(Robert Smith) Elements of Ecology

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Elements of
ECOLOGY
Elements of
ECOLOGY
GEORGE L. CLARKE
Harvard University
and
Woods Hole Oceanographic Institution
JOHN WILEY & SONS, INC.

New Yorfc
CHAPMAN & HALL, LTD.
London
COPYRIGHT, 1954
BY
JOHN WILEY & SONS, INC.
All Rights Reserved
This book or any part thereof must not

be reproduced in any form without
the written permission of the publisher.
THIRD PRINTING. APRIL. 1959
Contribution No. 704 from the Woods Hole Oceanographic Institution
Library of Congress Catalog Card Number: 54-12205
PRINTED IN THE UNITED STATES OF AMERICA

To
M. S. C.
Preface
"Live Alone and Like It" is a slogan that no living thing can adopt.
Every plant and animal is
subject to both the living and the non-living
influences of its surroundings. Every organism depends upon its
environment to supply it with vital materials and energy. Every
living being must share its world with members of its own species and
with members of other species be they friend or foe.
Man is no
exception. Man is surrounded by many kinds of living
things, and he must derive his needs from the world around him. Man
must learn to live in adjustment with his fellow men, and with the
plants and animals of his environment, and to use his natural resources
judiciously, or he will be exterminated.
Obviously, then, the interrelations of the organism and its environ-
ment are crucial; and ecology, which is the study of these interrela-
tions, is of great significance. Yet few books are available which deal
with the general principles of the whole subject. Most of the books
in the field of ecology treat
primarily either "plant ecology" or "animal
ecology." But animals cannot get along without plants, and plants
are almost always vitally influenced by animals. Man is dependent
upon both.
Accounts of the flora and fauna of various regions and descriptions
of a variety of habitats have long been available. Early ecologists
have investigated the effects of environmental factors on the activities
of living things. But not until relatively recently have we realized
that plants and animals also react on their surroundings in such a way
that they form a reciprocating and integrated system with their
environment. The modern ecologist focuses his attention on the
of the forces in the living community.
dynamic interplay
The purpose of this book is to bring together in one place and in a
simple way the elements of ecology with special emphasis on the
modern viewpoint of the science. It is desired to stress the unity of
ecology and the necessity for including the influence of both plants
and animals as well as the physical forces as part of the environment.
The subject is old, but efforts to crystallize the basic principles
in any
exact way are relatively new. The potential scope of ecology is very
vii
viii
Preface
great, and no attempt at any exhaustive treatment of the

subject is
made here. Rather, this book presents an introductory account of the
fundamental relationships of ecology upon which an understanding
of the life of the individual and of the community
depends. Since
many of these relationships are seen in a relatively simple form in the
marine environment, many illustrations are taken from marine habitats.
The book is written primarily for students in ecology. It is also
intended as background material for those interested in conservation,
forestry, agriculture, fisheries, wildlife management, and other
branches of "applied ecology," -The general biology student should
also derive value from the book since a knowledge of the principles
governing the real lives of animals and plants in nature is a vital
backdrop for
problems physiology, morphology, genetics, and
in
evolution. Furthermore, it must be remembered that every one of us
is an
organism with an environment and that each of us forms part of
the environment for other living beings. The general reader, as well
as the biology student, is interested to know how the world of life
works. Populations of men follow many of the same laws as popula-
tions of microbes. The need for a knowledge of the proper adjust-
ment among men and between man and his natural environment was
never greater than it is
today.
The drawn in large part from a course in
material of the book is
ecology given by the author at Harvard University. Many ideas and

illustrations have been derived from discussions with colleagues at
Harvard University, the Woods Hole Oceanographic Institution, and
elsewhere, and acknowledgment is made of this material. I am also
indebted to W. C. Allee, A. Cain, J. T. Curtis, R. W. Dexter, W. H.
S.
Drury, E. H. Graham, D. R. Griffin, H. C. Hanson, A. G. Huntsman,

S. C. Kendeigh, B. H. Ketchum, P. C. Lemon, D. Merriman, R. S.
Miller, E. W. Moore, E. T. Moul, O. Park, T. Park, H. M. Raup,
A. C. Redfield, F. A. Richards, G. A. Riley, J. H. Ryther, P. S. Sears,
J.
G. Steel, H. C. Stetson, and J. M. Teal for reading parts of the
manuscript and offering helpful criticisms. I am particularly grateful
to Marion S. Clarke, Donald Kennedy, and J. E. G. Raymont for
assistance in preparing the manuscript and for many valuable
suggestions.
GEORGE L. CLARKE
August, 1954
Contents
Chapter 1 VIEWPOINT OF MODERN ECOLOGY 1
The Meaning of Ecology 2

The Meaning of Environment . 3
The Critical Environment 5
The Development of Ecology 13
The Ecology of Plants and Animals 13'
The Ecology of Habitats and of Individuals 14
The Ecology of Populations and of Communities 14
The Ecological Complex 16
The Scope of Ecology 18
Approach to the Study of Ecology 20
Chapter 2 THE MEDIUM 23
Contrasting Qualities of Air and Water 24

Pressure 27
Pressure Reduction with Altitude 28
Pressure Increase with Depth 29
Support and Resistance to Motion 32
Effects on Structure and Size ,32
Effects on Locomotion through Medium 34
Passage of Medium through Organism 35
Existence of Plankton 36
Transportation by Medium 42
Sessile Existence 44
Distribution by Medium 45
Transport by Air 45
Transport by Water 48
Harmful Transport 52
Abrasive Action of Medium 53
Chapter 3
-
THE SUBSTRATUM 59
Significance of the Substratum 59

Needs Provided by the Substratum 59
Attainment of the Substratum 60
Reactions to the Substratum 61
The Variety of Substrata 62
Rock, Sand, and Mud in Aquatic Environment 66
Influence of the Aquatic Substrata 66
Breakdown of the Substratum 69
ix
x Contents
Build-Up of the Substratum 69

Rock, Sand, and Soil in Terrestrial Environment 71
Influence of the Land Substrata 71
Land Surfaces and Animals 72
Soil in Its Action on Plants 73
Action of Organisms on Soil 76
Abundance of Organisms in Soil 76
Formation
Soil 77
Humus and the Colloidal Complex 79
The Soil Profile 80
Soil-group Divisions 84
Chapter 4
-
WATER 90
Water Problem in the Aquatic Environment 91
Composition of Natural Waters 91
Methods of Meeting Osmotic Problem . .
........ 95
Limiting Effects of Salinity 97
Amphibious Situations 99
Swamps and Temporary Pools 99
Tidal Zone 101
Water Problem in the Terrestrial Environment 107
Occurrence of Water in Land Environment 109
Moisture in the Soil Ill
Moisture in the Air 112
Microclimates 117
Meeting Water Problem on Land 119
Influence of Moisture on Growth and Distribution . . . . . . 121
Chapter 5 TEMPERATURE 129

Distribution of Temperature 129
Extremes of Temperature and of Tolerance 129
Changes in Temperature 131
Changes in Time 131
Horizontal Changes 134
Vertical Changes 134
Seasonal Changes in Vertical Temperature Structure .... 138
Biological Action of Temperature
. 141
Extreme Temperature 142
Minimum Temperatures 142
Maximum Temperatures 143
Methods of Meeting Temperature Extremes 144
Morphological and Physiological Adaptations 144
Thermal Migrations 148
Action within Effective Range 152
Effect of Temperature on Biological Rates 152
155
Optimum Temperature
Other Effects of Temperature 156
Action of Temperature on Distribution 159
Mode of Temperature Limitation 159
Control by Extremes 160
Contents xi
Control by Need for Minimum Amount of Heat 164

