A SERIES OF BOOKS IN AGRICULTURAL SCIENCE

Animal Science
EDITORS
G. W. Salisbury
E. W. Crampton (1957-1970)
WILDLIFE ECOLOGY
AN ANALYTICAL APPROACH

AARON N. MOEN
Cornell University

with a foreword by Douglas L. Gilbert

Colorado State University

W. H. FREEMAN AND COMPANY

SAN FRANCISCO

rn
 Copyright © 1973 by W. H. Freeman and Company

No part of this book may be reproduced by any mechanical,

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private use without written permission of the publisher.

Printed in the United States of America

12345 6 789

Library of Congress Cataloging in Publication Data

Moen, Aaron N 1936-

Wildlife eco logy.

Bibliography: p.
1. Zoology - Ecology. 2. Wildlife management.
3. Ruminantia. I. Title.
QH541.M54 599'.735'045 73-6833
ISB N 0-7167-0826-4
 CONTENTS

Foreword XIII

Preface XV
Acknowledgments XVII

PART 1
LIFE, INTERACTIONS, AND ECOLOGICAL
MODELING

1 PRODUCTIVITY GRADIENTS 3

1-1 The concept of life 3

1-2 The role of the analyti ca l ecologist 4
1-3 Dead or alive 5
1-4 Productivity of the individual 5
1-5 The natural mosaic 7
1-6 Interspecies interaction 10
1-7 Intraspecies interaction 11
1-8 Reproductive patterns 12
1-9 A theoretical mosaic 13

2 INTERACTIONS BETWEEN ORGANISMS AND

ENVIRONMENT 16

2-1 Functional re la tionships 16

2-2 The scope of the environment 21
OPERATIONAL, POrENTIAL, AND HISTORICAL RELATIONSHIPS 21
TIME 24
ASSOCIATED RELATIONSHIPS 25
HABITAT EVALUATION 26

3 ECOLOGICAL MODELING AND SIMULATION 32

3-1 Mathematical models 33
3-2 The analyti cal mode l in ecology 36
vi CONTEN TS

PART 2
THE DISTRIBUTION OF MATTER AND ENERGY
IN TIME AND SPACE

4 SOIL, WATER, AND TOPOGRAPHY 43

4-1 Soil 44
PHYSICAL CHARACTERISTICS 44
SOIL PROFILE 46
SOIL WATER 46
CHEMICAL CHARACTERISTICS 47
BIOLOGICAL CHARACTERISTICS 48
4-2 Soil classification in transition 51
4-3 Eutrophication 52
4-4 Biogeochemical cycles 53

5 WEATHER IN RELATION TO
PHYSICAL CHARACTERISTICS 57

5-1 The distribution of sunlight 57

THE AT MO SPHERE 60
5-2 Atmospheric water 62
5-3 Precipitation 62
RAIN 62
RAINFALL IN RELATION TO SOIL AND TOPOGRAPHY 66
SNO W 68
CONDUCTIVITY 69
5-4 Snow cover in relation to kinetic energy 69
WINDPACK 69
THE EFFECT OF WINDBREAKS 70
SNOWFALL INTERCEPTION 72
THE ROLE OF SNOW IN A PRECIPITATION-CANOPY-SUBSTRATE MODEL 72
THE EFFECT OF SNOW DISTRIBUTION ON ANIMALS 72

6 WEATHER AND THE PROCESSES OF

THERMAL EXCHANGE 75

6-1 The four modes of heat transfer 75

6-2 Radiant energy exchange 76
THE ELECTROMAGNETIC SPECTRUM 76
SOLAR RADIATION 77
INFRARED RADIATION 78
ATMOSPHERIC TRANSMISSION AND ABSORPTION CHARACTERISTICS 78
ATMOSPHERIC EMISSION 80
RADIATION FROM PHYSICAL AND BIOLOGICAL OBJECTS 84
EMISSIVITY AND REFLECTIVITY OF SNOW 85
RADIANT TEMPERATURE IN RELATION TO AIR TEMPERATURE 85
 CON T EN TS vii

