used for applying the poison
solution, but special tools are available for those who plan to do
very much of it. The "Cornell tree killing tool" is made by the Ire-
land Machine & Foundry Co., Norwich, N. Y. This instrument makes
the cut and applies the poison in one motion. However, it operates
with a valve that must be kept clean and pliable. The Council Tool
Co., Wananish, N. C, makes a special ax for tree poisoning which is
used in conjunction with an oil can applicator.

It is recommended that the poisoned trees not be removed until
completely dead. And again it should be warned that containers of
this poison should not be accessible to animals or left where children
might find them.

SHRUBS ALONG THE WOODS ROAD

The association of shrubs in the woodland interior that is analo-
gous to the shrub border is found only in openings of one sort or
another. The simplest way to develop a well-distributed brushy
cover is in connection with the woodland road system (see Plate
43B ) , ( page 335 ) . Every woodland needs roads to facilitate the har-
vest. By keeping them well brushed out, a narrow crown opening is
maintained through the woods. If more effort can be afforded here
for the improvement of game cover, these road openings may be
widened to thirty feet or even more. This is the easiest way of mak-
ing a little work go a long way in creating openings in the woods.
Experience indicates that an open lane, thirty feet wide, makes a
good set of edges and admits plenty of light for the development
of shrub cover. It is desirable to keep the road in the center of the
lane, or on the shady side.

Small openings in the woods may be made in connection with
various types of wood product cuttings. Improvement cuttings made

Management of tlie Ruffed Grouse 343

by the removal of large-crowned "wolf" trees leave scattered small
clearings which quickly develop into thickets of brambles, shrubs,
and sprouts. Where two or three such trees are removed close to-
getlier, an opening results that will be particularly valuable to
grouse as a feeding area. When removing large, badly formed trees,
caution should be used not to take good den trees, i.e., those with
well-developed hollows in the trunk or major branches. The squir-
rels, owls, and raccoons that a good den tree may support are worth
far more than its wood value. At least two per acre should be left
if they are available.

The group selection system of woodland harvest is conducive to
the improvement of food conditions in the woodland. Instead of
choosing crop trees singly, they are taken in small groups of three to
eight or ten (see Plate 45). When the group of trees removed is
small— up to about a quarter of an acre— the result is a woods with
many small glades. When the groups are larger— up to an acre or
more in size— the woodland is broken up with numerous small clear-
cuttings or slashings. These provide good grouse food conditions.

The two conditions most to be avoided are extensive clear-cuttings
and the opposite extreme of no cutting at all.

So far emphasis has been placed on the use of woodland-manage-
ment metliods for wood product harvest that will provide the best
distribution of openings and brushy cover. This is tlie most economi-
cal and practical method of improving and maintaining grouse cover,
and little or nothing need be charged to grouse management. If
more idealized grouse cover is desired, a deliberate system of cut-
tings for game management may be undertaken. Such cuttings will
seldom return their cost, for the added increment of grouse and
other game to the woodland income will not suffice. In circumstances
where the economics of cuttings for game management do warrant
it, a system of game clearings may be made.

CLEARINGS FOR FOOD AND COVER IMPROVEMENT

Studies have shown that grouse seldom penetrate a uniform wood-
land cover more than three hundred feet from an edge having some
brushy or open cover ( Edminster, 1935 ) . A plan for proper distribu-
tion of edges and clear-cuttings should thus provide that such units
be within six hundred feet apart to provide for full use by grouse of

344 The Ruffed Grouse

the intervening cover (Edminster, 1934). Slashings (used here to
mean small clear-cut units) cut specifically to improve the grouse
habitat provide better food conditions by permitting sunlight to
reach the ground and develop a suitable herbaceous and shrubby
cover ( see Plate 46 ) . As has already been noted, such cuttings may
also improve shelter conditions in areas deficient in conifers or other
evergreens. They may be handled in a system of imits cut in rotation.

In planning the location for a system of game slashings, the deter-
mining factors to be observed are: (1) Space the units about six
hundred feet apart, or from natural openings, for ideal distribu-
tion. (2) Cuttings are best when next to two or more other types
of cover. ( 3 ) Sites with good fertility will grow a good bushy vege-
tation cover quickly, while the poorer soils will return to wood-
land slowly. Therefore the units in the poorer soils will require less
upkeep cost. ( 4 ) Where possible they should be located conveniently
to the woods road system in such manner that the wood products
may be removed on skid trails running along the contour, thereby
reducing erosion. (5) Steep slopes should be avoided.

The units should be no larger than necessary to accomplish the
objective. One-quarter acre to one acre is usually adequate. The
cutting of the unit should be done so as to take advantage of the
natural features of the area. Avoid clumps of conifers but cut the
margin close to them. Leave seed trees of good food and shelter
species. Make the shape irregular, thereby providing more edge.

The maintenance of such a system of openings may be accom-
plished by the repeated cutting back of the area. This eliminates a
useful wood crop after the first cutting. An alternative is to allow
each unit to grow into cordwood before cutting, and rotate in a
series with other contiguous units. The latter method requires more
area to be devoted to this use, but it is better forestry practice. Each
unit is cut at the pole stage, usually twenty to thirty years old. The
cutting time of the several units is staggered five to ten years apart
so that the series consumes the entire rotation period. The length
of usefulness of a game clearing depends upon the fertility of the
site and upon the species that prevailed before cutting. Generally a
fertile site will lose its opening value in about eight to ten years;
poor soil sites last longer. The best gauge is the vegetation itself.
When the brambles, pin cherry, and similar pioneer plants are elimi-
nated, the site is no longer of much value as a summer-fall food area.

Management of the Ruffed Grouse 345

In making game clearings it is well to follow the general rules of
forest cover improvement, and remove the trees of least value and
retain the best. Since we wish to keep a few mature trees in any
clear-cutting, these should be selected from the best shelter and
food species. As a general thing, if the cutting is in a stand predomi-
nantly hardwood, favor the conifers, and vice versa.

When the usable wood products are removed from the cut-over
area, most of the slash may be disposed of by piling and then burn-
ing, preferably on snow. Some of the slash piles may be permitted to
remain as shelter for rabbits and grouse, say three or four piles per
acre. The burning will serve to stimulate the germination of certain
plants, thus adding variety to the cover. Among those commonly
taking over burns are the blueberries, scrub oak, and pin cherry.
Where burning is too hazardous, the surplus tops may be lopped
and scattered over the ground.

SELECTIVE CUTTING TO IMPROVE FOOD
CONDITIONS IN THE V^HOLE WOODS

If a woodland is being managed on a selective cutting basis, the
crop trees in an imeven-aged stand being removed when mature,
it is desirable to give some attention to the species that are impor-
tant in furnishing grouse food. These are mainly certain of the hard-
wood trees and they are primarily important as food sources in the
winter and early spring.

Unfortunately, many of the important trees that furnish these
foods are considered weeds by many foresters because they are im-
important in the present-day lumber market. Seven tree species are
among the twenty-five primary food producers, and seven more are
in the group of forty-two secondary food sources. Among these more
important ones, beech, oak, apple, and cherry furnish nuts and
fruit; the birches, poplars, apple, hophombeam, and cherries supply
buds. Only the oaks are consistently regarded as "crop" trees for a
woodland harvest. In some areas black cherry is made into lumber
and the poplars are used for pulp. The rest are merely tolerated or
are often "persecuted" by those who have visions of forests of pure
lumber. Apple, of course, is an exception since it is not a woodland
tree.

However, the concept of forests composed exclusively of the bet-

346 The Ruffed Grouse

ter hardwood lumber species is impractical. There is a need for
variety in the woodland, if for no other reason than as insurance
against excessive insect or disease loss. A woodland composed of
trees of all ages will contain many "filler" trees, those that are not
intended for ultimate timber harvest but are needed to maintain the
stand density as the crop trees mature. Much of this filler stand will
be used only for cordwood, and may as well as not be good grouse
food species.

I recall a recent visit to a forest being managed by a young pro-
fessional forester who was anxious to develop a game crop as well
as wood products. In some of his improvement cuttings he had re-
moved hophonibeam and beech in favor of elm and white birch.
In fact, he had planned to eliminate the hophombeam as completely
as possible, seeing no use for it. It was pointed out that the beech
and hophornbeam were as good as the others for fillers and were
much more useful as winter food for grouse. He immediately ad-
justed his cutting plan in this detail.

It should be particularly noted that those woodland tree species
that produce fruit or nuts, as the oaks, beech, and black cherry,
require much sunlight to seed properly. If they are crowded their
crops will be small or wholly lacking. Releasing them from the com-
petition of adjacent trees helps by allowing their tops to develop in
the open crown. If the trees to be cut in the release are of no value
they can merely be girdled with satisfactory results.

When one is concerned with a typical woodland of the Northeast,
say of the beech-birch-maple-hemlock association, where there is
plenty of opportunity for species selection in improvement cuttings
or thinnings, the question arises as to how many good bud-producing
trees are needed to support a satisfactory population of grouse
through the winter. Theoretically this could be computed mathe-
matically if we knew the average number of buds per tree in a
balanced woodland, the daily consumption of buds per grouse, and
the proportion of buds a tree may lose without serious damage. The
result would actually be conservative, for the birds do not feed
evenly over the whole woods. In portions in, or next to, good conifer
shelter the consumption would, no doubt, exceed the average rate,
and in areas remote from coniferous or brush cover, use would be
low. The application of the principle is obviously most important
in hardwood stands close to other cover, particularly coniferous

Management of tlie Ruffed Grouse 347

types. On the basis of an average daily requirement of four hundred
buds over a fourth-month intensive budding season, a stand of a
grouse per four acres would consume twelve thousand buds per
acre per season. If we assume a fifty per cent safe removal of buds
from the trees, and add another twenty-five per cent safety factor
to cover the additional birds that survive for only a part of the win-
ter, an annual requirement of thirty thousand buds per acre results.
Tliis need can be met with a minimum of three trees per acre of
average submature size. They should be of tlie better buds species,
particularly the following: apple, birches (except gray), cherries,
hophornbeam, and popples.

So far we have developed the silvicultural practices for game
management on the basis of furnishing shelter and food. While these
are the primary facilities of the environment, and provide a handy
objective for the discussion of management measures, they are by
no means the birds' entire needs. In discussing silvicultural prac-
tices it has been rather essential to consider the details of cover.
It is also necessary to fit these practices into the creation of suitable
range in a broader sense. The characteristics of the land cover must
exist in a well-interspersed arrangement of all the major cover
types: hardwoods, conifers, brushland, and open edges. We must
not lose sight of the cover type pattern while dealing with indi-
vidual trees.

PROVISION OF DRUMMING AND DUSTING PLACES

Among the needs of grouse are suitable drumming and dusting
places. In most good grouse range these needs are well met with-
out particular attention by man. But many woodland areas definitely
lack one or the other, and here it is well to give attention to these
requirements. If the woodland floor does not have a scattering of
old, large-diameter logs lying around, or moss-covered boulders, it
is advisable to leave lying a few butt logs from "wolf" trees, logs
tliat are of little or no value as a wood product.

Dusting spots are furnished either by exposed dry mineral soil
or by dry rotted wood at old logs or stumps. If these are lacking, the
turning up of mineral soil eveiy few hundred feet along the woods
roads or burning brush piles will aid in providing dust bathing
facilities.

348

The Ruffed Grouse

PLANTINGS AND SEEDINGS

The pioneers who cleared the land for farming in the Northeast
opened up some lands that were physically unable to sustain cul-
tivation or grazing, with the result that many thousands of acres I
had to be abandoned when they no longer could be economically
famied. Exploitive use and changing economic conditions caused
other farms to become submarginal to profitable agriculture. Many
of these lands are coming into government ownership, either through
purchase or tax delinquency. The proper use for most of these areas

Table 11. Important Site Requirements of Trees and Shrubs

FOR Plantations

Fertility

Moisture

require-

Geographical

Species

tolerance ^

ment^

range ^

Conifers:

Norway spruce

Well-drained

Low

All except coastal
plains— Pa., N. J.,
and north

Red spruce

WeU-drained

Low

Except coastal plain

Red pine

Dry or well-drained

Low

All except coastal
plain and low alti-
tudes south of Pa.