Control by Need for Chilling 168
Results of Temperature Limitation 172
.
Special Cases of Common Boundaries 172
Frost Line 173
Tree Line 173
Life Zones 175
Temperature and Moisture Acting Together 181
Chapter 6 LIGHT 185

Distribution of Light 186
Light on Land 186
Spectral Composition 186
Intensity of Light 187
Duration and Amount of Light 188
Light in Water 191
Extinction and Modification of Light 191
Changes in Transparency 197
Biological Effects of Light 198
General Effects 198
Protective Coloration 199
Activity and Vision 202
Photokinesis 202
Vision 202
Bioluminescence 204
Orientation 207
Periodicity 216
Diurnal Periodicity 216
Lunar Periodicity 219
Seasonal Periodicity 224
Ultraviolet Light 230
Ecological Aspects of Photosynthesis 233
Land Plants 233
Aquatic Plants 235
Chapter 7 OXYGEN AND CARBON DIOXIDE 242
Oxygen 242
Availability of Oxygen
243
Terrestrial Environment 243
Aquatic Environment
244
Oxidation-Reduction Potential 251
Effects of Oxygen Availability 253
Environment
Terrestrial 253
Environment 255
Aquatic
Carbon Dioxide 259
Carbon Dioxide in Terrestrial Environment 260
Carbon Dioxide in Aquatic Environment 262
Reactions of Carbon Dioxide in Water 262
Ecological Effects of Carbon
Dioxide 265
xii Contents
Hydrogen Ion Concentration 268

Calcium Carbonate 270
Chapter 8 NUTRIENTS 277

Nutrients and the Environment 277
Modes of Nutrition 278
Influence of Nutrients on Green Plants 280
Nutrients Required 280
Law of the Minimum 281
Limitation by Nutrients in Nature 283
Influence of Nutrients on Animals 292
Decomposition and Regeneration 297
Processes of Decomposition and Transformation 298
Place of Decomposition 300
Stagnation in Cycles 303
Regeneration 304
Rate of Regeneration 305
Ratio of Regenerated Materials 306
Chapter 9
-
RELATIONS WITHIN THE SPECIES 309
Origin of Groups 309

Reproduction 309
Passive Transport 311
Active Locomotion 312
Common Orientation 312
Mutual Attraction . 314
Effects of Increased Numbers 316
Harmful Effects 317
Beneficial Effects 322
Protection 322
Influence on Reproduction 323
Division of Labor 327
Population Development 333
Natality and Mortality 333
, Biotic Potential and Environmental Resistance 334
Form of Population Growth ^ 336
Logistic Curve . < 337
Equilibrium and Fluctuation 340
Optimal Yield 344
Spatial' Relations of Populations
348
Space Requirements 348
Home range and Territory 350
Homing 353
Return Migration 355
Emigration 357
Chapter 10 RELATIONS BETWEEN SPECIES 362

364
Symbiosis
Mutualism 364
Mutualism with Continuous Contact 364
Mutualism without Continuous Contact 370
Contents xiii
Commensalism 374
Commensalism with Continuous Contact 374
Commensalism without Continuous Contact 377
Antagonism 381
Antibiosis 383
Exploitation 385
Parasitism 386
Predation . . 392
Competition . . 396
Chapter 11 THE COMMUNITY 401

Community Concept . 401
Community Dominance . 408
Ecotone . 410
Community Composition 114
Stratification of the Community 120
Chapter 12 SUCCESSION AND FLUCTUATION 125
Ecological Succession 125

Dispersal and Invasion 127
Barriers 428
Ecesis 429
Succession and Climax 430
Types of Succession 432
Primary Succession 432
Secondary Succession 435
Convergence 438
Succession in Special Habitats 438
Modification of Succession 444
Community Classification 447
Community Type 447
The Biome 448
Fluctuation within the Community 451
Irruptive Fluctuation 453
Year-Class Analysis 454
Cyclic Fluctuation
457
Causes of Fluctuation 458
Origin of Cycles
460
Chapter 13 DYNAMICS OF THE ECOSYSTEM 465

Fundamental Operation 465
Principal Steps and Components
466
Niches 468
Trophic Levels and Relations 469
Ecological Cycle in the Ocean
473
Producers 474
Consumers 476
Herbivores 476
Carnivores 479
Decomposers and transformers 481
xiv Contents
Productivity of the Ecosystem 481

Concepts of Productivity 482
Standing crop 483
Relation to Population Growth 483
Determination of Equilibrium Level 485
Regional differences 487
Material Removed 489
Production rate 493
Turnover 496
Efficiencies 497
Productivity of Land and Water 500
Conclusion 503
References 505
Index 523
Viewpoint of
Modern Ecology
Every is surrounded
living thing by materials and forces which con-
stitute environment and from which it must derive its needs.
its
Contact with the environment is inescapable. Protoplasm, the essen-

tial constituent of an
organism, is a dynamic substance, requiring a
continuous exchange of energy and materials. In order to remain
alive the organism, too, must secure energy and materials, and these
can be procured only by exchange with the outside world. Thus the
animal or plant cannot live completely sealed in an impervious skin or
shell but requires from its surroundings: (1) a supply of energy; (2) a
supply of materials; (3) a removal of waste products. The exchanges

that the organism has with its environment may be thought of as an
"external physiology," and they are just as essential as its internal
physiological adjustments.
The supplying of the vital needs of the organism is by no means
the only action of the environment. Since the animal or plant must
leave its borders open to foreign trade, as it were, the possibility
exists that harmful materials will enter the organism or destructive
Influences act upon it. For example, algae in a river must be suffi-
ciently permeable to take in the water needed for their metabolism,
but, they drift into the sea, the higher salinity will cause a fatal loss
if
of water from their tissues. Animals in the environment of these

algae tend to add materials to the water which serve as plant nutri-
ents, but these same animals may also feed upon the algae and
eventually destroy the population.

In order for the an organism to continue, the environment
life of
must be satisfactory on two counts: (1) it must provide the minimum
requirements for life; (2) it must contain no influence incompatible
with life. If there is too little water, as in the Sahara Desert, or too
oxygen, as at the top of Mt. Everest, or if nutrients are not avail-

little
able, as on rocky plateaux, animals or plants cannot obtain their

1
2 Viewpoint of Modern Ecology
minimum needs. In other situations an ample supply of food and
light may be found, but the temperature may be so high as to exclude

all life. This is true in some hot springs. Certain soils of southwestern
United States are completely devoid of vegetation because of the
Death Valley on the Dieng
excessive amounts of salts or "alkali."
Plateau in Java populated with plants but no animal life can exist
is
because of high concentrations of carbon dioxide issuing from subter-

ranean crevices. Such influences are prohibitory to life and others
are harmful without being lethal. In a complex natural situation it
is not always easy to discover the particular role each factor plays,
but we can be certain that in every situation dealings with the environ-
ment are inescapable.
THE MEANING OF ECOLOGY