6-3 Instrumenta ti on for measuring rad iation 90

6-4 Convecti on 92
A IR MOVEMEN T 92
MEAN VELO CITY W I ND PROFILES 93
6-5 Co nve ctive hea t loss 94
6-6 Cond uction 97
6-7 Heat loss by eva porati on 98
6-8 Concl usio n 103

PART 3
METABOLISM AND NUTRITION

7 ENERGY METABOLISM 109

7-1 Basal me tabo lism and associated terminology 110
USES FOR MEASUREMENTS OF BASAL METABOLIC RATE 110
CONDITIONS FOR MEASUREMENT 110
ASSOCIATED TERMINOLO GY 112
7-2 Measurements of basal metabolic rate 112
DIRECT METHODS
112
INDIRECT METHODS
112
CALCULATIO N OF HEAT PRODUCTION
113
7-3 Metabolic rates of ruminants 115
RELATIONSHIPS OF BODY WEIGHT
115
META BOLIC RATES OF WILD RUMINANTS 116
FASTING METABOLISM OF DOMESTIC RUMINANTS 120
7-4 Metabolic rates of other animals 120
7-5 Factors influencing energy me ta bolism and heat production 123
THE RELATIONSHIP BETWEEN HEAT PROOUCTI ON AND SURFACE AREA 126
HEAT INCREMENTS DUE TO DIET 127
HEAT INCREMENTS DUE TO ACTIVITY 127
NONSHIVERING THERMOGEN ESIS 127
SEX DIFFERENCES 128
REPRODUCTIVE CONDITION 128
RHYTHMIC CH ANGES I N THE BASA L METABO LI C RATE 129
I NSULATION CHARACTERISTICS
129
WEATHER
130
PATHOGENS AND PARASITES
130
7-6 Social and psychological effects on heat production 130

8 DIGESTION 135
8-1 The definition of digestion 135
8-2 A research philosoph y 135
8-3 Chemical composition of food ma terials 136
WATER 137
NITROGENOUS SUBSTANCES- CRUDE PROTE I N 137
CARBOHYDRATES-NITROGEN - FREE EXTRACT 138
viii CONTENTS

CARBOHYDRATES- CRUDE FIBER 138

ETHER EXTRACT 138
ASH 138
SUMMARY OF PROXIMATE ANALYSIS 139
8-4 The nutritive evaluation of forages 139
8-5 The alimentary canal 141
ANATOMY 141
HISTOLOGY 144
8-6 Mechanical and secretory processes in digestion 145
INGESTION 147
SALIVATION AND MASTICATION 147
R UMINATION 147
STOMACH AND INTESTINAL MOVEMENTS 147
DEFECATION 149
8-7 Chemical processes of digestion 149
RUMEN MICROORGANISMS 150
FERMENTATION 151
pH 153
8-8 Products of fermentation 153
HEAT ENERGY 153
GASES 153
VOLATILE FATTY ACIDS 154
PROTEIN 156
VITAMINS AND MINERALS 157
8-9 Passage of digesta through the gastrointestinal tract 157
8-10 Digestion and absorption in the gastrointestinal tract 159
8-11 Summary 159

9 INGESTION AND NUTRIENT UTILIZATION 164

9-1 Variations in nutrient intake 164
SEASONAL VARIATIONS 164
INDIVIDUAL VARIATIONS 165
9-2 Regulation of nutrient intake 166
RELATIONSHIP TO WEIGHT
166
RELATIONSHIP TO ENERGY EXPENDITURE
167
CONTROL OF INTAKE
167
9-3 Energy utilization 171
PATHWAYS OF ENERGY UTILIZATION
172
NET ENERGY FOR MAINTENANCE
175
NET ENERGY FOR PRODUCTION
176
SUMMARY OF ENERGY UTILIZATION 176
9-4 Protein utilization 176
9-5 Efficiency of nutrient utiliza tion 177
DIGESTION COEFFICIENTS 177
NET ENERGY AND PROTEIN COEFFICIENTS 177
METABOLIC EFFICIENCY 178
9-6 Body growth 179
9-7 Summary 182
r