White pine

Dry to moist

Ixiw

All except coastal

Arborvitae

Well-drained to moist

Moderate

plain
All except coastal

plain

Hardwoods:

Red maple

Well-drained to moist

Moderate

Entire region

Sugar maple

Dry to well-drained

Moderate

Entire region

Black cherry

Drv to well-drained

Low

All except northern
Maine

White oak

WeU-diained

Moderate

All except northern
New England

Red oak

Well-drained

Moderate

All except northern
New England

^ Moisture tolerance is rated as: drtj, if adapted to a soil that normally becomes
crumbly moistureless in dry season; well-drained if the soil holds enough moisture in
dry seasons to maintain adherence of particles when squeezed; moist if the soil drips
when squeezed even in dry season.

^Fertility requirement is rated as: low if species will tolerate almost complete loss
of topsoil; moderate if considerable good topsoil is required for satisfactory growth.

" The range covered here is from Virginia and West Virginia northward to New
York and Maine, including adjacent portions of Canada.

Management of the Ruffed Grouse

349

Shrubs 6- Vines:
Bittersweet
Silky dogwood

WeU-drained Moderate

Well-drained to moist Low

Flowering dogwood Dry to well-drained Moderate

Graystem dogwood

Hazelnut

Hawthorn

Apple and crabapple
Bayberry
Virginia creeper

Bear oak

Dwarf sumac

Staghom sumac
Swamp rose

Movmtain ash

Nannyberry

Blackhaw

Highbush cranberry

Fox grape

Riverbank grape

Dry to well-drained

Dry to well-drained

Dry to well-drained

Dry to well-drained
Dry to well-drained
Well-drained

Well-drained

Dry to well-drained

Dry to well-drained
Dry to well-drained

Dry to well-drained

Dry to well-drained

Dry to well-drained

WeU-drained to moist

Dry to well-drained

Well-drained to moist

Low

Moderate

Low

Moderate

Low

Moderate

Low

Low

Low
Low

Moderate

Moderate

Moderate

Moderate

Moderate

Moderate

Entire region

All except northern

New England
Mass. to central New
York and south-
ward
All except Maine and

coastal plain
All except northern

New England
All except coastal

plain
Entire region
Entire region
Mass. to central N. Y.

and southward
Mass. to central N. Y.

and southward
All except northern

New England
Entire region
All except northern

Maine
All except coastal

plain
AU except coastal

plain
Mass. to central N. Y.

and southward
All except coastal

plain
All except coastal

plain
All except coastal
plain

seems to be to keep them in woodland, hence the former croplands
and pastures present a problem in re vegetation.

The same condition of unproductiveness from overuse is found
on parts of many farms that are, in their entirety, sound economic
imits. Whole fields or portions of fields that were inherently un-
suited to farming or that have been worn out by farming should
be retired to woody vegetation.

In both these cases the problems and principles of revegetation
are essentially similar. We will assume for the purposes of this dis-
cussion that the area is in grouse range and that grouse produc-

350 The Ruffed Grouse

tion is an important objective to be accomplished in connection with
planting the land. However, it should be noted that by far the
greater opportunity for grouse cover improvement Hes with existing
woods rather than in plantations.

There are certain basic principles that must be followed in all
plantation work. First, the species used should be well adapted to
the climatic and site conditions. Most important of site conditions
are the level of fertility and degree of dryness of the soil. These re-
quirements for all species recommended for use are summarized in
Table 11. When the correlations of range and site are cared for,
the next requirement is good planting stock: large and sturdy
enough to withstand competition, but not too large to handle
easily; well balanced as to root-top ratio; and properly cared for
after removal from the nursery. Finally, it should be set in the
ground with the roots well spread, and with the root-collar at the
ground line. In most planting work a large "scalp" of the sod should
be removed (at least twelve inches in diameter) and the plant set
in the middle of this area. Still better is a plow furrow every six
feet ( closer for shrubs ) , made on the contour. These enable the seed-
ling to make good grov^h before the competition of herbaceous
plants can retard it.

Pattern of Plantations. Plantations may be divided into types ac-
cording to the nature of the resulting cover. Three types of plantings
are needed for grouse: conifers, hardwood trees, and shrubs. These
correspond to similar natural stands in furnishing cover and food.
The first need in planting for new cover is to correct the deficiencies
in existing stands. In the Northeast the most prevalent shortcom-
ing is in coniferous cover. The greater part of the planting area is
likely to be conifers for this reason, as well as because several of
the conifers are among the most profitable for planting from the
forestry standpoint. In many areas there may also be a lack of ade-
quate brushy cover, or overgrown land. Shrub plantings may be
made to fill this need.

The positional relationship between the several types in the plan-
tation is of great importance. The outside of the planting area, that
is, where it bounds on open field, road, or other opening, should
receive the shrubs, thus creating a border. Behind the shrub bor-
ders a band of hardwood trees provides good nesting and winter-

Management of the Ruffed Grouse 351

feeding cover, while the interior of the unit should be dominantly
conifers. This principle may be followed either in a square unit or a
parallel band system (see Plate 50). When the planting unit ex-
ceeds twenty acres and is quite evenly proportioned in shape, it may
be desirable to consider the need of more shrub bands than merely
along the outside borders. Areas of forty acres or more may well be
planted in two or more units of bands and blocks of shrubs, hard-
woods, and conifers.

Integrally related to the pattern of large plantations is the need
for open land. So far as possible interior openings may be supplied
by roads maintained for transportation of products, administration
and patrol, and fire control. These roads through plantations and
woodland also serve somewhat as firebreaks and provide an oppor-
tunity to improve grouse food conditions. The need for roads of
this type depends upon the lack of open-field edges, pond borders,
and wide stream margins. When points in the plantation (or woods)
are three hundred feet or more from the outside, there is need for
open edges in the interior. Six hundred feet apart is as close as these
openings are recommended for grouse habitat.

The Coniferous Plantings. Reforestation has been practically syn-
onymous with conifer plantings until very recent years (see Plates
51, 52). The problem of reforesting public land had been attacked
almost entirely from the forestry and watershed protection points
of view. So habitual had the practice become that it was generally
advocated by sportsmen and others interested in wild life. Within
the last decade wild-life studies have indicated the limitations of
coniferous plantings as desirable cover for game (Edminster, 1935).
The principle of edges, so well enunciated by Leopold and other
pioneer wild-life ecologists, was gradually clarified as it applied to
reforestation work. It was shown that when a uniform type plan-
tation exceeds a certain size, its center becomes a biological desert
as far as game is concerned. The distance that a bird or animal will
habitually penetrate this cover varies with the different species.
Grouse are more tolerant than many. They utilize the outer two
hundred-foot band of conifer stand fully, and make considerable
use of the next hundred feet. Beyond this it is practically worthless
to them. We may then proceed on the basis that a coniferous plant-
ing, not exceeding six hundred feet in the narrow dimension, will

352 The Ruffed Grouse

be well used by grouse. Fortunately, most plantings will not ex-
ceed this size, the exceptions being mainly on public lands.

In planning the coniferous plantings, as well as others, it must
be remembered that a mixture of species is desirable. Single-species
plantings of any great size are an immense hazard. Just one bad
insect infestation and the whole thing may be lost. It's a good exam-
ple of the old adage of not putting all one's eggs in a single basket.
Don't base the future of your plantation on a single species.

The selection of species will be based mainly on the wood prod-
ucts desired. Not only is it a good idea to mix several kinds of
conifers in the planting, but it is also good for grouse to have some
clumps of hardwoods in the conifer cover. This may be achieved by
not planting spots aheady seeded to hardwoods, by leaving the
fail spots that are likely to occur on poor land (seventy per cent
survival may be considered a satisfactory establishment if quite
evenly distributed), by planting them in scattered clumps of ten
to one hundred each, or by leaving such spots unplanted and trust-
ing to natural seeding in places where desirable seed trees are
nearby.

Species of conifers best for grouse shelter are hemlock, white
pine, red, white, and Norway spruce, red pine, and arborvitae. All
are practicable to grow for this use or are purchasable at reasonable
prices. Other species may be used to give variety. Where enough
of those recommended are not adapted to the conditions, scotch,
pitch, jack, and Virginia pine, any of the other spruces, and firs, are
suitable.

The Hardwood Plantings. We have suggested that the best ar-
rangement for the planting of hardwoods is in a band between the
conifers and the shrub border. This is ideal from the point of view
of cover type arrangement, but is, of course, not essential. In large
plantings it may be more expedient to plant hardwoods in alternat-
ing bands with conifers (see Plate 50). The width of the band, what-
ever its arrangement with respect to the conifers, should be a hun-
dred feet or more in extensive reforestation work. In that arrange-
ment it will act as a break for crown fires. In small farm plantings
of only a few acres, it may be necessary to reduce this dimension
to fifty feet or even less, or handle the whole tree arrangement in
clumps. In a small planting there may be set out three or more rows

Management of the Ruffed Grouse 353

of hardwoods, then a few rows of conifers, and so on, in order to
develop a mixed stand. The total proportion of the planting to be
given over to hardwoods will depend largely on the need for bal-
ancing the existing woods and upon the type of wood products
wanted. The end result should be a fair balance between hard-
woods and conifers, the two taking eighty to ninety per cent of
the area, with the remainder in shrub borders and openings.

Estabhshment of the hardwoods stand will be guided by existing
seed trees and by site conditions. Many of the hardwoods are no-
toriously diflBcult to establish on poor sites by planting. As far as
possible the better, that is, the more fertile and well-drained soils
should be given over to hardwoods. While many hardwood trees
are slow and difficult to establish by hand planting, many establish
themselves readily by natural seeding or by sprouting. Popples, cher-
ries, ash, maples, and sometimes many others, are often found to be
taking over old fields in the Northeast a few years after abandon-
ment. If seed trees of satisfactory species are located within a few
hundred feet (or one hundred feet for the more heavily-seeded
species), we may count on nature taking care of the hardwood
planting for us. Under those conditions, merely refrain from plant-
ing conifers where hardwoods are wanted ( see Plate 53A ) . On the
other hand, if seed trees of desired species are not close by, the
probability of getting a stand of good hardwoods where you want
them is remote. Then it is best to plant.

A good mixture of several species is particularly important in the
hardwoods planting, both from the standpoint of ecology and from
the grouse food needs. Among the more important grouse foods that
come from trees, the most practical to establish by planting are:
the oaks (particularly white and northern red) (see Plate 54),
black cherry, and the maples, both sugar and red. Others may be
aided in establishment by planting seeds. A light spot-burning will
prepare the ground for the popples and pin cherry, and usually no
seeding will be required. The owner of the land will probably be
interested in other hardwoods too for their wood products or for
their value to other forms of wild life. Black walnut, hickory, but-
ternut, Asiatic chestnut, and oaks may be planted by using the
nuts— setting them twice the depth of their own long diameter in
the soil. (Here one should be cautious about rodent damage; plant
in years of low rodent abundance, or keep away from edges of

354 The Ruffed Grouse

existing woods. ) Black locust, basswood, ash, tulip and others may
well be included in the planting although of little food value to
grouse. These should be handled as nursery-grown seedlings, usually
one year of age. Occasional clumps of conifers may also well be
planted among the hardwoods.

The Shrub Border. The advantages of a border of shrubs along a
woodland margin have aheady been noted. The best way of insur-
ing a good shrub border on new woodlands is to provide for it
in the planting (see Plate 53B). This should be arranged on the
edges that are to remain next to an open field, road, or other open
area.

The width of the border will depend upon the desire of the owner
to provide this type of cover. At least twenty-five feet of the bor-
der is needed as a matter of good land use between fields and woods.
Beyond this minimum any extra width of shrubs is largely a matter
of planning for more game food cover. On public lands it may be
desirable to plant borders up to a hundred feet in width, using as
much as ten or fifteen per cent of the area. Planting of the taller
shrubs and small fruit-bearing trees should be at standard five to six
foot spacing. These species are most useful in the part of the border
next to the forest tree species. Here they will get plenty of light
for good fruiting, yet will not be in the way of the lower-growing
shrubs. The smaller shrubs may be placed next to the open field
and can be planted three to four feet apart. The rows may be the
same spacing.

By using lower shrubs on the outside, and slightly higher species
in each successive row, we may develop a graduated border that
will serve to protect the woods from drying winds, and the crop
field or pasture from the shading effects of the high trees and the
food and moisture competition of the tree roots.

Since a major purpose of the shrub border is to furnish summer
and fall food, we should select shrubs that best serve this use. Those
that are among the better grouse foods and are also practical to
propagate and plant are listed on the next page. As a general rule
it is well to use at least four species in each border. Most successful
of these species for planting are bittersweet, swamp rose, silky dog-
wood, graystem dogwood, hazelnut, bayberry, nannyberry, black-
haw, and highbush cranberry.