The study of these interrelations of plants and animals with their
environment constitutes the science of ecology. The environment in-
cludes the influences of other plants and animals present as well as

those of the physical features. In order to investigate the exchanges
and interdependencies involved, necessary to have a knowledge
it is
both of the organisms themselves and of the environments inhabited

by them. The ecologist must know the material with which he works.
He must have a grasp of the classification and the structure of plants
and animals, and he must understand what makes them tick. At
the same time the ecologist must be thoroughly aware of the nature
of the environment both living and non-living. He must be familiar
with the different types of terrain on land and with the different
qualities of water in the ocean, lakes, and rivers. He must appreciate
the special environmental conditions provided by the various kinds
of vegetation. He must have a knowledge of the circulation of water
and air and of the dynamic processes going on in the soil. But the
taxonomy, morphology, and physiology of organisms and the physi-
ography of land and sea, although providing a necessary background,
do not form the central core of ecology. The ecologist focuses his
attention primarily on the interrelationships between the organism
and its environment:
Ecology
/4-\
Organism 4-V Environment
The word "ecology" comes from the Greek "oikos," meaning "home"
or "estate" hence, ecology is the study of the home, or how the house-
The Meaning of Environment 3
hold of nature is
kept in order. Interestingly enough, although
ecology comes from the same root as our word "economics," the sub-
ject that we now call ecology was not given a name until a century
later, Man, being egocentric, began this type of study in his im-
mediate surroundings. Not until long afterwards did he realize that
man's economics is but a special case of the broader subject. In the
words of Wells, Huxley, and Wells (1939), "Ecology is really an ex-
tension of economics to the whole world of Economics and
life."
sociology might be thought of as the "ecology of man" in a broad sense.

The realization that the relations of man to his environment, both
physical and social, form a distinct and most important study is re-
flected in the increasing use of the term human ecology in sociology
and in other fields.
Our goal in ecology is to understand the interrelations of organisms
and their environments under natural conditions. Many biologists
of have tended to lose this point of view. As Elton (1939)
all sorts
stated, "The discoveries of Darwin, himself a magnificent field

naturalist, had the remarkable effect of sending the whole zoological
world flocking indoors, where they remained hard at work for fifty
years or more, and whence they are now beginning to put forth cau-
tious heads again into the open air!" Laboratory tests and field ex-
periments are, of course, used in studying the reactions of animals and
plants as an aid to understanding their actual and possible behavior
under natural conditions. Modern ecology thus goes beyond the mere
description of the habitat, or the listing of its inhabitants, to an anal-
ysis of causal relationships and a coordinated understanding of con-
structive and destructive processes in the community.
THE MEANING OF ENVIRONMENT

In referring to the natural environment one tends to think first of
the broad aspects of the landscape, such as water, soil, desert, or
mountain. These types of environment can be more exactly described
in terms of physical influences differences in moisture, temperature,
texture of material, and the like and of biological influences. Other
organisms form part of the environment just as much as the soil or the
rocks. No animal can live entirely as a hermit; every animal must
have other organisms within its range to serve as food. All animals are
dependent directly or indirectly upon green plants. Many plants are
dependent upon animals those that require pollination by insects, for
example. Some green plants could live independently for a tinie,
deriving their energy from the sun and their nutrients from the soil,
but, as soon as seedlings start to grow, relations of competition appear.
Every organism thus has other organisms as a necessary, or an un-
avoidable, part of its environment. Animals and plants compete
with each other, devour, or aid one another. Fellow inhabitants
cannot be disregarded as part of the environment, as is clearly ap-
parent in a thick stand of trees (Fig. 1.1) or in the slum conditions of
U, S. Forest Service
FIG. 1.1. Grove
of giant redwoods near Crescent City, California,
showing the
intense competition of the trees with each other and their profound influence on
the conditions beneath the forest canopy.
a sea-bird rookery (Fig. 1.2). In other situations the effects of the
presence of other animals and plants are more subtle. Although such
influences cannot be photographed, they are often just as crucial, and
the environment must be understood to include both physical and
biological influences whether conspicuousorobscîTe^
*5-*."
.
:
-.-
Jf '
''
.
, >"
'"'"'
FIG. 1.2. Gannet rookery on Bonaventure Island, Quebec, showing the keen
competition for nest sites.
The Critical Environment

Are the necessary dealings of the organism with the environment
usually successful? Can all living things
find their needs in the out-
side v/orld and avoid dangers? In most instances a large fraction of
the young plants and animals produced are unable to cope with the
environment, and relatively few survive. A striking example of the
magnitude of natural mortality is found in the accompanying data
from a study of the growth of mackerel eggs and young off the east
coast of the United States (Sette, 1943).
Young Mackerel
Stage of Duration Mortality
Spawning to 10-mm length 40 days 14% per day
Transition to post-larva few days 30% per day
Post-larva to 50-mm length 40 days 10% per day
These values show that mortality was high throughout development
and that a particularly critical period occurred during the transition
to the post-larval stage. Out of each million mackerel eggs spawned
in the area of this
investigation only four survived on the average to
reach a size at which the young fish could forage for themselves
effectively. Others would, of course, succumb before they grew to

spawning size.
The U. S. Fish and Wildlife Service estimated that early in 1944

there were roughly 125 million ducks in North America. These ducks
If all the adults and all the
ordinarily produce 10 to 16 eggs per pair.
eggs produced in 1944 had survived, there would have been about
900 million ducks inhabiting the continent in 1945. Actually the
population was not significantly larger at the next census, early in
1945, with again about 125 million ducks present. This means that
something like 775 million ducks had died during the year. The
sportsman's kill, which is estimated from licenses and bag reports, was
about 20 million during 1944, or less than 3 per cent of the total
mortality ( Griscom, 1947 ) Again
. it is obvious that a very large por-
tion of this bird population died during the year, and most died of
"natural causes" not by the hand of man.
Oneprocess developed during the course of evolution that acts to
insure the survival of species in the face of this high mortality is the
production of huge numbers of young. The prodigious fertility of

nature is a measure of the destructive action that is received from the
environment. In species in which each pair produces thousands or
millions of eggs or seeds, only two survive on the average to reach
the adult condition as long as the population is not continually in-
creasing. Many examples could be given of the huge excess number

of young that are produced and the small number that survive
(Fig. 1.3). Insects are notoriously prolific. The queen termite in an

African species after only one mating is reported to lay eggs at the
rate of one every few seconds and to continue egg laying at this
terrificpace for the rest of her life. A calculation shows that at this
rate each termite queen would lay approximately 30,000 eggs per day
or 100 million in her lifetime (Wheeler, 1923). According to esti-
mates, an oyster may discharge 500 million ripe eggs in one spawning.
If all these eggs developed into mature oysters and all subsequent
progeny survived, after only four generations we would have a pile of

oysters about eight times the size of the earthl
Plants also have the capacity to reproduce at very high rates, and,
if all
spores or seeds grew to maturity, plant populations would in-
crease with tremendous rapidity. Pine trees liberate such quantities
of pollen that a yellowish dustis often

deposited on all surfaces in the
vicinity.
A
tropical American orchid of the genus Maxillaria is re-
ported to produce as many as 1,756,000 seeds per capsule. Spores are

produced in astronomical numbers by many lower plants. A conserv-
ative estimate made for the downy mildew, Sclerospora, which attacks
maize in the Philippines, indicates that as many as 6 billion conidia
(spores) are liberated from the fungi parasitizing a single maize
plant during one night. Since discharge of conidia continues night
after night for months, the intensity of the
reproductive capacity of
.
these species surpasses the imagination (Weston, 1923).
an/ Eschmeyer, 1938

FIG. 1.3. Schematic representation of natural mortality in small-mouthed black
bass, Only a few fish survive to maturity from hundreds of eggs laid.
The adaptations of some plants and animals are more specifically per-
fected to their environments, and their reproductive rates can con-
sequently be relatively low, but even slow breeders tend to increase.