CONTENTS ix

PART 4
BEHAVIORAL FACTORS IN RELATION TO
PRODUCTIVITY

10 THE ORGANISM AS A FUNDAMENTAL UNIT

IN A POPULATION 189

10-1 Energy, matter, and time 190

10-2 Biological chronology 191

11 INTRASPECIES INTERACTION 206

11-1 Sensory perception 206

11-2 Protective behavior and family ties 213
11-3 Movement patterns 214
11-4 Feeding behavior 216
11-5 Rest 216
11-6 Play 217
11-7 Social order 217
DOMINANCE PATTERNS 217
11-8 Radio telemetry and behavioral analyses 218

12 INTERSPECIES INTERACTION 225

12-1 Predator-prey relationships 226

12-2 Factors affecting predation rates 228
12-3 Energetic considerations 230
TROPHIC LEVELS 232
12-4 Man as a predator 235
12-5 Parasites and pathogens 236
12-6 Competition 238
12-7 Conclusion 238

PART 5
ENERGY FLUX AND THE ECOLOGICAL
ORGANIZATION OF MATTER

13 THERMAL ENERGY EXCHANGE BETWEEN

ORGANISM AND ENVIRONMENT 245

13-1 Thermal energy exchange 245

13-2 The concept of homeothermy 246
REGIONS OF THERMAL EXCHANGE 247
ANALYSES OF HOMEOTHERMIC RELATIONSHIPS 249
13-3 Measurement of thermal parameters 250
CONDUCTION COEFFICIENTS 250
TEMPERATURE-PROFILE MEASUREMENTS 253
CONTENTS

TEMPERATURE PROFILES IN FREE CONVECTION 254

TEMPERATURE PROFILES IN FORCED CONVECTION 254
TEMPERATURE' PROFILES UNDER RADIANT ENERGY LOADS 255
13-4 Radiant surface temperature related to air temperature 258
13-5 The concept of thermal depth 261
13-6 Geometry and surface area 262
13-7 The calculation of heat loss 265

14 PHYSIOLOGICAL, BEHAVIORAL, AND GENETIC

RESPONSES TO THE THERMAL ENVIRONMENT 273

14-1 The role of the animal in thermal energy exchange 273

14-2 The thermal regime and the critical thermal environment 274
14-3 Heat loss in relation to heat production 275
14-4 Physiological responses to the thermal environment 276
THERMOGENIC RESPONSES 277
THERMOREGULATORY RESPONSES 281
14-5 Behavioral responses 286
I ND IVIDUAL RESPONSES 286
GROUP RESPONSES
292
14-6 Genetic responses 293
14-7 Summary 296

15 THE ORGANIZATION OF ENERGY AND MATTER

IN PLANT AND ANIMAL COMMUNITIES 299
15-1 Taxonomic relationships 300
15-2 Morphological relationships 300
15-3 Chemical relationships 306
15-4 Temporal relationships 308
SUCCESSION
314
15-5 A plant-production model 316
A GEOMETRIC MODEL
31 .6
A SEASONAL MODEL
318
A PHYSIOLOGICAL MODEL FOR PREDICTING NET PHOTOSYNTHESIS 321
15-6 Perturbations 325
OVERGRAZING 325
FIRE 325

PART 6
PRODUCTIVITY, POPULATIONS, AND
DECISION-MAKING
16 A BIOLOGICAL BASIS FOR THE CALCULATION
OF CARRYING CAPACITY 333

16-1 The conceptual design 333

16-2 Protein requirements of the individual animal 334
PROTEIN REQUIREMENTS FOR MAINTENANCE 334
PROTEIN REQUIREMENTS FOR PRODUCTION 336
 CONTENTS xi