Management of the Ruffed Grouse 355

Low shrubs and vines Medium shrubs Tall shrubs and small trees

Bittersweet Silky dogwood Flowering dogwood

Virginia creeper Graystem dogwood Hawthorn

Dwarf sumac Hazelnut Apple and crabapple

Swamp rose ^ Bayberry Staghom sumac

Fox grape Bear oak Mountain ash

Riverbank grape Nannyberry

Blackhaw
Highbush cranberry

^ Multiflora rose is even better suited for plantations but no information is avail-
able as to its value in fimaisliing grouse food.

The thicker the shrubs develop the easier will be the maintenance.
But since this association of plants is not ordinarily a climax in the
plant succession, some tree species will encroach eventually even
if not at the start. These should be periodically cut out or poisoned.
This is preferably done when the adjacent woodland or tree plant-
ing is being thinned, improved, or market products removed. Little
extra effort will then be required to keep the shrub border in good
condition.

Open Edges. Any open area of herbaceous cover, water, or bare
ground next to woody cover furnishes an open edge. In grouse cover
these edges serve several important functions: food production,
notably through herbaceous plants and insects; keeping adjacent
cover exposed to the sun to the end that it produces more food; and
providing dusting and sunning spots.

Most of these edges will be supplied where farm fields meet wood-
lands and where roads traverse them. To the degree that fields con-
tinue to be farmed will most of these edges be maintained. We have
already stressed the importance of woods roads in the woodland
proper for the opening and cover change that they furnish. These
woodland roads offer an opportunity for improving grouse food con-
ditions by the simple expedient of disking the road center and
seeding with wild white clover {Trifoliurn repeiis). The soil must
be fairly high in lime content, or at least neutral, or lime will have
to be added to make this treatment successful. The seeding may be
made at a rate of about four pounds per acre, or one pound for each
thirteen hundred feet of seeding eight feet wide.

In extensive forest areas where open field edges are lacking, road-
way openings are particularly important. A good system of logging

356 The Riiffed Grouse

roads kept well brushed out will serve this need. Clover will often
come into these roadways naturally as a result of the horse manure
dropped during wood harvest operations. It then needs only con-
tinued sunlight to thrive and provide grouse with a source of
succulent green food.

PROPAGATION OF WOODY PLANTS FOR PLANTATIONS ^

Seedlings growoa in a nurser)^ are generally most suitable for
woody plantings. The problems of growing the trees and shrubs
for field use are many and can be discussed only briefly here. In the
following subheadings the essential steps in growing seedlings for
field use are summarized. Propagation information for trees and
shi-ubs recommended for plantings for grouse cover in the Northeast
are given in Table 12. It will be noted that with a single exception
(Norway spruce), all the species listed are natives. It is generally
best to use indigenous plants, although there are a few instances
where exotics may better serve the purpose.

Procuring the Seed. The first problem is obtaining good quality
seed. It is generally wise to gather the seed as near as possible to
the nursery or, if it is to be planted elsewhere, in the same climatic
zone where it will be used. Botli altitude and latitude of the seed
source should be as close as possible to that where the plants must
live. Only thrifty specimens should be utilized as sources of seed
as the character of the mother plant often has a marked influence on
the development of the transplants.

If the collection of seed from wild plants is not feasible, it may
be possible to purchase it through commercial seed dealers. In this
transaction it is well to state carefully the desired specifications as
to seed source and germination percentage.

The most difficult seed collection problems arise with the larger
trees, especially the pines, spruces, and maples. The seed should be
picked from the tree, often requiring a difficult climb. The opera-
tion should begin as soon as the seed is ripe in order to avoid losses
from shattering, or from animal feeding. Periodic inspection of the

^ This sectiun was written by Robert Thornton, formerly Regional Nurseryman,
U. S. Soil Couservatiou Service.

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358 The Ruffed Grouse

seed trees every few days during the ripening season will assure
accurate determination of the best collecting time.

Cones of the pines and spruces should be taken just before they
open. When squirrels begin cutting off the cones and dropping a
few to the ground, maturity of seed is indicated. Red pine cones
turn a deep purple, with brown on the scale tips at ripening, while
those of white pine turn yellowish-green with brown scale tips. Ma-
turity of these pine cones may also be ascertained by their specific
gravity. When the majority of red pine cones float in kerosene, they
are ready for picking. White pine cones may be picked when they
float in linseed oil. The best method of verffying the maturity of
spruce cones is by the general browning of the cone scales. Pine
cones can be removed most easily by using either hand or pole
pruners. The spruces and arborvitae are best gathered by hand pick-
ing or raking onto a tarpaulin.

Since maple seeds are winged and disperse quite widely upon
falling it is generally advisable to collect from the tree. In towns it
may be more eflBcient to collect from places of concentration such as
gutters. Tree collection is done by hand stripping from ladders from
the outside of tlie tree canopy.

Oak acorns can easily be raked up from the ground after they
have fallen. Since the acorns of white oak start germinating imme-
diately after falling, they should be gathered promptly.

Fruits, nuts, or seeds of most shrubs and vines may be collected
with httie trouble either from the ground or from the plants by the
aid of stepladders. Of the shrubs listed in Table 12 the hazelnut
and dogwoods require the most careful watching. The dogwoods
drop much of their fruit quickly upon maturing while the hazelnuts
are gathered quickly by squirrels. Fruits of the other species remain
on the plant long enough so that speed of collection is not lu-gent.
Rose, bayberry, bittersweet, and the siunacs generally remain for
some months while the others hang at least one or two months dur-
ing which collections may be made.

A word of caution in respect to handling quantities of freshly col-
lected fruit. Keep them well aerated, especially when in contain-
ers, in order to prevent decomposition through heating.

The times for collection of seeds given in Table 12 are general
and will vary some according to location and weather conditions.

Management of the Ruffed Grouse 359

By keeping notes on dates of maturity for a few years, these local
variations may be learned.

Cleaning and Extraction of Seed. Seeds should be removed from
their coverings before storing or planting.^ With the fleshy fruits
this may be done by maceration and fermentation. Some kinds, how-
ever, must be treated carefully. Mountain ash and crabapple de-
teriorate rapidly if they remain in the fermentation bath more than
forty-eight hours. Hence they should be very thoroughly macerated
before being subjected to fermentation. The fermentation bath may
be prepared by adding a small quantity of yeast to the water and
leaving it standing in a warm room.

The dry nuts are prepared by removing the husks (hazel) or
cups ( oak ) . Sumac "bobs" or heads should be broken apart before
removing the thin outer coat of the seeds. After separating the trash
from the seeds, the outer coat may be taken off by abrasion.

Extraction of seeds from cones should be done soon after collec-
tion is completed. This is most true of the pines and spruces which
become very pitchy ff allowed to he about. Arborvitae cones are
very dry and do not form pitch. The cones are placed in thin layers
on trays having screen bottoms for air circulation. They should be
kept in a hot dry room (with the temperahu-e not exceeding 120° F. )
and stirred occasionally. When the cones have opened, the seed
may be shaken from them. After the wings are removed they are
ready for either storage or planting. Sometimes the scales will stick
together through the first drying attempt. These cones should be
sprinkled with water and then redried, repeating the process as
many times as necessary.

Care of Seed Prior to Sowing. After the seed has been cleaned and
dried or extracted it is advisable to make a simple cutting or ger-
mination test at once. Germination tests immediately after seed col-
lection are not satisfactory for certain species having a period of
seed dormancy. In the cutting test, viable seed is recognized by the
clear white meat. If the interior is discolored, deep yellowish or
brown, the seed is considered nonviable. The percentage of seeds
that are viable as shown by cutting or by a trial germination is an
indication of production that may be anticipated. The actual num-

^ This is not essential for black cherry if it is sown immediately after collection.

360 The Ruffed Grouse

ber of seedlings produced, however, will be much lower than the
number of viable seeds as shown in these tests. Table 12 gives
average figures for viability tests and for the corresponding num-
bers of resultant usable plants per pound of seed. By comparing the
actual viabihty test witli the average, one may easily compute the
probable seedling production per pound of the seed on hand. The
rate of seeding may then be adjusted from that recommended in
order to compensate for the variation in viability.

Some seeds, such as tliose of the pines, spruces, arborvitae, and the
maples, are capable of immediate germination after maturing. All the
shrubs and vines under consideration have a period of dormancy
during which certain physiological changes take place prior to the
time when germination can take place. Also, some seeds have hard
coats that prevent absorption of moisture and thus delay germina-
tion. For those species that can be collected early enough for fall
planting, the over- wintering in the seed bed provides the necessary
conditions to carry them through the dormancy period. Those that
are to be spring-planted must be stratified over winter to prepare
them.

Seeds to be stratified should first be soaked for two hours in a
saturated chloride of lime solution to prevent molding. The medium
for stratification may be acid peat or an equal mixture of sand and
peat. The best medium for each species and the number of days
of stratification needed are listed in Table 12. The desuable tem-
perature for stratification is about 41° F.^ and the seeds should be
placed in thin layers in the shghtly moist medium. In handling large
kinds, tlie seeds may be mixed generally through the medium. For
very small seeds, as bittersweet and mountain ash, it is advisable
to keep a layer of cheesecloth above and below to facilitate recovery
at the end of the stratification period.

The hard-coated seeds of hawthorn and sumac must be scarified
before planting if quick germination is desired. The seed should
be thoroughly dried and then immersed in a solution of concen-
trated sulphuric acid ( H2SO4 ) . The liquid should cover the seed to
twice its depth and should be stirred occasionally for the twenty-
minute treatment. After this the acid is poured off and large quan-
tities of water are quickly added. The operator should be very care-
ful when adding the water because of the violent boiling that would

^ Mountain ash seed should be kept at about 32°— 34" F.

Management of the Ruffed Grouse 361

ensue if any acid remains, possibly causing acid bums. After the
seeds are thoroughly washed, add one tablespoonful of washing
soda for each quart of water to neutrahze the acid. Then rinse again
thorouf^hly in clean water after which the seeds are ready for
planting.

Bayberry seed should have the outer wax coating removed just
before stratification. This may be done by abrasion. Sand paper is
useful for this pinpose. Separation of the cleaned seed from the
granular bits of wax may be done through a screen.

The seeds of the northern viburnums also require special con-
sideration. They go through two stages of germination instead of
one. The root develops after a period of warm weather and the
leaves are not released until after a period of cold. The seeds should
not be stratified but rather held in dry storage until midsummer
of the year following collection when they should be sown.

Preparation of the Nursery Seed Beds. Selection of suitable soils
for the nursery beds is of the utmost importance. For all species the
soil should be deep and well drained but not drouthy. A light
sandy soil is the best for growing the conifers and small-seeded
hardwoods. Heavier loam soils are better for the large-seeded hard-
woods, although tight clay soils should be avoided.

Location of the seed beds should be made with great care. Sites
that can be worked early in the spring and soon after rains are best.
Friable soils free of stones make operations easier. Fields in sod
should be put through a cultivated crop before they are used for
seedling beds in order to reduce the weeds and white grubs, and
to eliminate clods.

The area should be deep-plowed and thoroughly haiTowed a
month in advance of seeding time so as to eliminate as many weeds
as possible. Do not use manure in the prepaiation of the beds. After
thorough disc-harrowing it is convenient to lay out the area in
standard beds four feet wide and a hundred feet long. Bomiding all
sides a pathway two feet wide should be left. Soil from the paths is
thrown up on the beds so as to make them four to six inches higher.
This insures good drainage and helps to prevent washing. When
the bed has been built up, it should be raked thoroughly, filling in
the low spots, and making a level surface. Beds should be raked
out a little wider than the four-foot dimension so that a full four-

362 The Ruffed Grouse

foot vWdth can be planted without breaking the shoulders. After
it is thoroughly raked and leveled, the bed should be rolled to
form a firm soil.

Beds to be used for conifers should have all nutrients added prior
to seeding. Never apply lime to conifer beds at any time. Those de-
voted to hardwoods may be top-dressed after seeding and at inter-
vals through the summer. Fertilization practices will vary with the
needs of the soil in each location and the rapidity of growth, and
should be about the same as for ordinary farm crops. Likewise, the
need for lime will depend on the present acidity of the soil and upon
the species to be grown. The conifers need an acid soil while the
hardwoods mostly prefer a neutral one. Bayberry is an exception.