If the
elephant produces only six young in its life time, as has been
estimated, these creatures would nevertheless eventually increase to
such numbers, if all progeny survived, that they would cover the
whole surface of Africa. An even slower breeding rate is exhibited
by the California "big trees" (Sequoia gigantea] which do not pro-

duce their first seeds until they are 175 or 200 years old! And yet
even has the capacity to enlarge its population.
this species
Since no species of plant or animal goes on increasing indefinitely,

we are forced to the conclusion that, whether reproduction rate is fast
or slow, supernumerary individuals are killed off. Usually most of

the young produced die at an early age. In the terms of the insurance
salesman, their life expectancy is low. Man is a notable exception.
Civilization has brought about an increasingly
higher life expectancy
in man and with economic and which
sociological consequences to
we must adjust.
The great majority of plants and animals in nature die young, not
because of flaws in their internal mechanism, but because of their
failure to cope successfully with the external world. When fish eggs
are raised under the best conditions in a hatchery, most of them sur-
vive; very few die because of a mechanical or a physiological break-
down in their embryological development. When seeds are sown
in a well-prepared greenhouse bed, germination of most of the popu-
lation is expected. On a bag of grass seed, you will usually find a
statement that tests of the seed have shown a germination of 80 to 95
per cent. In a laboratory culture the larvae of a marine copepod were
raised to the adult stage with an 80 per cent survival, although
a survival of less than 0.5 per cent would have been sufficient to main-
tain the population on the basis of the number of eggs laid (Johnson
and Olson, 1948 ) In the laboratory the copepod was protected from
.
predators, diseases, and the exigencies of the physical and chemical

factors of its usual environment, which would normally have killed
off more than 995 out of every 1000 young animals.
Of course, some individuals are born with congenital flaws in their
anatomy or physiology. Little information exists on the frequency

with which lethal genes occur in wild populations, but their influence
is
probably very slight. Studies of wild fruit flies indicate that only
about 20 per cent of the eggs laid fail to develop into adult insects
because of genetic causes. Most of these "lethals" succumb because
the genetic changes cause failures in their relations to the environment,
rather than failures in their internal adjustments.
The few animals or plants that have escaped death long enough
to reach maturity are leading a very precarious existence. sword A
of Damocles hangs over their heads! The threat is ever present that
the action of the environment may become a little more severe and
wipe the population out. This sometimes happens. A striking ex-
ample of wholesale destruction of a bird population occurred in Min-
nesota when hundreds of Lapland longspurs were killed during a late
winter storm (Fig. 1.4). Another illustration of mass mortality is the
suffocation of fish under the ice of a lake (Fig. 1.5). In these in-
stances certain physical features of the environment became too
severe and the entire local population was killed.
Sometimes a biological influence goes beyond tolerable limits.
Older residents of northeastern United States

,
tell of Sunday excur-
^JOT^* ^
^m^^'^l:^^^^^^;^
FIG, 1,4.
;
:\îi ,,,^
-V:;!. *>
.
Dead Lapland longspurs on the frozen surface of a Minnesota lake

after a late winter storm.
Prescott, 1939, AAAS Pub. No, 10

FIG. 1,5 Masses of dead fish washed ashore breakup of the ice in Lake
after the
East Okoboji in northwestern Iowa. Bass, perch, carp, and other fish were killed
by the depletion of oxygen under the ice following the decay of a heavy growth
of algae.
sions into extensive groves of edible chestnut trees to gather nuts.
Now an edible chestnut tree is a rarity because a blight on this species,
caused by a fungus, spread throughout the northwestern states and
wiped out the chestnut groves between 1910 and 1930.
If the slight, but crucial, increase in the destructive action of the
environment extends beyond a local area to the whole range of a
species, its effect becomes much more serious, and it may even result
in the extinction of the species. This, too, has happened, not once,
but many Within
times. the memory of man such species as the
Labrador duck and the passenger pigeon have become extinct; many
more, such as the American bison, would have been entirely wiped
out if it were not for protected preserves. No one knows how many
species during the ages failed to meet the challenge of the environ-
ment. We
do know that about 21,000 species of extinct vertebrates
and an even larger number of extinct higher plants have been de-
scribed. we add to this figure a guess as to the number of extinct
If
vertebrates and higher plants whose remains were never found, and
of the extinct species of invertebrates and of lower plants, we shall
have some impression of the precariousness of existence in this world.
As if the destructive action of the environment were not serious
enough under natural conditions, man has added immeasurably to it
as civilization has "advanced." As will appear many times in our
further discussions, man need not necessarily be the destroyer. With
an adequate understanding of ecological principles he can utilize
many natural resources without impairing the*n. In some instances
the abundance and variety of the natural fauna and flora have im-
proved as a result of man's activities. If we include cultivation, a

very great development of plant and animal life has, of course, been
brought about by the hand of man. Unfortunately the intelligent use
of biological resources has not been the rule. Whole populations of
animals and birds have been slaughtered for their fur or their feathers.
Grasslands have been ruined, forests have been cut down (Fig. 1.6).
Besides direct devastation of much of our natural vegetation and
wildlife, we have wrought even more harm through causing serious
pollution of lakes, rivers, and harbors, irreparable damage to the soil,
and loss of ground-water reserves (Fig. 1.7). The sad story of man's
destruction of his own natural resources throughout the world has
been effectively told. The need for intelligent conservation and the
steps being taken in the United States to achieve it have been sum-
marized by Gustafson, Guise, Hamilton, and Ries (1949), by G. H.
Smith (1950), and others. Suffice it here to stress once more the fact
that animals and plants in nature are living dangerously. The natural
U. S. Forest Service
FIG. 1.6. Advanced degree of land destruction near Leadville, Colorado, where
forest has been ruined by clear-cutting, and the complete removal of trees has
resulted in gullying and serious soil erosion.
FIG. 1.7. and consequent farm abandonment in Oklahoma, showing

Soil erosion
disastrous economic and sociological effects of agricultural practices carried on
without regard to the ecological limitations of the region.
environment causes the death of most young organisms brought into
the world, and sometimes it may vary so as to kill all members of a
population. Man is an unavoidable part of the environment. With

intelligence he may help guard the danger spots he may even improve
the productivity of natural environments. Without intelligence man
accelerates destruction.
In the preceding examples of mass mortality and extinction the
devastating action which the environment can have, with and without
the presence of man, is obvious to all. For every spectacular instance
of this sort, there are thousands of instances in which no cataclysmic
destruction is
taking place, but in which the forces of the environ-
ment are nevertheless exerting a crucial influence in some subtle or
obscure manner.
This undercover work of the agents of the environment may perhaps
be most easily appreciated first in its action on individual species.
The geographical range of each species, for example, is controlled by
the pruning action of external forces. Every species is
pressing
against its boundaries and always tending to extend its range.
is
It is held in check
by the physical and biological factors of the environ-
ment which kill off those individuals which spread out too far into
areas where conditions arc no longer tolerable. During periods when
no unusual fluctuations occur at the limits of the range inhabited by
a species, the destructive effect of the environment may not be spec-
tacular because ordinarily only a few individuals are eliminated at any
one time or place.
The environment may thus control crucially, but quietly, the
geographical range of a species. Fluctuations in the environment
may even allow the limits of distribution to change somewhat as

conditions vary without a particularly noticeable mortality. One
example of that possibility is found in the commercial fishery records