PROTEIN REQUIREMENTS FOR GESTATION 336

MILK PRODUCTION 341
SUMMARY OF THE PROTEIN REQUIREMENTS OF THE INDIVIDUAL 344
16-3 Energy re qui rements of the individual animal 347
ENERGY REQUIREMENTS FOR MAINTENANCE AND ACTIVITY 347
ENERGY REQUIREMENTS FOR PRODUCTION 352
ENERGY REQUIREMENTS FOR GESTATION 352
MILK PRODUCTION 354
SUMMARY OF THE EN ERGY REQUIREMENTS OF THE INDIVIDUAL 356

17 MATHEMATICAL ANALYSES OF FACTORS

AFFECTING CARRYING CAPACITY 365

17-1 The carrying-capacity model 365

17-2 The program format 36 7
INPUTS FOR ANIMAL REQUIREMENTS 367
INPUTS DESCRIBING THE RANGE SUPPLY 368
DECISIONS 369
17-3 Constraints in the animal-range relationship 370
17-4 Program outputs 371
17-5 The dynamic characteristics of the animal-range
relationships 372
17-6 Weight changes 372
ABSOLUTE VALUES 372
PERCENTAGE OF BODY WEIGHT 373
LACTATION EFFECTS 374
17-7 Forage ingested 375
RELATIONSHIP TO BODY WEIGHT 375
MA INTENANCE-GAIN COMPARISONS 375
FORAGE CONSUMPTION DURING GESTATION 377
FORAGE I NGESTED AS A FRACTION OF THE PHYSICAL RUMEN CAPACITY 378
17-8 Physiological efficiency 380
ME TABOLIC FECAL NITROGEN AND NET PROTEIN RELATIONSHIPS 380
META BOLIC FECAL NITROGEN, FORAGE INGESTED, AND NET
PROTEIN COEFFICIENT RELATIONSHIPS 380
METABO LIC FECAL NITROGEN, BODY WEIGHT, AND NET PROTEIN
COE FFICIENT RELATIONSHIPS 38]
BODY WEIGHT, NET PROTEIN COEFFICIENT, AND PHYSICAL RUMEN
CAPAC ITY RELATIONSHIPS 382
17-9 The expression of carrying capacity in deer-days 383
EFFECTS OF BODY SIZE 384
EFFECTS OF SNOW DEPTH 386
FORAGE PRODUCTION AT DIFFERENT STAGES IN SUCCESSION 388
I7-1O Summary 388

18 N-DIMENSIONAL POPULATION STRUCTURES 390

18-] Sex and age ratios 39]
18-2 Parasitism 397
18-3 Geometry and posture 398
xii CONTENTS

18-4 Population requirements for protein 400

18-5 Population requirements for energy 400
18-6 Other population dimensions 400

19 PREDICTING POPULATION DYNAMICS 404

19-1 Ecological productivity gradient 404

19-2 The relative importance of different variables 405
ANIMAL REQUIREMENTS 405
RANGE SUPPLY 406
FACTORS AFFECTING BOTH ANIMAL REQUIREMENTS AND RANGE SUPPLY 406
19-3 Factors to consider in population analyses 408
19-4 Population analyses for n =1, 2, ... n 408
MORPHOLOGICAL AND PHYSIOLOGICAL CONSIDERATIONS 409
BEHAVIORAL CONSIDERATIONS 410
A THEORETICAL AVERAGE DEER 410
19-5 Time in relation to biological events 411
SEASONAL CHANGES IN ANIMALS AND PLANTS 411
THE IMPORTANCE OF SEASONAL CHANGES 412
VARIATIONS IN THE TIME OF BREEDING 413

20 ECOLOGICAL ANALYSES AND DECISION-MAKING

PROCEDURES 418

20-1 The simulation of management practices 418

BIOLOGICAL CONSIDERATIONS 419
SOCIAL AND ECONOMIC CONSIDERATIONS 419
POLITICAL CONSIDERATIONS 420
20-2 Conclusion 420

APPENDIXES

A-I Weights and Measurements 425

A-2 Weather, Thermal Factors, and the Julian Calendar 428
A-3 Weight and Metabolic Weight 432
A-4 Radiant Temperature in Relation to Air Temperature 433
A-S Surface Area in Relation to Weight 436
A-6 Symbols 438
A-7 Reference Books 440
A-8 Instructions for Contributors to the Professional Literature 442