Seeding and Seedling Culture. The time of seeding and rate of
seeding are given in Table 12 for each species. It is recommended
that the seed be sown by hand broadcast, mixed with sand to get
an even distribution where necessary, except v^th bear oak and
mountain ash. These two species should be seeded in rows about
ten inches apart.

The method of covering the seed of the conifers is to spread on
top of the seed a layer of sand %-% inch thick. For the hardwoods
the covering is either soil alone, or soil covered by a mulch. Clean
grain straw is suitable for the mulch. Soil covering depth is three
times diameter of clean seed or at least a quarter inch. The mulch is
removed when the seeds start to germinate.

Conifer beds should be covered with slat frames about a foot off
the ground during the early growth period to give about half shade
from the sun. The screens should be removed occasionally during
cloudy damp days to prevent the "damping off" of the young
plants. The frame covering is gradually abandoned after about mid-
summer.

One of the great problems in most nurseries is providing adequate
water during the dry season. Wliere operations are large enough to
warrant, an overhead irrigation system should be installed. For small
nurseries hand watering or a portable irrigation system must suffice
when rainfall is inadequate.

Competition of weeds with the young seedlings is a big problem,
especially during the first half of the summer. Continual vigilance
and lots of handwork are necessary to prevent them from stifling the

Management of the Ruffed Grouse 363

crop. Careful and frequent weeding during the early season will
reduce the problem to minor proportions in late summer.

As the seedlings develop, it is very important to reduce their
density to a desirable level so that each plant will have adequate
space and food. The optimum densities for each species are given
in Table 12. The thinnings should be started as soon as the plants
are large enough to begin crowding. For conifers this is three to
four weeks after germination; for hardwoods, after they form their
second true leaves. It is required to insure an evenly spaced stand.

Weeding ( or cultivation if row-seeded ) and irrigation should be
gradually tapered off beginning at midsummer and should be
stopped completely about six weeks before frost is expected. This
gives the plants an opportunity to harden before cold weather.

Various species of trees and shrubs require different periods of
time for the attainment of size suitable for field planting. A number
of the hardwoods grow to sufficient size in a single summer. These
are known as one-year seedlings and are designated "1-0," meaning
one year in the seedling bed and no transplanting. All the northern
conifers require more than one year and those recommended in
Table 12 require three or four years, depending upon the growing
conditions in the individual nursery and upon the weather. These
pines, spruces, and arborvitae are usually kept in the seedling bed
for two years and then transplanted in field rows. Two more years
as a transplant give the young tree its final designation "2-2." When
they are to be grown to usable size in three years, it is generally best
not to transplant.

Some of the hardwoods require two years and are designated
in Table 12 as "2-0." Others may sometimes, in good years, reach
usable size in one year and then in other seasons grow so slowly
as to require a second year. Those hardwoods requiring a second
year in the nursery may be transplanted to become 1-1 plants ff
desired. This is usually done only with beds of uneven growth where
some plants are ready for field use after one season while others
are too small. The small ones may be saved, after the bed is dug,
by transplanting.

The final column of Table 12 gives the smallest size recom-
mended for field-planting stock. The first figure is the height in
inches above ground, the second is the diameter of the stem one-
half inch above ground. In all cases it is assumed that the plants

364 The Ruffed Grouse

have a good root-top ratio, that is, roots about equal to top in
development. Stock may be laiger than the dimensions given as
long as the plant is small enough to be easily handled in planting.

CONTROL OF THE HARVEST

Proper and Flexible Laws. Any sound system of game manage-
ment requires a framework of good, workable laws in which to
function. Since the grouse is a very dynamic resource, the laws
pertaining to it should be flexible enough to provide for quick ad-
justments when needed. Fundamental is the centering of discretion-
ary authority for harvest limitations in the executive of the conserva-
tion agency of the state. This requires the delegation of this author-
ity by the state legislative body. And to be successful it requues an
intelligent, responsible, and independent executive branch backed
by an able staff of technical men.

Most state legislative bodies up to the present have been unwill-
ing to release this authority. But it is manifestly impossible for a
large pohtical body to gather the necessary facts that bear upon
hunting restrictions, seasons, bag hmits, and the like, or to take
timely action. When changes are needed they may be required
quickly and often at times when legislative action is out of the
question. The only sensible system is to place the responsibility
where it can be used with speed and be guided by impartial atten-
tion to the biological facts.

The histoiy of grouse laws, as with all game laws, is one of grad-
ually increasing restrictions on the freedom of the individual to
hunt. Changing conditions of habitat and human populations made
these laws a necessity if the sport and species were to be preserved.

However, it should be borne in mind that the purpose of game
laws is fundamentally to protect a living resource and to provide
for an equitable harvest of the annual crop. Both objectives are of
the utmost importance, but it is also true that they tend in opposite
directions. The greater the protection the smaller is likely to be the
harvest; and the bigger the harvest the more likelihood of endan-
gering the safety of the species. Game laws are therefore a compro-
mise. They should be designed to permit the greatest possible sus-
tained harvest without endangering the status of the breeding stock.

In states having adequate grouse range, there should be an open

Management of the Ruffed Grouse 365

season in most years. Only in years of catastrophic losses will it he
necessary to close the season— but then it should be closed quickly.
Ordinarily one or two years' protection are then needed to eflFect
sufficient recovery to warrant renewal of open seasons. With grouse
we should be able to anticipate an average of eight years or more
of open seasons out of ten.

The limitations on the individual in an open season must be de-
termined by existing conditions. In states like New York, New Jer-
sey, Connecticut and Pennsylvania where the hunting pressure is
generally intense, bag limits must be low. The present New York
law of three a day and fifteen a season is probably safe and yet
satisfactory to the fair-minded sportsman. Length of season is
the subject of divergent opinions in various states. Most experience
has shown that a season of moderate length produces little, if any,
more hunting than a short, intense season. In most places with good
range a season of at least a month is warranted. In years of good
abundance it is actually better to lengthen the season and harvest
a bigger crop. Two months of continuous hunting might be war-
ranted in some years, provided such a hberalization did not add
unduly to problems of conservation of other species.

I think we can fairly say that under modem conditions of hunt-
ing the sale of grouse and taking of the birds by means other than
gun or bow and arrow are definitely not warranted.

Refuges. In our discussion of man as a conservationist in his re-
lation to grouse, we pointed out that the refuge principle is not as
applicable in grouse management as it is to that of some other
species. Protection of this type during the breeding season is of
little value in most grouse range; if any protection is needed then
it is from cattle, fire, or lumbermen, not from the hunter. The es-
tablishment of grouse (or game) refuges to protect breeding birds
will rarely be justified. Protection from hunting through the means
of escape, or "seed stock," refuges will only occasionally be war-
ranted, and then in areas of high hunting pressure, or where there
is a combination of hmiting intensity with inadequate natural
escape cover. In many areas of grouse range it may be desirable
to establish refuges primarily for other game species. This discus-
sion does not refer to that problem.

Because of the rather short daily cruising radius of grouse, any

366 The Ruffed Grouse

refuges that are to benefit this bird should be small— say from ten
to one hundred acres. Witliin these limitations the exact size is not
of much importance. Of more import is the location of the refuge
with respect to cover. It should contain the best available shelter
and some good feeding area of the available fall foods. This usually
means that the refuge will be in tlie outer part of a woodland.

The distribution of refuges need be no more than one to three
per square mile under most circumstances. In range with rough
topography and variable cover, the pattern may be quite irregular.
The refuges should be placed where grouse are most sure to use
them regardless of whether this provides an even distribution.

Most of the intensive hunting in the Northeast is in the discon-
nected cover range, hence refuges wiU most often be needed on
small units of land ownership— the predominant condition in this
partly farmed country. A man who owns only a hundred or so acres
will not be able to give much consideration to the needs of a refuge
pattern. The most he is likely to require on his own land will be a
single unit. It is therefore quite largely a problem for the organized
sportsmen of the locality or for the state game agency to plan and
organize for tlie grouse refuge needs in any particular range.

The mechanism of making a refuge is simply the posting around
the boundaries of a unit of signs that will legally prohibit the use
of firearms therein. The type of sign, and the distance apart that they
must be posted, is generally specified in the state conservation laws.
When making a refuge, many states bound it v^dth a single strand
of smooth wire, hung about breast height. This is not essential, but
it is important that the boundary be apparent to one approaching it
if it is to serve its purpose. This means tliat expediency will often
alter the selection of its location and size from what might otherwise
be the more ideal. Roads, trails, woods edges, fence-rows, and sim-
ilar natural boundaries should be used so far as possible. When it is
necessary to have the line traverse woody cover it must be kept
brushed out if people are to see and respect it.

Use of Censuses. A census is essentially an enumeration of a pop-
ulation. In field studies of animal populations it is of fundamental
importance to know the number, type (sex, age, etc.), and location
of the individuals comprising the population. The greatest possible
accuracy is needed, therefore time and cost are not the most im-

Management of the RuflFed Grouse 367

portant considerations. The use of the census in practical wild-life
management is quite a different matter. It must be simple, quick,
and inexpensive. Its basic purpose is the same— to determine the
size and distribution of a population— but the accuracy required
is not so great and no supplementary data are needed. The use of
a management census is to determine the harvestable crop. This
may be the crop on a single area of cover, such as a private holding
or a public hunting ground, or the crop over a whole state. In the
latter case, the estimate would be based on sample censuses from
representative areas and would furnish the basis for seasons and bag
limits.

Accurate determination of a grouse population in most of the
northeastern range requires detailed coverage of the entire census
area by trained observers. In most circumstances this expenditure
is not warranted. The most expedient method is one of using indexes
and samples.

The period of census-taking for the estimation of shooting season
populations is from mid-August to mid-October. The middle of this
period falls in the height of the moult when the birds are difiBcult
to flush. It is generally best to avoid the month of September and
confine the counts to late August and early October.

One of the best indexes of grouse populations is the productivity
of the year. A comit of brood size in late August is a quickly available
method for gauging the productivity. If the brood size, female in-
cluded, averages four, the probability is for a population the same
size as the year before ( if other factors are equal ) . If the brood size
averages five or more, the probability is for an increasing popula-
tion; if three or less, the likelihood is for a decline.

The second figure needed in addition to brood size is brood den-
sity. To obtain a fair index of this figure the census should be made
in good weather: moderate in temperature for the season, at least
partly sunny, and with little or only moderate wind. The best hours
are from an hour after daylight to an horn* before sunset. The cov-
erage should be confined primarily to summer cover: brushy cover,
and hardwood edges. Covering fifty acres per man-day of eight
hours, an average of two broods per man-day would constitute an
average good year record. This, with an average unit size of five,
would give a full brood count of ten birds per man-day on fifty
acres. If the brood density is one per man-day on a fifty acres per

368 The Ruffed Grouse

man-day coverage, or less, tlie population may be gauged as quite
low, the degree by which it falls below the 1.0 average indicating
how low.

It is apparent that population estimates made in late August may
not trutlifully portray the condition of October or November. Of
course there is a normal loss during September and October that is
regularly anticipated, but there may be extra losses that would make
the August prediction for fall conditions invahd. However, this is
not commonly the case, as most of the serious losses occur in late
winter or in summer. Few serious late-summer, early-fall declines
have been observed in the East. But there apparently was a serious
autumn decline in at least some parts of New York in 1926, and a
similar one was observed in Wisconsin in September, 1933.

In view of the chance of error in depending upon an August brood
count for predicting grouse shooting season populations, it is a good
thing to check conditions again in October. At this season we can-
not use a brood count but must depend upon a time-index of adult
flushes. Again the work should be confined to balmy, calm days,
and to the best early fall coverts. These are nearly the same as those
used in the August count, but with more attention to the overgrown
shrub cover, including adjacent orchard and hedge-rows, and a lit-
tle less to the slashings.

The cover should be traversed systematically, as in August, but
whereas a close coverage of lines twenty-five to forty feet apart is
best for checking broods, lines of fifty to seventy-five feet are more
suitable for the autumn work.

The cover with a normally good brood count in August (two
broods per man-day vdth a ten-bird total) would ordinarily result
in an average rate of grouse flushes in October of fourteen per man-
day, excluding reflushes where determinable. Thus this figure may
then be used as a criterion for checking the population at this
season. The degree to which the average daily flush total varies
from this number may be taken to indicate whether a high popu-
lation or a lower-than-good population prevails. If the average flush
is seven or lower, the population is low and consideration may be
given to curtailment of the harvest.