for the landings of the weakfish along our middle Atlantic coast.
This fish, which is sold in New York markets as "sea trout," was
caught only in waters south of Cape Cod previous to 1895. During
that year and in succeeding seasons weakfish were taken north of the
Cape, and by 1901 the catch was large enough to support a regular
fishery. By 1907, however, the numbers of this fish had dropped
again to such a low figure that the fishery was abandoned, and never
established north of Cape Cod. There was no dramatic
again
change ocean
in the waters around the Cape during those six years.
Some subtle, unnoticed variation in the ecological conditions allowed
the weakfish to extend its range a short distance for a little while, and
then caused the northern limit of the species to be drawn back again.
Ecology of Plants and of Animals 13
However, the environment sometimes changes so as to allow huge

variations in the numbers of organisms, such as plagues of mice or
insect pests, as will be discussed later. More generally, however,
the fluctuations in the surroundings and in the abundance of plants
and animals are less violent. In the great majority of these unspec-
tacular oscillations it is nevertheless the environment that is
exerting
the vital control of the numbers of each species. In order to discover
the causes of fluctuations, which are sometimes of great economic
importance, a clear understanding of the essential relations between

the organism and its environment must be obtained.
The everyday action of the environment in curtailing the geographi-
cal spread and the abundance of organisms has other significant
effects. The production of more young plant seedlings and young
animals than can survive causes a continual struggle for existence,
with the resulting survival of the fittest. These are essential factors
in the evolutionary process as was pointed out
by a prominent "ecol-
ogist" by the name of Darwin, long before the term ecology came into
common use. Through its influence on the individual the environ-
ment indirectly controls the natural community. It determines the
complexion of the whole population by determining which species

can exist in each area and the relative numbers of each. Through the
control of feeding, growth, and other activities of each living com-
ponent, the environment regulates the dynamic operation of the

community.
THE DEVELOPMENT OF ECOLOGY

The Ecology of Plants and of Animals
The term "oecology" was first used by the German zoologist Haeckel
in 1869. The word did not appear again until 1895, when a report
on ecological plant geography was published by Warming, a Danish
botanist. The term in its modern spelling was taken up again later
by the zoologists. In those early days and for a long period there-
after botanists and zoologists were often working quite separately.
It is not
surprising, therefore, that plant ecology and animal ecology
tended to develop independently, but it is unfortunate that this divi-
sion into two separate fields tended to persist. We have now come to
realize that a proper understanding of the ecology of animals neces-
sarily involves a
consideration of the plants of the environment, and
that astudy of the ecology of plants would be incomplete without in-
cluding the influence of

animals. However, ecologists, and biologists
in general,have been slow to point out that many of the principles of
ecology are the same for both plants and animals.
The modern ecologist strives to understand the fundamental in-
fluence of the factors of the environment and to delineate such gen-
eral concepts as limiting action, competition, population growth, and
the like.These principles may be applied to the plants or to the
animals of the region under investigation, but for many concepts,
such as those of the food chain and the dynamics of energy exchange,
both the plants and the animals must be considered. The term ecol-
ogy thus necessarily includes the interrelations of all kinds of organisms
with the environment.
The Ecology of Habitats and of Individuals
In the early development of the subject, one group of ecologists

concentrated on the relations of the habitat. They strove toward a
better description of the habitat and of the influences of the habitat
on the plants and animals that lived there. First the physical fea-
tures and then the biological influences were investigated. The study
of the habitat and its effects is often spoken of as habitat ecology.
The work of other early ecologists, instead of beginning with a
description of environments, took its departure from an investigation
of the individual plant or animal. Attention was focused on the needs
and the reactions of the organism and the influence of environmental
factors upon it. Thus was developed a study of the ecology of the
individual, or to use a specially coined term, autecology.
The Ecology of Populations and of Communities
While the habitat ecologists were hard at work, other biologists with
ecological leanings were turning their attention to the fact that new
interrelations appear when groups arise. No animal or plant lives as
a completely isolated individual. When groups of the same species
are formed new effects appear. A simple example will illustrate this
point. Suppose that 100 trees are growing as individuals widely
spaced in a pasture. The shade from each of these trees will move
around during the course of the day so that the ground beneath the
trees will receive direct sunlight at least for a time each day. On the
other hand, if these same 100 trees were growing close together in a
grove, the shadow of one would overlap that of the next, with the
result that continuous shade would exist underneath. The effect of
the trees in the grove on temperature of the soil, evaporation, and
Ecology of Populations and Communities 15
wind conditions would be entirely different from that produced by

widely spaced trees. The interdependencies resulting from an aggre-
gation of individuals of the same species may become very complex,
colony. The size of a population
as for example in an insect of ani-
mals or plants and its rate of growth are regulated by the reactions
of the members of the population to each other and to the environ-
ment. The study of these and similar relationships of groups of or-
ganisms termed population ecology.
is
When
several species of
plants and animals are present,
as is usual
in a natural
community, still further complications arise. In the
example above a very different vegetation would exist beneath

cited
the 100 trees growing as isolated individuals from that found on the
floor of a dense grove. The species of animals associated with the
two situations. Widely varying
plants will also differ widely in the
combinations of plants and animals coexist in the many different
habitats of the world. It is found that certain species live together
in mutual adjustment, and these are spoken of as a natural community.
The study of the relationships of the animals and plants making up a
community is termed community ecology or, again to use a coined
term, synecology.
The development of the different viewpoints in ecology mentioned
above is due in part to the fact that the plant ecologist and the animal
ecologist have tended to work rather independently, The community

of plants is perhaps more obvious than the community of animals.
With plants the vegetation as a whole is often more striking, whereas
with animals the individuals tend to be considered first. Since animal
communities were generally less apparent, they were delineated at a
later date. The result of these influences was that animal and plant
communities were first thought of and studied quite separately.
As ecology developed, it came to be realized that the animals of an
area do not constitute a community entirely distinct from the plants
It is true that in some situations, as for
of that area. example in the
desert, the interrelations between the animals and the plants may be
less critical than the dependence of both upon the physical factors of
the environment. Nevertheless, because of the fundamental depend-
ence of animals upon green plants and the influences commonly
effective in the reverse direction, the plants and animals of a region
should be considered as one integrated community. The animal
taxonomistlists the fauna in a region and the plant taxonomist records
the flora. The fauna and flora together are spoken of as the biota of
the region. In an analogous fashion the modern ecologist considers
the integrated community of plants and animals as the biotic com-
It cannot be
munity. emphasized too strongly that the community
exists because of the suitable reactions of the individuals which make
up the community. Therefore, no sharp line exists between the ecol-
ogy of the individual and community ecology. All the foregoing
concepts will be discussed more fully and illustrated in subsequent
chapters.
The Ecological Complex
We have traced the development of the concept of the biotic com-

munity as the complete assemblage of interdependent plants and ani-
mals inhabiting an area. As a further step it came to be realized that
the physical conditions of the area must be considered for the com-
munity just as for the individual. (fThe organisms interact with each
other and also with the physical conditions that are present. Thus
organisms and the physical features of the habitat form an ecological

complex, or, more briefly, an ecosystem. J
In ecology, as in other subjects, the descriptive view appeared first.