INDEX 445
 FOREWORD

In recent years, environmental problems have created great general concern. Thus,
the time has come when a revitalized and more effective approach to the manage-
ment of natural resources is necessary. This is especially true in light of increased
human populations.
In the past, individual abuses of the natural resources have been treated as
isolated problems-an approach doomed to failure . Instead, individual abuses can
be seen as parts of a larger problem: the increasing pressure of an expanding
population on dwindling nature resources. That problem often appears over-
whelming. Ia seeing it, many have given up in despair. But a great problem may
be broken down; each part can be attacked separately and perhaps solved. Bit
by bit the big problem becomes solvable. The importance of each issue, whether
it be protein availablility, harvest of females, or disposal of waste pollutants,
depends on the particular role of the issue in the overall environmental structure.
Wild animals, and the management of them are a vital part of the environ-
mental "machine," a part that also is made of smaller parts. Age, sex, and time
of year affect the physiology of an individual animal. These, together with nutri-
tional factors, genetic history, and features of the physical environment, combine
in the complex system that determines the interactions between an animal, other
organisms, and the land.
It is the essence of the wildlife manager's job that he understand the system
and be able to work with it. H~ must understand how an organism fits into the
ecosystem. He must understand the effects of the organism on its total environ-
ment and the effects of the environment on the organism.
In Wildlife Ecology: an analytical approach, Professor Moen has analyzed this
natural system. He evaluates each component and welds them together into a
unified whole . Although most of the examples deal with white-tailed deer, the
concepts are applicable to the other wild ruminants and, indeed, to all organisms.
xiii
xiv FOREWORD

Professor Moen's creative research and dedication have produced a work in

which traditional pieces of wildlife management- numbers and conditions of
animals, nutritive values of range plants, behavior patterns-are at last presented
as parts of a greater whole . This book should be made available to every wildlife
professional, wheth..er technician, manager, biologist, conservation officer, admin-
istrator or researcher. It is an important publication and the time for it has come.

Douglas L. Gilbert

Colorado State University

Fort Coll ins, Colorad o
September 1972
 PREFACE

Rapid advances in analytical capabilities within the last fifteen years have made
it possible for the ecologist to do things within a time dimension that were unheard
of a few yea rs ago . The capabilities for rapid analyses pose a threat to the
discipline of ecology, however, because there can be a tendency to use numbers,
large quantities of them, hoping by some magical means of computer analysis
to find some relationships emerge.
The reorganization of numbe~s within a computer program of storage and
compu tation is nothing more than a rapid bookkeeping system. Computers used
in such a way do n ot usually help much in gaining insight into the mechanisms
that are operating in the natural world. They tend to promote a false sense of
security.
The real benefits of computer analyses emerge if they are used to extend the
analyst' s capabilities for analyzing the relationship between one factor or force
and another factor or force in the ecosystem. It is important to realize that the
human min d must always be ahead of the computer, with the electronic system
doing rapid computations that are too numerous and time-consuming to do in
any other manner. This suggests that the first models built by analytical ecologists
are of necessity very simple ones . Let them be no more complex than the model
builder can full y comprehend, insuring that he knows not only the capabilities
of his analytical model but also its weakness. A progression of such simple models
will result in more complex, working models that represent a known portion of
the ecosystem.
In this book I have aimed at promoting the building of simple but workable
models . They do not require large computer centers for their use; small desk-top
computing systems are entirely adequate. In fact, many of the models suggested
can be done manually, with the principles of model building illustrated just as
xv
xvi PREFACE

well. Thus the book should be of interest to ecology classes in many types of
educational institutions, from the small college to the major university. I am
convinced that, wherever the student is located, the major factor that will deter-
mine his progress in ecology is his ability to think, along with the guidance of
a professor who stimulates thinking about meaningful ecological relationships.