It should be emphasized that the validity of the indexes used will
depend in good measure upon the conditions under which they are
derived, the equahty used in covering the area involved, and the

Management of the Ruffed Grouse 369

number of times it is covered. It is never safe to draw conclusions
from a single day's census. Of course tlie more times the area is cov-
ered tlie more accurate will be the average. As a general thing, four
surveys should suflBce.

If the area for which information on grouse populations is desired
is large, it may be well to census portions as samples and then apply
the data to the whole. Again, tlie accuracy will be greatest when we
cover most of the area; that is, get the best sample. We well know
that coverts only a mile or so apart will sometimes vary widely. In
using samples of cover, we must also be careful to select imits that
are fairly representative of cover conditions on the whole area.
When we attempt to gauge conditions in counties or states, the dis-
tribution as well as the adequacy of the samples used for censusing
is very important. Representative units of all types of range and of
all geographical areas must be surveyed, each with sufficient thor-
oughness to yield a reliable result. What is considered "reliable" will
depend upon circumstances. If a close knowledge of the population
of the coverts is needed, a sample of a hundred acres or more out
of each ten thousand is about the minimum. On a state-wide basis,
a group of four or five samples in each range type should suflBce,
if well distributed.

The application of the census information to local areas is in
safeguarding the grouse population during the hunting season. This
means hunting control.

Hunting Control. Control of hunting implies the limitation of the
harvest to a predetermined number of birds— within the framework
of applicable state laws, of course. Such control should be based
upon a knowledge of the hunting season population based on
censuses. The need for hunting control implies a very high hunt-
ing pressure. Otherwise there would be no need for this particular
attention. And it should be remembered that under normal circum-
stances in good grouse range, no special local control is needed. The
grouse is an adaptable creature an'd has a faculty for becoming more
wary as hunting pressure increases.

If the hunting take is to be held to a stated limit, the trespass
problem must be thoroughly under control. In most instances this
requires posting of the land under the legal requirements of the
state. It often means patrol in order to prevent illegal poaching.

370 The Ruffed Grouse

It means that some system of checking in the hunters must be
maintained and records kept of their bag. It may be that the hunt-
ing time or the take of each ehgible hunter will have to be appor-
tioned. In any case, the take will have to be kept to the desired
limit by stopping the hunting entirely when the full number of
birds has been taken.

If, with the approval of all these administrative problems and the
expenses they entail, we still find it worth while to insure a limited
grouse kill on an area, what take should be allowed? For a normally
good hunting season population of about a grouse per eight acres
or thereabouts, a take of twenty-five per cent is warranted. In years
of high populations when densities of a bird per four acres or better
are attained, the kill may safely be allowed to reach forty or even
fifty per cent. In seasons of low populations the kill should be kept
well below the twenty-five per cent figure, and if the density is lower
than a grouse per thirty acres, no hunting at all is warranted.

Predator Control. In our analysis of experimental work aimed at
an appraisal of the possibilities of increasing grouse by controlling
predators, the conclusion was that it not only doesn't pay; it doesn't
even work. This is true of various methods that have actually been
applied, especially bounty payments. When we add to this the very
serious questions concerning the unfavorable effects of predator
destruction on the rodent balance, the prospects for improving
grouse conditions through this medium are thoroughly discourag-
ing. And this in the face of the very important part that predators
play in decimating grouse. Even stripping our minds of all senti-
ment for the persecuted predators, we cannot honestly recommend
any systematic destruction of predators in the interest of the grouse.
This is especially true as regards the predatory birds, accurate con-
trol of which by species is itself impossible and indiscriminate kill-
ing very unwise.

The mammalian predators that have valuable furs not only can,
but should be, controlled (i.e., harvested) without regard to their
relations to grouse. Foxes, skunks, weasels, and raccoons are among
those grouse predators whose taking by trapping should be consid-
ered as a part of the wild-life harvest. Here, as with game, the take
should be confined to the annual increment in order that the stock
may not be endangered.

Management of the Ruffed Grouse 371

Restocking of Grouse. The hberation of game birds is usually done
in order to build up a population that has, presumably, been over-
shot, or has for some other reason been reduced below a safe sur-
vival level; or at least what the sportsmen think is a safe level. There
are many questions concerning the efficacy of this restocking which
we may not take time to discuss now, but it is clear that it can be
carried out successfully only with species that are easily produced
on game farms at a low cost per head. This is not true of the ruffed
grouse. It may conceivably be the case some day, but it is far from
true in the present state of our knowledge of grouse propagation,
and the prospects do not seem much brighter for the near future.
Thus we must dispense with any notion of building up the native
stock of grouse by liberations of more birds. Fortunately this is
seldom needed with this species for it has a great ability to recover
its numbers even when reduced to a remnant.

There is one place where the stocking of grouse is warranted;
namely, on areas of formerly good grouse range where the species
has been extirpated, where the cause of its extirpation has been re-
moved or corrected, and where there is no reasonable probability of
infiltration of grouse from surrounding range. Under these circum-
stances the reintroduction of grouse may prove well worth while.

As an example we can cite the occasional area of hilly or moun-
tainous land that has been subjected to repeated, complete burning
of the forest with consequent elimination of the grouse. If, in sub-
sequent years, people have left the region or have changed their
habit of burning the land so that this destructive influence no
longer prevails, it may be feasible to attempt the re-establishment
of the ruffed grouse.

In restocking such lands, either wild-trapped or hand-raised
grouse may be used. They will be expensive in either case. As far
as we know, either type is able to establish itself provided the speci-
mens are in good health. This is absolutely essential. If the birds are
even shghtly droopy or lazy when hberated they are almost surely
doomed.

The best time to stock grouse is at the very first break of spring.
At least four pairs should be stocked together in order to give fair
assurance of mating and nesting in the approaching spring season.
Since the birds will inevitably move away some distance, a few hun-
dred yards to several miles from the point of release, several pairs

372 The Ruffed Grouse

of birds in a vicinity will be needed to give assurance of establishing
a colony.

REFERENCES AND CITATION SOURCES ON
RUFFED GROUSE MANAGEMENT

Bump, Gardiner. New York's Ruffed Grouse Survey, Game Breeder, Vol. 36,

No. 2, Feb., 1932.
Bump, Gardiner. Forest Cover in Relation to Upland Game Bird Management,

Joum. Forestr)', Vol. XXX, p. 834, 1932.
Bump, Gardiner. Wanted— More Desirable Woodlands, Outdoor America, Dec.,

1935.
Bump, Gardiner. P'actors to be Considered in the Location and Development

of Artificial Slashings in Woodlands, Bur. of Game Circular No. 5 (mimeo.),

N. Y. S. Cons. Dept., March 23, 1936.
Chapman, H. H. Forestry and Game Management, Joum. Forestry, Vol.

XXXIV, No. 2, 1936.
Davenport, L. A. Timber vs. Wildlife, Joum. Forestry, Vol. 39, No. 8, Aug.,

1941.
Edminster, F. C. Man as a Predator on Ruffed Grouse Areas, Game Breeder,

Vol. XXXVII, No. 4, April, 1933.
Edminster, F. C. Developing Ruffed Grouse Areas, Trans. 20th Am. Game

Conf., 1934.
Edminster, F. C. Tlie Effect of Reforestation on Game, Trans. 21st Am. Game

Conf., 1935.
Edminster, F. C. An Analysis of the Value of Refuges for CycUc Game Species,

Joum. Wildlife Mgt., Vol. 1, No. 1-2, July, 1937.
Edminster, F. C. Improving Farm Woodland for Wildhfe, Soil Cons., Vol. IV,

No. 9, March, 1939.
Edminster, F. C. Wildlffe Management Through Soil Conservation on Farms

in the Northeast, Farmers' Bull. No. 1868, 1941.
Engstrom, H. E., and Stoeckeler, J. H. Nursery Practice for Trees and Shrubs,

U.S.D.A. Misc. Public, No. 434, 1941.
Forbush, E. H. Birds of Massachusetts and Other New England States, Vol. II,

1929.
Gabrielson, I. N. The Correlation of Forestry and WUdlife Management,

Joum. of Forestry, Vol. XXXIV, pp. 98-103, 1936.
Hosley, N. W., et al. Some Preliminary Game Management Measures for New

England Conditions, Report of New England Section Comm., S. A. F.,

Journ. Forestry, Vol. XXXII, No. 8, Nov., 1934.
Hosley, N. W., et al. The Essentials of a Management Plan for Forest WildUfe

in New England, Joum. Foresby, Vol. XXXIII, No, 12, December, 1935.
King, R. T. Ruffed Grouse Management, Joum. Forestry, Vol. XXXV, No. 6,

June, 1937.
Leopold, Aldo. Game Management, 1933.

Management of the Ruffed Grouse

373

McAtee, W. L. Forest Management and Wildlife Management, Trans. 1st

N. A. Wildlife Conf., 1936.
Miller, J. Paul. The Correlation of Game and Forest Management in New

England, Trans. 21st Am. Game Gonference, 1935.
Tourney, James W., and Korstian, Glarence F. Seeding and Planting in the

Practice of Forestry, New York, 1931.
Webb, W. L. A Method for Wildlife Management Mapping in Forested

Areas, Journ. Wildlife Mgt., Vol. 6. No. 1. Jan., 1942.

Index

References to authors cited are printed in all capital letters. Index items, other
than authors, are species of animals and plants, and subject matter. Indexed phrases
of two or more words are arranged in normal order, except where tliis might make
it difficult to find an item. Plants and animals are listed according to common names.
Scientific names are cross-referenced for plant genera, and for orders of insects used
as food by grouse. Disease agents are given by specific names.

ABBOT, 311

Abies. See Balsam fir

Acarina, 248

Acer. See Maples

Achillea. See Yarrow

Acorns, 105, 111, 128, 172

Actaea. See Cohosh, White baneberry

Adaptability to artificial propagation and

changing environment, 55
Agamodistomum sp., 237
Agrimonia. See Agrimony
Agrimony, 148, 170
Air-sac mites, 248
Alar tract, 14

Alder, 65, 91, 139, 154, 169
ALDRICH and FRIEDMANN, 7
ALLEN, 24, 25, 28, 29, 30, 32, 53, 56,

104, 148, 168, 171, 188, 224, 226,

227, 228, 229, 230, 231, 233, 235,

236, 238, 239, 241, 242, 243, 244,

247, 248, 287, 308, 318
ALLEN, E. A., 238
Alnus. See Alder
Alula, 14; coverts, 14
Alumroot, 68, 150
Amelanchier. See Serviceberry
American Game Protective Association,

23, 56
Amerindians, 1, 19, 254
Amphicarpa. See Hogpeanut
Andropagan. See Broomsedge
Angelica, 148

Antennaria. See Everlasting
Appalachian ruffed grouse, 6
Apple, 65, 105, 106, 107, 108, 112, 122,

159, 168, 172, 180, 345, 347, 349,

355
Apple budding, 180
Apple tent caterpillar, 311

375

Aquilegia. See Coliunbine

Arachnida, 107, 110, 167

Aralia. See Sarsaparilla

Arborvitae, 161, 337, 348, 352, 357, 358,

359, 360
Arctic fox, 315

Arctostaphylos. See Bearberry
Arrow-leaved tearthumb, 152
Arrowwood, 136, 158
Artificial propagation of grouse, 55, 371
Asarum. See Wild ginger
Ascaridia: bonasae, galli, lineata, 228
Asclepias. See Milkweed
Ash, 144, 161, 349, 354, 355, 357, 359,

360
Aspergillosis, 242
Aspergillus fumigatus, 242
Aster, 66, 140, 148, 169
Astilbe. See False goats beard
AUDUBON, 20, 255, 287
Automobiles, related to grouse hunting,

265
Avens, 141, 169
Azalea, 157

Bacteria, 226

Bags of grouse, daily and seasonal, 259

BAILEY, 7

BAILEY, W. W., 104

BAIRD, BREWER and RIDGEWAY, 287

BALL, 314

Balsam fir, 69, 159, 337

poplar, 125, 160
Banks' pine, 69, 86, 337, 352
Barberry, 140, 155
Barren strawberry, 66, 146, 154, 170
BARTLETT, 122
BARTRAM, 20, 55
Basswood, 161, 354