Lists of the animals and plants present in characteristic situations
were prepared, and values were reported for the physical and chemical
conditions of the areas. In time all the important habitats became
subjects of special study, including the forests, the grasslands, the
deserts, the mountains, and the aquatic regions. Investigations of
certain definite habitats gave rise to such sciences as forestry, ocean-
ography, and limnology. In other instances studies of special types

of environment were not given specific names.
As various characteristic environments and their biotas were in-
vestigated more intensively, the realization developed that the eco-
logical complex should not be viewed as a static group of animals and
plants with the accompanying climatic conditions. The ecosystem is

not a museum group remaining immovable and unchanged as genera-
tions of observers pass by the plate glass windows. The community
cannot continue to exist without exchanges and interdependencies
any more than the individual plant or animal can. The community,
as well as the individual organism, is "something happening." Thus
the functional viewpoint gained momentum as modern ecology de-
veloped. Sears (1939) nearly emphasized this point by stating,

"When the ecologist enters a forest or a meadow, he sees not merely
what is there, but what is happening there."
As the action of the environment was studied in further detail it was
found convenient to list its influences as ecological factors. The stu-
The Ecological Complex 17
dent may have at first gained

an impression of all the factors of the
environment impinging upon the
organism in a one-sided action.
With further understanding of the natural situation, however, it came
to be realized that not
only do these factors affect the organism, but
also the organism affects its environment. This reciprocal action is
seen of all in relation to the For

physical features of the habitat.
first
example, light plays a part in controlling the growth of trees, and

at the same time the trees control the amount of beneath
light
them. Similarly, the dissolved nutrients and in water pond
oxygen
aflect the growth of the aquatic organisms that live there, but the very
activity of these organisms in turn modify these factors. The growth
of plants depletes the
supply of nutrient salts. The respiration of ani-
mals consumes oxygen and increases the amount of carbon dioxide in
the water. Thus the environment receives materials from the organ-
isms living in it and loses material to them as they grow. The fact
that regular changes in the environment are brought about by the life
activities of the inhabitants was
especially emphasized in a series of
Lowell Institute lectures given by A. C. Redfield in 1941, which he
entitled The
Physiology of the Environment.
The animals and plants modify the biological features of their
environment do the non-living factors. In thinking of the
just as they
;
activity of a carnivore, such as a fox preying on rabbits, we realize first
perhaps the influence of the predatory action in killing the rabbits and
depleting their numbers^ But by turning the picture around it be-
comes clear that the abundance of rabbits also influences the fox popu-
lation. A rapid growth of the individual foxes and a high rate of
reproduction is made possible if the supply of rabbits for food is
large,
but the number of foxes may be drastically curtailed if rabbits be-
come extremely scarce. In the same way when we observe a flock of
sheep in a pasture, we tend to think first of the activity of the sheep
in grazing down the grass. It is also true, however, that the sheep's
sharp teeth are clipping off tree seedlings
which may be sprout-
ing in the turf and the animals are adding manure to the pasture.
If it were not for the presence of the sheep, the continued existence
of the turf would often no longer be possible. In many situations
trees would seed in, and as a forest grew up the turf would be killed
off.
Clearly the sheep
and the plants of the turf form an integrated
system.
The concept that organisms and their environment form a recipro-
the viewpoint of most modern ecologists.
cating system represents
In every natural situation the environment affects the organisms pres-
ent and to a greater or lesser extent, the organisms affect the environ-
ment. The accompanying terms were proposed by Clements to de-
scribe the several aspects of the foregoing
relationships.
"Action" = habitat >

organism
"Reaction" = organism >
physical factors
"Co-action" = organism organism
The community, which Sears (1950) aptly refers to as "the living
landscape," maintains itself as a working unit with all the necessary
exchanges going on, more or less in balance, but in a dynamic and not
a static balance. The functional concept of the community and of
the two-way reaction between the environment and its inhabitants
carries us far beyond the descriptive view. The improvement gained
from this modern approach in ecology is analogous to the better
understanding of the conditions inside an individual animal or plant
is obtained when the
that physiologist's viewpoint and technique are
added to those of the anatomist. Modern ecology might thus be
thought of as the "physiology" of the ecological complex in the sense

that it deals with the functional aspects of the interactions, exchanges,
and adjustments of the members of the community arid of their
environment.
THE SCOPE OF ECOLOGY

Taylor (1936) has said "Ecology is the science of all the relations
of all the organisms to all their environment." Since the plant and
animal inhabitants very abundant and diverse, and since en-
may be
vironmental conditions are extremely variable, the possible scope of
ecology becomes very great. The central task of ecology, however,
is to delineate the
general principles under which the natural com-
munity and its
component parts operate. These may then be applied
to the interpretation of the activities of the particular plants and ani-
mals present under the existing specific conditions of a given stiuation.
Although the fauna and flora of an area must be identified and
enumerated, and although the physical forces at work in the area must
be recognized, neither an account of the biota, nor a description of
the habitat constitutes an ecological investigation. Similarly, if a man
arises at daybreak and makes a list of the birds he sees without any
consideration of the relation of the occurrence of these species to
other factors, he is not an ecologist. Modern ecology is concerned
with the functional interdependencies between livinfe things and their
The Scope of Ecology 19
surroundings. Ecology is primarily a field subject. Nevertheless, a

knowledge of the principles and problems of ecology should be ac-
quired before attempting to evaluate a natural situation, where the
multiplicity of ecological activitiesmay be bewildering. Many eco-
logical relationships can be effectively analyzed under the simplified
and controlled conditions of the laboratory.
Because of a lack of understanding of ecological principles the
efforts of well-intentioned conservationists and
agriculturalists are
frequently badly misdirected. Astory is told of certain sheep ranchers
who became convinced that coyotes were robbing them of their young
sheep. As a result, the community rose up and by every possible

means slaughtered all the coyotes that could be located for miles
around. Following the destruction of the coyotes, the rabbits, field
mice, and other small rodents of the region increased tremendously

and made serious inroads upon the grass of the pastures. When this
development was sheep men executed an about-face,
realized, the
abruptly stopped killing the coyotes, and instituted an elaborate pro-

gram for the poisoning of the rodents. The coyotes filtered in from
surrounding areas and multiplied, but finding their natural rodent

food now scarce they were forced to turn to the young sheep as their
only available source of food!
An understanding of ecological principles provides a background
for further investigations not only into the fundamental relationships
of the natural community but also into sciences dealing with particular
environments such as the forest, soil, ocean, or inland waters. Many
practical applications of ecology are found in agriculture, biological
surveys, game management, pest control, forestry, and fishery biology.
Knowledge of ecology is
critically important for intelligent conserva-
tion whether in relation to soil, forest, wildlife, water supply, or
fishery resources.
Ecology is significant also in a wider sense for us as citizens. It
gives us an insight into how the world works. In addition, man him-
self is a most important element in the environment. Man almost
always has a modifying influence, and, without proper regulation, he
often has a destructive effect. Man is himself an organism with an
environment, and this fact has been particularly emphasized in the
development of human ecology. A knowledge of the general prin-
ciples of ecology thus provides a background for the understanding
of human relations just as a study of general zoology is necessary as a
groundwork for medicine. Like other animals man is influenced by

the physical features of his environment, he is absolutely dependent
upon other species, and he must adjust to other individuals of his

own species, At the moment man is
suffering from lack of these
adjustments.
APPROACH TO THE STUDY OF ECOLOGY