Aaron N . Moen
April 1973
 of
1m
ACKNOWLEDGMENTS
er-
of
ps.

oen

The completion of a book is not possible without the help of many people. My
own -efforts have been made possible through the kind direction and guidance
given to me by my parents on their farm in western Minnesota. The opportunities
for contact with wild animals and native plants in that area stirred within me
an interest to pursue an understanding of the relationships between organism
and environment.
My academic career in the field of natural resources began under the guidance
of Dr. Max Partch at St. Cloud State College. His enthusiasm for teaching in the
field impressed me greatly. Dr. William H. Marshall, of the University of Minne-
sota, gave me opportunities, freedom, and responsibility as I pursued a PhD.
The most significant academic work that permitted me to delve into the energy
relationships of deer at that time was that of Helenette Silver and her colleagues
at the New Hampshire Fish and Game Department and the University of New
Hampshire. Without her pioneering efforts in the field of energy metabolism of
white-tailed deer, my PhD. dissertation could not have started me on the chal-
lenging research on the energetics of a free-ranging animal.
I wish to thank the many friends I have made in the field of wildlife manage-
ment, especially the deer biologists in the State of New York who always provide
stimulating interaction as we proceed together to understand this most important
resource in New York State. My colleagues at Cornell, especially Dr. Peter Van
Soest of the Department of Animal Science, have provided many insights into
the animal-environment relationships currently under investigation. Dr. Douglas
L. Gilbert, formerly at Cornell and now at Colorado State University, has discussed
big-game management with me on many occasions. Dr. Donald Ordway and his
staff of aerodynamic engineers have been of great help in our thermal analyses
at the BioThermal Laboratory.

xvii
xviii ACKNOWLEDGMENTS

Dr. Dwight A. Webster, former head of the Department of Naturai Resources,

and the administrators of the Agricultural Experiment Station at Cornell have
all been most helpful as I established a research program at the BioThermal
Laboratory. Funds for research at the Laboratory have been contributed through
the Pittman-Robertson Federal Aid program, Project W-124-R, and the New York
State Department of Environmental Conservation. Additional funds from the
Agricultural Experiment Station at Cornell, the Cornell Research Grants Commit-
tee, the National Science Foundation, The Loyalhanna Foundation, and the Na-
tional Rifle Association have helped support the work at the BioThermal Labora-
., tory.
The staff at the Laboratory has contributed significantly to the work that is
described in this text. My respect for the abilities and dedication of my students
cannot be fully expressed by acknowledgment but will be manifested by their
contributions in the future. I must recognize the help and accomplishments of
former students, especiq.lly Dr. Keith. E. Evans and Dr. Deborah S. Stevens. The
work of Nadine L. Jacobsen and Charles T. Robbins, both PhD. candidates
studying the energy relationships of deer, has provided much insight into the
complex animal-environment relationships that are the focus of study at the
Laboratory. William Armstrong, laboratory technician, has helped in the design
and construction of research equipment and in the care of our experimental deer
herd. Richard E. Reynolds, foreman at the Ithaca Game Farm, has contributed
much to the program with his help in the construction of the deer pens, mainte-
nance of the facilities, and continual attention to our needs . Eleanor Horwitz
offered many fine suggestions on ways to improve the manuscript. I appreciate
her efforts to convince me to say things in the simplest way possible.
Students in my courses have raised many stimulating questions. I wish that
each one of them could participate actively rather than passively in the educational
process of research and discovery.
Finally, the help and encouragement of my wife, Sharon, and of Ronald,
Thomas, Daniel, and Lindy cannot be fully expressed in words . It has often been
impossible to keep up with some of the domestic duties confronting every hus-
band and father because of the urgency of research according to a biological clock
and my own intense interest in the subject. As Tom (age 9) said when I suggested
I might write another book, "Oh no, not another five years of that!"
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