376

Index

BAUMGARTNER, 25

Bayberry, 65, 142, 156, 349, 354, 355,

357, 358, 361, 362

Bear oak, 127, 156, 345, 349, 355, 357

Bearberry, 155

Beaver, 114, 116, 127, 133

Bedstraw, 150

Beech, 68, 74, 88, 105, 108, 112, 118,

159, 169, 172, 174, 338, 345, 346
Beech-drops, 140, 148
Beggar-ticks, 148
Belmacanda. See Blackberry lily
BENNETT, 202, 203
BENT, 256
Berberis. See Barberry
Betula. See Birches
BEZDEK, 203
Bidens. See Beggar-ticks
Birches, 68, 74, 88, 105, 106, 107, 108,

112, 113, 159, 169, 172, 345, 346,

347
Bishop's cap, 142, 151, 169
Bitter nightshade, 144, 157, 170
Bittersweet, 140, 155, 349, 355, 357.

358, 360

Black bear, 119, 127, 128, 132, 135
birch, 113, 159
cherry, 68, 126, 160, 175, 345, 346,

348, 353, 357, 359
duck, 298

gum, 142, 160, 181
huckleberry, 156
locust, 160, 354
walnut, 353
Blackberry, 66, 107, 108, 132, 157
Blackberry lily, 148

Blackhaw, 136, 158, 349, 354, 355, 357
Blackhead. See Histoinonas meleagridis
Bloodroot, 153
Blue beech. See Hornbeam

cornel, 115, 155
Blueberry, 65, 68, 70, 105, 106, 107,
112, 134, 138, 158, 172, 175, 340,
345
Bluet, 150

Bobcat, 197, 208, 263, 264
Bobwhite. See Quail
Bonasa umhellus: affinis, 8; brunnescens,
6; canescens, 6; castaneus, 7; helmei,
7; incanus, 8; jobsii, 6; i7iedianus,
6; monticola, 6; ■phaios, S; sabini, 5,
17; thayeri, 5, 17; togata, 5, 17;
umbelloides, 5, 17; umbellus, 1, 3,
16; ijukonensis, 6, 17
Botrychium (Grape fern). See Fenis
BOUGHTON, 225, 228, 229, 231, 2-32,
233, 235, 236, 237, 238, 239, 2.51

Bounty systems, 215, 263

Brachylaemus fuscatus, 238

BRADLEY, 171

BRAESTRIP, 318, 320

Brambles, 65, 67, 90, 112, 132, 157, 169,

172, 175, 270, 338, 340, 344
Brassica. See Wliite mustard
Breeding refuge, 274
Breeding years, 289
Brome grass, 148
Bromus. See Brome grass
Brood cover, related to: openings, 80;

slope, 80; time of day, 79; weather,

80
Broods, number of, 291, 302; related to

cover types, 77
Brood spot, 16
BROOKS, 156
Broomsedge, 148
Brush piles, 345, 347
Buckwheat, 64, 141, 150, 169
Buffalo berry, 157
Buffer species, 215, 319
Bulbous buttercup, 152
BUMP, 56, 73, 77, 82, 186, 187, 239
Bunchberry, 68, 70, 115, 149, 340
BURNHAM, 24

Burning, affecting grouse food, 175
Burning bush, 155
Bush clover, 151

honeysuckle, 155
Buttercup, 66, 107, 110, 143, 152, 153,

169
Butternut, 353

Calamint, 153

Calls of mother and young, 39

Canada blueberry, 134, 158

lynx. See Lynx

mayflower, 68, 105, 107, 112, 121,
151, 189, 340

ruffed grouse, 5

spruce grouse, 22

yew, 158
Capilluria anrmlata, 231
Capital spaces, 14

tract, 12
Carex. See Sedges
Carpal remex and remex coverts, 14
Carpmus. See Hornbeam
Carpa-metacarpal coverts, 14
Carrion flower, 134, 153
Carrying capacity, 78, 298
Carya. See Hickory
Castanea. See Chestnut
Cat, 197, 208, 212, 269
Catbrier, 133, 157

Index

377

Catmint, 151

Catnip, 151

Caudal tract, 12

Ceanothus. See Jersey-tea

Cedar. See Juniper

Celastrus. See Bittersweet

Celtis. See Hackberry

Censuses of grouse, 366

Cerastium. See Mouse-ear chickweed

C eratophyllus sp., 248

Cestodes, 226, 234

Chamaecyparis. See Southern white cedar

CHAPMAN, 317

Cheilospirura spinosa, 230

Cherry, 105, 106, 107, 108, 110, 111,

112, 126, 160, 168, 172, 174, 270,

345, 347
Chestnut, 74, 140, 159, 169, 353
Chickweed, 153
Chicory, 149

Chiogenes. See Creeping snowberry
Chipmunk, eastern, 116, 120, 127, 132,

137, 197, 208
CHITTY and CHITTY, 314
Choanotaenia infundibulum, 236
Chokeberry, 139, 155, 170
Chokecherry, 126, 156
Chrysanthemum. See Oxeye daisy
Chrysogonum. See Goldenstar
Chrysosplenium. See Golden saxifrage
Ciclwrium. See Chicory
Cinquefoil, 66, 143, 152, 156
Cirsium. See Thistle
Cissus. See Heartleaf ampelopsis
CLARK, 63, 104, 209
CLARKE, 25, 225, 228, 229, 230, 232,

234, 235, 237, 239, 240, 241, 247.

248, 314, 316, 318, 320
Clearings for food and cover, 343
Climbing bittersweet. See Bittersweet
Cloacal infection, 244
Clover, 66, 145, 154, 169, 355
Club mosses, 69, 70, 151
Coccidia, 223, 238, 239
Cohosh, 148

Coleoptera, 107, 110, 146, 163
Columbian ruffed grouse, 8
Columbine, 148
Commercialism of grouse, 20
Common names of grouse, 2
CONOVER, 6
Contracaecum,, 233
Control of grouse harvest, 364
COOK, 202
COPE, 174

Coptis. See Tliree-leaved gold thread
Coralberry, 158

Com, 154, 169

Cornus. See Dogwoods

Corrodentia, 162

Corylus. See Hazelnut

Corynebacterium perdicium, 244

Cottontail rabbit, 114, 116, 118, 119, 123,
124, 130, 131, 132, 135, 137, 138,
215, 314

Courtship, 28

Cover, interspersion of, 70

Cover, related to: openings, 91; predator
mortality, 98; seasons, 82; slope,
93; time of day, 92; weather, 94

Cover types, 63, 73, 77; coniferous wood-
land, 69, 79, 80, 81, 86, 95, 173;
hardwood woodland, 67, 79, 81, 89,
95, 173; mixed woodland, 68, 79, 81,
88, 95, 173; open land, 64, 81, 91,
173; overgrown land, 64, 79, 81, 90,
95, 173; related to food supply and
predation, 172, 209; slashings, 66,
79, 81, 89, 95, 173

Cowberry, 135, 158

Cow wheat, 151

Crabapple, 122, 159, 849, 357, 359

CRAM, 226, 228, 230

Cranberry, 135, 158

Cranesbill, 150

Crataegus. See Hawthorn

Crazy flight, 43, 308

Creeping snowberry, 155

CRIDDLE, 313, 315, 317, 318, 321

CROSS, 316, 320

Crow, 197, 207, 212, 265

Crowberry, 155

Crucihulum. See Fungus

Crural tract and space, 14, 15

Cryptogamia. See Moss

Cultivation affecting grouse, 176

Cultural operations affecting grouse, 174

Cyathosoma striatum, 241

Cycles of abundance, 310, 322

Cyperus. See Galingale

Cytoleichus nudus, 248

Dandelion, 153, 169
Dangleberry, 156

DARROW, 104, 122, 128, 133, 139,
140, 142, 143, 144, 145, 146, 147
Daucus. See Wild carrot
Davainea tetraoensis, 235

proglottirm, 235
DAVENPORT, 328
DAVIE, 291
DAVISON, 146
Deadnettle, 151
DEARBORN, 202, 205, 208

378

Index

Decimating agencies, 300

Deer. See White-tailed deer

Deerberrv, 135, 158

Deervetch, 151

Degeeriella camerata, 248

DE LAHONTAN, 20, 21

DE LURY, 311, 320

Desmodium. See Tick trefoil

Dewberry, 107, 132, 157

DiervUIa. See Bush-honeysuckle

DILLON, 312

Diptera, 110, 146, 165

Disease agents, characteristics of, 225

Diseases, losses from, 308, 318; of the

ruffed grouse, 223, 250
Dispharynx spiralis, 24, 224, 226, 249,

308, 309, 319
Distribution of grouse, related to food

supply, 174
Dog, as grouse predator, 197, 208, 269
Dogs, used in grouse hunting, 21, 205
Dogwoods, 105, 106, 107, 108, 112, 115,

155, 168, 181, 340, 349, 355, 357,

358
DOUGLAS, 6
DOUGLASS, 312
Drumming, 28, 29, 30, 32
Drumming log, 33, 53, 326, 347
Duck millet, 149
Dust bathing, 54, 326, 347
DWIGHT, 11

EADIE, 203

Eastern hemlock. See Hemlock

red cedar, 159
EATON, 186

Echinochloa. See Duck millet
Ectoparasites, 226, 245
Edge, 20, 61, 74, 76, 355
EDMINSTER, 91, 99, 156, 164, 258,

267, 274, 277, 283, 296, 331, 343,

344
Egg-binding, 244
Eggs, 17, 287; fertility and hatchability,

288; mortality, 301; number in first

nests and renests, 287, 288
Eimeria: angusta, bonasae, and dispersa,

238
Elder, 65, 144, 157, 168
ELLIOTT, 255
Elm, 161, 346
ELTON, 314, 319, 320, 821
Empetrum. See Crowberry
ENGLISH, 202, 203
Enteritis, 244
Environment, related to productivity, 300;

limiting effects of, 309

Epifagus. See Beech-drops
Epigaea. See Trailing arbutus
Epilobium. See Willow herb
Epizootics, 223, 244, 251
Equisetum. See Horsetail
ERICKSON, 228, 231, 235, 238, 240
Erigeron. See Fleabane
ERRINGTON, 202, 204, 262, 282, 299
Eupatorium. See Thoroughwort
Evening primrose, 151
Everlasting, 139, 148, 150
Evonymus. See Strawberry bush, Burn-
ing bush
External parasites. See Ectoparasites

Fagopyrum. See Buckwheat

Fagus. See Beech

FaU shuffle, 43

False goats beard, 148

miterwort, 68, 108, 145, 154
Solomon's seal, 144, 153, 170

Feathers, number of, 16

Femoral tract, 14

Fencing affecting grouse, 176

Ferns, 67, 68, 70, 105, 107, 111, 112,
124, 125, 148, 151, 152, 153, 154,
156, 169, 172, 189, 340

Feverwort, 154

Field mouse, 215, 314
penny cress, 154

Filoplumes, 10

Fir, 86, 88

Fire, affecting grouse, 270, 330

Fireweed, 67

FISHER, 25, 225, 228, 229, 231, 232,
235, 238, 240, 241, 246, 296, 297,
816, 317

Flagellates, 240

Fleabane, 150

Fleas, as ectoparasites, 226, 248

Flies, as ectoparasites, 226

Flight, 52

Flukes. See Trematodes

Food, affected by snow and sleet, 189;
as limiting factor, 176, 321; compe-
tition for, 178; economic aspects,
179; primary, 113; quality of, 171;
related to cover types, 172; related
to grouse health, 178; secondary,
138; use by seasons, 103, 172; water
as, 182

Food types: utilized, 101, 171; emer-
gency, 172, 177; incidental, 171;
occasional, 147; preferred, 171;
staple, 171

FORBUSH, 23, 186, 188, 190, 191, 255,
259, 270, 287, 292, 312

Index

379

Fowl cholera, 245

Foxes, 55, 119, 138, 197, 198, 211, 213,

264, 265, 319, 370
Fragaria. See Strawberry
Frontal space, 14
Frostweed, 150
Fungus as disease agent, 226; as food,

149, 150

Galiugale, 149

Galium. See Bedstraw

Gallberr\', 156

Gallifonnes, 1

Gastropoda, 167

Gaultheria. See Wintergreen

Gaylussacia. See Huckleberry

Geranium. See Cranesbill

GERSTELL, 215, 263, 264

Geum. See Avens

GILFILLAN and BEZDEK, 104, 106,

116, 148
Gizzard worm. See Dispharynx spiralis
Gnaphalium. See Purplish cudweed
Goldenrod, 153
Golden saxifrage, 149
Goldenstar, 149
GRAHAM, 151