The study of ecology is best begun through the analytical approach.
This involves the delineation of the individual influences of the en-
vironment and the recognition separately of the various activities of
the organisms present as steps toward building an understanding of
the entire dynamic interaction between the complete environment
and its inhabitants.
The fundamental relationships are most readily grasped by analyz-
ing the simplest situations first. Contrast, if you will, the ecological
dependencies of an alga living near the surface in the open ocean
with those of a tree growing on land. The tissues of the alga receive
theirenergy supply directly from the sun and they carry on their inter-
change of materials directly with the surrounding water, which is
uniform and extremely constant in respect to the ecological factors
concerned. The tree, on the other hand, is partly in the light and
partly in the dark. Part of the tree is surrounded by the atmosphere
with its
widely fluctuating temperature and humidity; part is in the
soil, where subject to a very different temperature and is alter-

it is
nately flooded with air and with water. The part of the tree that is
above ground must deal with one set of organisms, and the part of the
tree below ground is concerned with an almost entirely different set.
Another reason for adopting the analytical approach is that this
procedure is more likely to reveal limiting factors. All animals and

plants tend to grow, to reproduce, and to disperse until checked by
some influence of the environment. The factor that first stops the
growth or spread of the organism is called the limiting factor. It is
not always easy to single out the limiting factor, and sometimes two
or more factors combine to provide the limiting influence. Never-
theless, extremely desirable whenever possible to determine what
it is
agent or agents control the natural tendency of the plants and animals
present to increase in size, numbers, and range. In the investigation
of any natural area correlations will be found between features of
the environment and the activities of organisms present. Analysis of
the action of individual influences at work in the habitat is necessary
in determining which of the correlated factors are actually causal
factors. Suppose, for example, that we discovered a correlation be-
tween the occurrence of the factor A and organism B. Should we
conclude that A causes B?
Approach to the Study of Ecology 21
A-+B
It might very well be that no direct causal relation exists between
A and B whatsoever, but that both are controlled by a third influ-
ence, C.
Or factor A may influence C,

A
v
which
B
in turn influences B, thus:
A B Temperature Plant
\ C/ \ Moisture/
.
In considering the factors of the environment separately in order to
distinguish and to measure the influence of each, we must remain

thoroughly aware that in nature the factors are never acting alone.
Animals and plants are subject to many influences at the same time,
and the effect of one factor is often modified by action of other factors.
The "real life" of the organism, on which its
growth, distribution, and
multiplication depend, necessarily involves the simultaneous and con-
tinuing impact of all existing factors and also influences that occurred
at earlier
stages in the organism's experience.
An important difference exists in the extent to which factors can be
modified by living organisms. Some features of the environment are
largely unaffected by the activities of the organisms present; these are
unmodifiable or conservative factors. The salinity of the ocean is an
example of a conservative factor. The volume of the ocean is so great
that, although animals and plants living in it are continually adding
or withdrawing salts, the amounts have an immeasurably small effect
upon the total salt content of the water. The modifiable or non-con-
servative factors of the environment are susceptible to change caused
by the inhabitants of the area. The oxygen in a small pond, for ex-
ample, may be
so depleted by the respiration of a large population of
an unfavorable or even a lethal condition for the fish is pro-
fish that
duced; or the concentration of oxygen in the pond may be increased

of algae a modification that will benefit the
by the photosynthesis
fish. Heather (Calluna) tends to increase greatly the acidity of the
soil in which it is growing, and this condition favors the further de-
velopment of this plant, but it is unfavorable to most other plants.

Through modification of its own environment, heather often comes to
dominate the vegetation in large areas, as may be seen in Jutland.
No sharp division exists between modifiable and unmodifiable fac-
tors. All gradations exist, and a given factor may be modifiable in

some situations or for some organisms and quite unmodifiable for
others. In the succeeding chapters the more general, often unmodi-
fiable, factors will be scrutinized first, and will be followed by a
discussion of the more commonly modifiable factors. This will lay
the foundation for a consideration in the later chapters of the com-
positionand functioning of the community as a whole.
2
The Medium
The first of the

physical features of the environment to be considered
will be the medium that is, the material which immediately sjucrounds
;.
the organism and with which it has its all important exchange. At
first
sight one might think that many diverse media exist. Some
organisms live in the soil, and some in ponds; some thrive in manure
piles, and others enjoy a successful existence in the blood stream of
vertebrate animals. Certain nematodes live in vinegar, and a fly
larva of the genus Psilopa grows in petroleum. Once during a de-
partmental gathering at Cambridge University a member of the staff
entered the room waving a journal in which the habits of this larva
were reported. "Look here," he said, "in this report an insect is
described which lives in petroleum. The first thing you know it will
parasitize our motor cars!"
The medium in each of the above examples, and indeed that for
organisms in every natural situation, is either a liquid or a gas, and it
i.s
usually air or water. Although animals and plants inhabiting soil
or mud may at first appear to be exceptions, a closer scrutiny shows
that a film of air or water around each organism is actually the ma-
terial in immediate contact with it. An enlarged view of the small
animals living in the wet sand of the seashore shows that their essen-
tial exchange is with the water percolating between the sand
grains
and that the medium for these animals is sea water, not sand (Fig.
2.1). The term medium is thus used in a strict sense and is distin-
guished from the substratum, or surface on or in which the organism
lives.
The existence of air and water as the fundamental media divides
the world into two major environments: terrestrial and aquatic. The
media are not completely isolated from each other, however; some of
the atmospheric gases are dissolved in all natural waters, and some
in the atmosphere.
moisture is
present almost everywhere Differences
23
24 The Medium
in the amount of intermixture play a part in
subdividing the terrestrial
environment into arid and humid climates and the aquatic environ-
ment into stagnated and aerated water. Transition areas of special
interest exist such as
swamps and the tidal zonewhere sometimes
one medium and sometimes the other dominates the scene. Dams
builtby beavers occasionally result in the flooding of large tracts of
1mm.
Pennak, 1939, AAAS Pub. No. 10

FIG. 2.1. Enlarged diagram of sand in a beach habitat showing water-filled spaces
between the grains. 1= rotifers, 2 =gastrotrichs, 3
=
tartigrade, 4 nematode,
5 = harpacticoid copepods.
land, transforming them from terrestrial to aquatic habitats (Fig.

2.2); conversely, the growth of vegetation often tends to fill up a
shallow pond, gradually converting it into a swamp, and eventually
into dry land. Ordinarily, however, the medium is a highly inde-
pendent factor, for rarely do the activities of organisms cause a change
from one basic medium to the other.
CONTRASTING QUALITIES OF AIR AND WATER

The two fundamental media are very different in nature, and this
difference has important ecological consequences. is

composed
Âir
Contrasting Qualities of Air and Water 25
of 79 per cent nitrogen, 21 per cent oxygen, 0.03 per cent carbon
dioxide, and several other gases in much smaller quantities^ These
gases are not chemically combined, but exist as a simple physical

mixture. Water, by contrast, consists primarily of a single compound,
H 2 O. There is nothing especially unusual about the physical and
chemical properties of air and the gases of which it is
composed.
U. S. Forest Service
FIG. 2.2. Pond formed by beaver dam (left foreground), showing beaver house
(right center) and trees felled and stripped by the beavers. Cochetopa National
Forest, Colorado.
Water, on the other hand, is a unique substance from the ecological

viewpoint. H. B. Bigelow, formerly director of the Woods Hole
Occanographic Institution, when lecturing on oceanic biology once
stated: "The most important fact about the ocean is that it is full of
water!"
The unusual detail by Henderson
qualities of water are discussed in
(1924) in his classic book, The Fitness of the Environment. Suffice
it here to mention a few of the attributes of water that have a special

ecological importance. In the first place
water is the most abundant
substance of the earth's surface, covering more than 70 per cent of the
area of the globe. Because the oceans are about 2%
times more
extensive than the land, and because they are habitable throughout
their depth, the sea provides more than 300 times the living space.
Water has a higher specific heat, latent heat of fusion, and latent
26 The Medium
heat of evaporation than any other common substance. These facts
play a very important role in the heat regulation of organisms them-

selves and in the resistance of natural environments to temperature
change.
Another characteristic of water having crucial ecological significance
is its
relatively high freezing point. Because of the large amount of
heat which must be given up before water can turn to ice and because
of restricted stirring, oceans and lakes freeze only at the surface.
Even ponds rarely freeze to the bottom. The temperature of the
medium, therefore, can drop only to 0C in fresh-water environments,
or to a few degrees lower in the ocean. The biological reactions of
a great many plants and animals can still go on perfectly well at
temperatures down to the freezing point of water.