Grape, 65, 68, 105, 106, 108, 111, 112,
137, 157, 158, 159, 168, 172, 340,
349, 355, 357
Grasses, 150, 170
Gray birch, 65, 113, 159, 347

dogwood, 115, 155, 349, 354, 355, 357
fox, 116, 118, 123, 128, 132, 137, 138,

198, 212, 263, 264
rirffed grouse, 5

squirrel, 116, 117, 137, 215, 314
Greater primary and secondary coverts,

14
GREEN, 319
GREEN and EVANS, 317
GREEN and SHILLINGER, 243, 244,

245, 246
Greenbrier, 68, 105, 106, 107, 108, 112,
133, 134, 157, 169, 172, 181, 340
Gregariousness, 47
Grit, ingested, 180

GROSS, 24, 104, 108, 115, 142, 143,
147, 148, 162, 209, 224, 226, 227,
228, 229, 230, 231, 233, 235, 236,
238, 239, 241, 242, 243, 244, 246,
247, 248, 318
Groundsel, 153
"Grouse disease,** 244
Gum, 68
Guns used in grouse hunting. 265

Habitat development, 329
Hackberry, 159
Haemaphysalis chordeilis, 2A6

cinnabarina, 243, 246

leporis-palustris, 246

punctata, 246
Hamamelis. See Witch hazel
HAMERSTROM, 204, 299
HAMILTON, 201, 202, 205, 208
HANDLEY, 202
Hardback, 65

Harmostomum pellucidum, 236; sp., 237
HAWKINS, 282
Hawks, 43, 197, 206, 207, 208, 209, 211,

212, 213, 263, 264, 265, 315
Hawkweed, 150
Haws. See Hawthorn
Hawthorn, 65, 91, 105, 107, 108, 111,
112, 117, 155, 168, 172, 180, 340,
349, 355, 357, 360
Hazehiut, 65, 112, 117, 155, 340. 349,

354, 355, 357, 358, 359
Heal-aU, 143, 152
Heartleaf ampelopsis, 155
Hedeomn. See Mock pennyroyal
Heliantliemum. See Frostweed
Helianthus. See Sunflower
Hehninths, 226

Hemiptera, 110, 146, 162

Hemlock, 65, 69, 74, 86, 88, 161. 338,

352
Hepatica, 68, 150
Hepatitis, 244
Heterakis, honasae, 229

gallinae, 229
Heuchera. See Alumroot
Hickory, 68, 88, 159, 353
Hieracium. See Hawkweed
Highbush cranberry, 136, 158, 349, 354,

355, 357

Histomonas meleagridis, 241, 242

Hoary ruffed grouse, 8

Hobblebush, 136, 158

HODGE, 56

Hogpeanut, American, 154

Holcus. See Sorghum

HoUy, 141, 156

Homoptera, 162

Honeysuckle, 68

Hophombeam, 68, 105, 106, 107, 112,

124, 160, 345, 346, 347
Hornbeam, 68, 105, 140, 159
Horsemint, 151
Horse nettle, 153
HorsetaiU 150
HOSLEY, 104, 108, 114, 116. 117, 118,

119, 120, 123, 124, 128, 129, 130,

380

Index

HOSLEY, (Continued)

ISl, 133, 136, 137, 138, 139, 140,
141, 142. 143, 144, 145

Houstonia. See Bluet

Huckleberry, 65, 141, 156

Humeral tract, 14

Hungarian partridge, 135, 138, 282

HUNTER, 208

Hunting, 45, 256, 309; control, 369; sea-
sons, 256, 271, 272

Hymenolepis, carioca and microps, 234

Hymenoptera, 107, 110, 146, 166

Hypericum. See St Johnswort

Idaho ruffed grouse, 8

Hex. See HoUy

Impatiens. See Jewelweed

Improvement of woodlands, 331

Inbreeding, 24, 321

Incubation, 35, 290

Indian names, 1

Indians. See Amerindians

Infectious diseases, 242

Inferior space, 15

Insects, used for food, 146, 170, 180

Interplanting conifers, 336

Interspersion of cover, 70, 326, 332

Intimidation display, 28

Ironweed, 154

Ixodoidea, 245

Jack pine. See Banks' pine

Japanese honeysuckle, 156

JAYCOX, 6

Jersey-tea, 155

jewelweed, 68, 107, 110, 141, 150

JONES, 171

JUDD, 104, 105, 115, 117, 140, 147, 148,

162. 179
Juncus. See Rush
Juneberrv. 107, 110, 111
Juniper, 156, 159
Juniperus. See Juniper

Kalmiu. See Laurel

KELSO, 104, 106, 122, 128, 140, 143,

144, 145, 146, 147, 162, 181
KING, 25, 208, 287. 291, 297, 315,

317, 318, 322
Knntweed, 152, 169
KOZICKY, 201, 203
KRIEBLE, 270
KUHN, 104, 108, 118, 139, 141, 143,

144, 145, 146, 147, 148, 162, 181

Lactuca. See Wild lettuce
Lady's thumb, 152

Lamium. See Deadnettle

Laud clearing, affecting grouse, 267

Land management, for wildlife, 273

Large-tootlied aspen, 125, 160

Larix. See Tamarack

Lateral cervical space, 15

Lateral trvmk space, 15

Laurel, 105, 106, 121, 156, 170, 340

Law^ relating to grouse, 271, 364

Lead poisoning, 244

Lemming, 314, 316, 319, 320

LEOPOLD, 185, 219, 260, 277, 282,

292, 298, 300, 314
Lepidium. See WHd peppergrass
Lepidoptera, 107, 110, 146, 165
LESLIE and SHIPLEY, 223
Lespedeza, 68, 151
Lesser secondary coverts, 14
Leucochloridium pricei, 237
Leucocijtozuon honasae, 225, 239, 249,

318
LEVINE, 244

Lice, as ectoparasites, 226, 248
Life equation, 284
Hgustrum. See Privet
Limiting factor, 98, 176, 219, 250, 275
Linnaea. See Twinflower
Liverleaf. See Hepatica
Livestock, affecting grouse, 175, 268, 329
Longevity, 289

Lonicera. See Japanese honeysuckle
Lotus. See Deervetch
Louse flies, 247
Lower wing space, 15
Lowhop clover, 154
Lumbering, affecting grouse, 174
Lycopodium. See Club moss
Lynchia Americana, 247
Lynx, 208, 315, 320

MacGREGOR, 104, 139, 140, 142, 143,

144, 145, 146, 148
MacVICAR, 164

Maianthemum. See Canada mayflower

MalUu-d, 128

MaUophaga, 248

Malnutrition, 244

Malus. See Apple

Management of the ruffed grouse, 326;
clearings for food and cover, 343;
control of harvest, 364, 365, 366,
369; development of habitat, 329;
extensive and intensive, 327; im-
provement of food, shelter, and
woodland cover, 331, 337, 339, 345;
interplanting c-onifers, 336; open
edges, 355; plantings and seedlngs.

Index

381

S48, 351, 352, 354; predator control,
370; providing drumming logs and
dusting places, 347; restocking

grouse, 371; shrub borders, estab-
shment of, 340; along woods roads,
342; woodland protection, 329

Man's relation to grouse, 254; as con-
servationist, 254, 271; farmer, 254,
267; hunter, 254, 255, 263, 309;
limiting factor, 275; lumberman, 2.54,
267, 269; trapper, 254, 255. 263

Mapleleaf vibvumum, 68, 136, 158

Maples, 68, 88, 105, 110, 138, 159, S38

Marginal coverts, 14

Massachusetts Fish and Game Association,
24

MATHEWS, 32

Mating, 29, 287

McATEE, 2, 206, 207, 209

McDowell, 204, 206, 207

McLULICH, 316, 320
Meadow parsnip, 153

rue, 153
Meadowsweet, 157
Mecoptera, 164
Medicago. See Medick
Medick, 151
Megninia, 248

Metampyrum. See Cow wheat
Melilotus. See Sweet clover
MENDALL, 206, 208
Mentha. See Mint, Peppermint
Menziesa. See Minnie-bush
MERRILL, 56
MERRITTS, 104, 149
Mesoptiles, 10
Mice, 215, 314, 319
Microfilaria, 232
Middle primary and secondary coverts,

14
MIDDLETON, 318
Milkweed, 148
Milkwort, 143, 152
Mink, 208, 263
Minnesota ruffed grouse, 6
Minnie-bush, 142, 156
Mint, 151

Mitchella. See Partridge berry
Mitella. See Bishop's cap
Miterwort, 68, 108
Mites, as ectoparasites, 226, 248
Mock pennyroyal, 150
Monarda. See Horsemint
Moneses. See One-flowered pyrola
MONON, 255
Moose, 114, 135, 137
MORLEY and WETMORE, 244

Mortality, 76, 215, 256, 301, 303, 304,

306, 307, 308, 309
MORTON, 20
Mortis. See Mulberry
Moss, 149, 151
Moult, 11, 43
Mountain ash. See Ash

holly, 103, 156

laurel, 68, 107, 112, 121, 156, 338,
340
Mourning dove, 128, 130, 138, 298
Mouse-ear chickweed, 149
MOUSLEY, 189
MUELLER, 230, 235, 236, 237
Mulberry, 159
Multiflora rose, 355
MURDOCK, 171

Nannyberry, 68, 136, 158, 349, 354, 355,
357

NELSON, 104, 106, 134, 139, 140, 141,
142, 147, 148, 181

Nematodes, 223, 226, 233

Nemopanthtis. See Moimtain holly

Neossoptiles, 10

Nepeta. See Catnip

Nest desertion, 53

Nesting, 34, 53, 73, 74, 290; mortality,
301, 302; related to openings and
slope, 76

Neuroptera, 162

New England Ruffed Grouse Investiga-
tion, 24

New York game kill records, 217, 256

New York State Conservation Dept., 23,
25

NICE, 50

NICHOLSON, 314

NICHOLSON, S. B., 311

Nortliem ruffed grouse, 6

Norway spruce, 337, 348, 352, 356, 357

Nova Scotia ruffed grouse, 5

Number of grouse harvested, 255

NUTTALL, 20, 23

Nyssa. See Black gum

Oaks, 65, 68, 74, 88, 106, 108, 112, 127,
156, 160, 168, 175. 338, 345, 346,
348, 353, 357, 359

Oenothera. See Evening primrose

Oil gland tract. 12

Olympic ruffed grouse, 7

One-flowered pyrola, 151

Onoclea. See Ferns

Openings related to grouse, 76, 80, 91

OpUiones, 107, 110, 146

382

Index

Opossum, 119, 123, 128, 131, 132, 135,

138
ORD, 6

Oregon ruffed grouse, 5
O'Roke, 240

Orthoptera, 107, 110, 147, 162
Osmunda. See Ferns
Ostrya. See Hophombeam
Osijspirura petroioi and mansoni, 232
Owls, 197, 203, 209, 211, 212, 213, 263,

265, 315
Oxalis. See Wood sorrel
Oxeye daisy, 149

PALMER, 271

Panic grass, 152

Panicum. See Switchgrass, Panic grass

Parasites of ruffed grouse, 223, 249

Varthenocissus. See Virginia creeper

Partridge berry, 69, 70, 105, 107, 112,

123, 156, 168, 340
Pasteurella aviseptica, 245

tularense, 243
Pasture gooseberry, 157
Peach, 126, 160
Pear, 160
Pedal tract, 14

Pennsylvania grouse kill records, 256
Peppermint, 151
Peritonitis, 244
Persicaria, 152
Phaseolus. See Wild bean
Pheasant, relations to grouse, 55
PHILLIPS, 266, 270
Physaloptera, 232
Picea. See Spruce
Pin cherry, 65, 66, 67, 91. 126, 160, 175,

338, 344, 345, 353
Pine, 74, 160, 358, 359, 360
Pinus. See Pine
Pitch pine, 69, 86, 160, 352
Pkintago. See Plantain
Plantain, 152
Plantings and seedings, 348, 350, 351,

352, 354, 355
Plant succession, 71
Platanus. See Sycamore
Plum, 126, 160
Plumage, 4, 9, 11
Plumulae, 10
Pneumonia, 244

Poison ivy, 68, 107, 129, 157, 340
Poison used in silviculture, 341
Polygala, 70, 152
Polygda. See Milkwort, Polygala
Polygamy of niffed grouse, 287
Polygonatum. See Solomon's seal

Polygonum. See Smartweed

Polypodiaciae. See Ferns

Polypodium. See Ferns

Polystichum. See Ferns

Poplar. See Popple

Popple, 67, 68, 91, 97, 105, 106, 108,

112, 125, 160, 169, 172, 174, 175,

270, 338, 345, 347, 353
Population declines, 190
Population densities: of game birds, 298;

of grouse, 292, 293, 294; of nests,

292
Population, related to productivity, 292;

to weather, 190, 307
Population trends of grouse, 218, 278
Populus. See Popple
Porcupine, 119, 131, 137, 197, 208
Potentilla. See Cinquefoil
Poultry, relations with grouse, 242, 269
Prairie chicken, 282
Predation, 196, 307, 309; as limiting

factor, 219, 309, 819; related to

cover types, 209
Predator control, 211, 263, 370
Predators, agent of sanitation, 210; of

grouse, 197; related to nest cover,

76
Primaries, 14
Privet, 156
Productivity of grouse, 277, 281, 293;

related to environment, 300
Promiscuity of ruffed grouse, 287
Propagation, artificial, of grouse, 55, 371
Propagation of woody plants, 356; age

and size of stock for planting, 363;

care of seed, 359; cleaning and

extracting seed, 359; preparation of

seed beds, 361; secairing seed, 356;

seeding and seedling cvilture, 362
Prosthogonimus macrorchis, 237
Protozoa, 226, 238
Prunella. See Heal-all
Prunus. See Cherry
Pterylography, 9
Pterylosis, 9, 12
Ptilosis, 10

Ptycfwstoma honasae, 241
Purphsh cudweed, 150
Pyrola. See Shin leaf
Pyrus. See Pear

Quail, bobwhite, 116, 118, 119, 120,
123, 124, 127, 128, 130, 131, 132,
135, 137, 138, 282, 298, 317

"Quail disease," 244

Quaking aspen, 125. 160, 175

Quercus. See Oak

Index

383

Rabbits, 215, 345

Raccoon, 119, 123, 127, 128, 135. 137,

138, 197, 208, 212, 264, 370
Radicula. See Water cress
Raillietina tetragona, 236
Range, daily and seasonal, 51; geo-
graphic, 16; types of, 60
Ranunculus. See Buttercup
Raspberry, 66, 107, 108, 110
Rectrices, 3, 12
Red-backed mouse, 215
Red cedar, 339

fox, 123, 127, 128, 182, 138, 198, 212,
263, 264, 317

grouse, 223

maple, 159, 348, 353, 357

osier, 115, 155

pine, 65, 86, 88, 337, 348, 352. 357,
358

ruffed grouse, 5

spruce, 337, 348, 352, 357

squirrel, 123, 131, 132, 138, 197, 208,
212, 215, 265, 314
Reforestation, 91, 351
Refuges, 273, 365
Relations to other animals, 55
Renesting, 53, 302
Reptilia, eaten by grouse, 167
Reptilian grouse predators, 209
Restocking grouse, 371
Rhododendron, 144, 157, 338
Rhus. See Sumac
Rhynchosia, 153 *

Ribes. See Pasture gooseberry
Ring-necked pheasant, 116, 118, 119, 120,
123, 124, 127, 128, 130, 131, 132,
135, 137, 138, 282, 298
Roads, affecting grouse, 265
ROBERTS, 303
Robinia. See Black locust
Rosa. See Rose
Rose, 65, 105, 107, 112, 131, 157, 169,

181, 340, 358
Roundworms. See Nematodes
Rubus. See Brambles
Ruff, 1, 12

RUHL, 258, 260, 261
Rumex. See Sheep sorrel
Ruptures, 244
Rush, 151
RYDER, 256

St. Johnswort, 150, 156
Sale of grouse, 21, 271, 272, 273
Salix. See Willow
Sahnonbeny, 132, 157
Sambucus. See Elder
SAMUELS, 188

Sanctuaries, 273

SANDYS, 186, 303

Sanguinaria. See Bloodroot

Sanicle, 153

Sanicula. See Sanicle

Sarsaparilla, 154

Sassafras, 160

Saturation point, 298

Satureja. See Calamint

Sawbrier, 133, 157

SAWYER, 39

Saxifraga. See Saxifrage

Saxifrage, 153

Scapulars, 14

Scotch pine, 352

Scrub oak. See Bear oak

SEALANDT, 208

Secondaries, 14

Sedges, 65, 107, 110, 112, 114, 148,
149, 172, 340

Sedum. See Stonecrop

Seed-stock refuges, 274

Semiplumes, 10

Senecio. See Groimdsel

Septicemia, 244

Serviceberry, 68, 139, 159

Sex ratio, 291
rhythm, 30, 287

Sexual relations, 287

Sharp-tailed grouse, 317, 321

SHAW, 1

Sheep sorrel, 66, 105, 107, 108, 110,
111, 112, 133, 153, 340

Shelter, 60; as limiting factor, 98; re-
lated to adults, broods, nesting, and
openings, 73, 77, 80, 81, 91; related
to predation, slope, time of day and
weatlier, 76, 79, 80, 92, 93, 94, 98

Shepherdia. See Buffalo berry

Shin leaf, 68, 143, 152

Shortleaf pine, 69

Shrew, Blanna, 208, 215

Shrub borders, 340; established by cut-
ting and planting, 354; use of poison
in establishing, 341

Silky cornel, 155, 349, 354, 355, 357

Silviculture, 331, 337, 339, 341, 345

Siphonaptera, 248

Skunk, 123, 132, 135, 137, 138, 197,
205, 211, 212, 213, 264, 370

Skunk cabbage, 68, 145, 153, 169

Slope, related to adult grouse, broods,
and nests, 76, 80, 93

Smartweed, 143

Smilacina. See False Solomon's seal

Smilax. See Greenbrier

SMITH, 287

SMITH and BYER, 175

384

Index

SMYTHE, 17, 32, 104, 105, 106, 108,

115, 128, 139, 140, 141, 142, 143,

144, 145, 147, 148, 162, 181
Snakes, 197, 209
Snares, 265
Snowberry, 158
Snow-roosting, 46, 108
Snowshoe hare, 114, 123, 131, 133, 215,

319, 320
"Snowshoes," 4, 11
Society American Foresters, 257
Solanum. See Bitter nightshade. Horse

nettle
Solidago. See Goldenrod
Solomon's seal, 152, 169
Sonchus. See Sow tliistle
SoTbus. See Ash
Sorghum, 150
Sorrel. See Wood sorrel
Soutliem white cedar, 159
Sow bugs, 226

thistle, 153
Speedwell, 154
Spicebush, 155
Spiders, eaten by grouse, 107, 110, 167,

170
Spinal space, 15

tract, 12
Spiraea. See Meadowsweet
Spotted jewelweed, 141, 150
Spruce, 69, 86, 88, 160, 358, 359, 360
Squashberry, 136, 158
Squirrels, 119, 127, 128, 215, 358
STAFSETH and KOTLAN, 230, 241, 246
Starflower, 154
Stellaria. See Chickweed
STETSON, 311, 312
STEVENS, 1
STODDARD, 25
STODDART, 23, 186, 223, 312
Stonecrop, 153
Strawberr>', 66, 105, 107, 110, 112, 119,

150, 169, 172, 340
Strawberry bush, 155
Streptopus. See Twisted stalk
Striped maple, 159
STUDHOLME, 259, 298, 301, 303
Submalar space, 14
Subspecies, 5, 6, 7, 8
Subulura strongylina, 233
Sudan grass, 150
Sugar maple, 159, 348, 353, 357
Sumac, 65, 105, 107, 108, 112, 129, 157,

169, 181, 340, 349, 355, 357, 358,

359, 360
Sunflower, 150
Sunshine, related to nesting, 184

Sunspots, 179, 320

Superciliary space, 14

Survival factors, 3

SWAINSON and RICHARDSON, 20

Swamp rose, 349, 354, 355, 357

SWANSON, 178

Sweet birch. See Black birch

cherry, 126, 160

clover, 151, 170
Sweetfern, 142, 156
Switch grass, 151
Sycamore, 160

Symphoricarpos. See Coralberry, Snow-
berry
Symplocarpus. See Skunk cabbage
Syngamus trachealis, 2.33

Tamarack, 159

Tapeworms. See Cestodes

Taraxacum. See Dandelion

Taxus. See Canada yew

Teaberry, 155

Tearthumb, 152

Teleoptiles, 10

Temporal space, 14

Temperature, related to cover used, 94

Territory, 50

Tetrameres americana, 233

Tetrao fusca, 6

Tetraonidae, 1

Thalictrum. See Meadow rue

Thaspium. See Meadow parsnip

Thelypteris. See Ferns

Thimbleberry, 132, 1.57

Thistle, 149

Tlilaspi. See Field penny cress

Thomapple. See Hawthorn

THORNTON, 356

Thoroughwort, 150

Three-leaved gold thread, 149

Thuja. See Arborvitae

Tiarella. See False miterwort

Tick trefoil, 149, 169

Ticks, as ectoparasites, 226, 245

Tilia. See Basswood

TODD, 6, 7

TORREY, 56

Touch-me-not. See Jewelweed

Trailing arbutus, 140, 149

TRAINER, 11, 12, 13, 16

Trapping of grouse, 265; of fur-bearers,

255
Trematodes, 226, 236, 238
Trichomonas sp., 242
Trichoptera, 164
Trientalis. See Starflower
Trifolium. See Clovers

Index

385

Triosteum. See Feverwort
TRIPPENSEE, 258, 261
Trombicula microti, 248
Trypanosoma gaUinarum, 240, 241

sp., 240
Tsuga. See Hemlock
TUBES, 313
Tularemia, 242, 243
Tulip tree, 354
Timiors, 244
Tvvinflower, 156
Twisted stalk, 153
TYZZER, 229, 2S6, 238, 241, 242, 243

Ulcerative enteritis, 244

Ulmus. See Elm

Under carpal coverts: greater primary,
greater secondary, lesser primary,
lesser secondary, middle primary and
middle secondary, 14

Upper wing space, 15

Vaccinium. See Blueberry

VAN COEVERING, 260

VAN DERSAL, 60, 104, 114, 121, 148

Ventral tract, 12

Ventriculitis, 244

"Vermin" hunts, 265

Veronica. See Speedwell

Vetch, 154

Viburnum, 112, 136, 157. 168, ,340, 361

Vicia. See Vetch

Viola. See Violet

Violet, 66, 68, 110, 111, 145, 154

Virginia creeper, 143, 156, 169, 355, 357

deer. See White-tailed deer

pine, 69, 86, 352
Viruses, 226
Vitis. See Grapes

Waldsteinia. See Barren strawberry

Wand lespedeza, 151

Water cress, 152

Water, need for, 182

Waxmyrtle, 156

Wayfaring tree, 136, 157

Weasel, 197, 205, 211, 212, 213, 263,

264, 370
Weather, affecting grouse, 40, 46, 80, 95,

184, 185, 187, 190, 305, 307, 318,

321
Weather, related to: adult losses, 185;

brood mortahty, 185, 305; cycles,

318, 321; egg losses, 185; population

changes, 190, 318

WEBB, 332
Whippoorwill, 208
White avens, 150

baneberry, 148

birch. See Birches

cedar. See Arborvitae

clover, 154, 355

mustard, 148, 170

oak, 127, 160, 175, 348, 353, 357, 358

pine, 65, 69, 86, 88, 160, 337, 348, 352,
357, 358

spruce, 337, 352
White-tailed deer, 114, 116, 117, 118,

119, 120, 123, 124, 127, 128, 130,
131, 132, 135, 137, 138, 219, 330

Wild apple. See Apple

bean, 152

carrot, 149

ginger, 148

grape. See Grape

lettuce, 67, 151

peppergrass, 151

plum, 126, 160

raisin. 111

red raspberry, 132, 157

rose. See Rose

turkey, 116, 119, 128, 130, 131. 138
Willow, 144, 160

herb, 149
WILSON, 20, 255
WING, 320
Winterberry, 141, 156
Wintergreen, 69, 70, 105, 106, 107, 112,

120, 168, 340

Winter weather, related to food supplv,
189; related to grouse, 187, 307

Witch hazel, 141, 156, 170, 181

Witchhopple, 68

Withered, 68, 136, 157

"Wolf" trees, 343, 347

Wood duck, 128

Woodland management, 326. See also
Management of the ruffed grouse

WOODRUFF, 23, 190, 191, 223, 270

Wood sorrel, 68, 107, 142, 151, 169

Woods roads, 76, 342, 355

WRIGHT, 129, 130, 131, 134, 138

Yarrow, 148
YEATTER, 282
Yellow avens, 150
Yukon ruffed grouse, 6

Zea. See Com
Zizia, 154