Another unusual quality of water is its power as a solvent; no other
common substance compares with water in this respect. Many kinds
of material can pass into, through, and out of the body of an organism
in aqueous solution. Water provides a transporting medium that is
versatile as a solvent but not too active chemically. Very consid-

erable solution would take place if
sulphuric acid ran in our rivers or
coursed through our veins, but such a solvent would profoundly
alter the materials that
it carried. Furthermore, the extent of ioniza-
tion of solutes in water is extremely high, providing the possibility
of a great variety of radicals and of chemical recombination. Water
has the highest surface tension of any common substance except
mercury. This high surface tension has many ecological influences,

involving the movement of water into and through organisms as well
as the rise of ground water in the soil.
Many of the foregoing differences and special qualities of air and

water will be referred to again in connection with other factors of
the environment. Confining ourselves for the moment to the simple
physical differences between the two fundamental media, let us ex-
plore the ecological effect of the difference in density of air and
water. The densities of representative natural waters and of air,
and the approximate average density of protoplasm are as tabulated.
Pure water 1 . 000 g/cc at 4C

Pond water 1.001
Sea water (at salinity of 35%c) 1 028
.
Air (at sea level) . 0013

Protoplasm 1 . 028
The density of protoplasm is closely similar to that of sea water and

only slightly greater than that of fresh water, but it is more than 850
times greater than that of air. Associated with this difference in
Pressure 27
density are important differences in the pressure, inertia, viscosity,

and mobility of the media.
PRESSURE
The difference in the densities of the media results in a great
difference in the rate of change of pressure at increasing altitudes in
the atmosphere and at increasing depths in the water. Near the
earth's surface a rise of 300 m (1000 ft) in altitude results in a reduc-
tion of pressure of about 25 mm Hg, or a relatively slight change in
f- 25,400 m -
Rocket-powered plane
22 mm Hg I- 22,000 m -
Stratosphere balloon
235 mm Hg r 8840 m (29,000 ft)

-
Mt Everest
310 mm Hg - 7000 m Vultures and eagles
AIR 367 mm Hg - 5800 m Wild sheep and ibex
413 mm Hg - 4860 m Highest human settlement, Tibet

- 4420 m -
Mt Whitney, California
Rates of change:
25 mm Hg/300 m 1920 m -
Mt Washington, N. H.
760 mm Hg - 1 Atmos. Sea level
925 m Deepest dive of bathysphere (Beebe)

1400 m Deepest dive of benthoscope (Barton)
1 Atmos/10 m
370 Atmos. 3700 m Average depth of oceans

4050 m Deepest dive of bathyscaphe (Houot)
WATER
625 Atmos. 6250 m Ten species of animals taken by
"Challenger"
_ 10,500 m Various invertebrates taken by "Galathea"

1086 Atmos. L 10,860 m (35,640 ft) Greatest ocean depth
(Mariana Trench)
FIG. 2.3. Range of pressure in air and water in relation to the distribution of life.
28 The Medium
pressure. In contrast, for every increase of 10 m(33 ft) in depth in

the water, pressure is increased
by 760 mm
Hg, or 1
atmosphere. The
tremendous pressures existing at the average depth of the ocean and
in the ocean
deeps are indicated in Fig. 2.3. At a depth of only 900
m in the ocean Beebe's bathysphere was subjected to a total pressure
on its whole surface of more than 7000 tons. It is not that
surprising
in his
explorations of the earth man has not been able to descend into
the ocean to depths much
greater than 4 km, whereas he has ascended
in the
atmosphere to heights of more than 22 km. What are the
ecological effects of these differences in pressure and in the rates of
pressure change in the two media?
Pressure Reduction with Altitude
The reduction of pressure with altitude

seems to be of little impor-
tance for plants, invertebrate animals, and the lower vertebrates.
Insects have been subjected
experimentally to a reduction of pressure
from 760 mm Hg to 0.0001 mm Hg without harmful effect. Similarly,
frogs have withstood reduction of pressure down to 100 mm
Hg.
Beetles reach the highest meadows in the
Himalayas and earthworms
are found up to the snow line in the Andes. In most situations it is
not the reduced
pressure which limits the altitude at which plants and
cold-blooded animals can exist. The distribution of these
organisms
up the sides of mountains is ordinarily stopped by other adverse fac-
tors, such as low temperature, unsuitable soil, or lack of food,
long
before the influence of the reduced
pressure is felt.
For warm-blooded vertebrates the reduction of
pressure with alti-
tude becomes important primarily because of the lesser amount of
It is true that at
oxygen present. very high altitudes the thinness
of the air renders
flying more difficult, but the chief limitation imposed
on birds, and also on mammals, the impairment of
is
respiration.
Distinctly harmful effects are observed for man when the pressure of
the atmosphere has been reduced to about half that normal at sea
level. The highest permanent human settlement occurs in Tibet at
an altitude of about 5000 m. Even the best
adapted of other mam-
mals are not found living permanently much
higher than this altitude
(Hesse, Allee, and Schmidt, 1951, Ch. 24). Although vultures and
eagles have been reported at about a thousand meters higher, they
probably remain at such altitudes for only short periods of time.
Insects, which often abound near the rocky or snow-covered
peaks of
mountains, are undoubtedly blown there by the wind, a^ they could
Pressure Increase with
Depth 29
not survive long under the conditions of the low
temperature and
lack of food,
Pressure Increase with Depth
the relatively slight reduction in

pressure with altitude in air is
If
important to some organisms, the tremendously greater increase in

pressure with depth in water might be expected to have serious con-
sequences to all
aquatic organisms, When early calculations were
made of the magnitude of the pressure at the bottom of deep lakes,
and particularly of the ocean, it was believed that the stupendous
pressures would annihilate all living beings, so that the greatest depths
in the aquatic environment must be lifeless, This conclusion seemed
at first to be confirmed
by the early explorations, but, with the im-
provement of gear for investigating the bottom of the deep sea, animal
life was
gradually discovered at greater and greater depths in all the
ocean basins (Fig, 3.5), In 1951 the Danish Galathea Expedition
trawled 17 sea anemones, 61 sea cucumbers, 2 bivalves, and 1
crustacean from a depth of about 10,500 m off the Philippine Islands.
At this depth the pressure is 1050 atmospheres, or about 1 ton on each
square centimeter, but weight of water does not crush the
this terrific
organisms living at that depth because the pressure is the same inside
their bodies as outside.
When most deep-sea animals are brought to the surface, they are
dead or dying. The popular opinion is that they have been killed
by a violent release of pressure. In reporting the work of the re-
search vessel Atlantis, a newspaper once stated: "The sudden change
of pressure when deep-sea are brought to a higher level in the
fish
ocean causes them The fragments are then put together

to explode.
again!" Fish with air cavities within their bodies do indeed expand
when they are brought to a higher level, but most fish inhabiting the
ocean abyss have no air bladders. Their death is due primarily to
injury from the nets and
to the change in temperature experienced in
being brought to the surface.

The effect of the pressurechanges in the aquatic environment is
very different for organisms with
and without air cavities. Aquatic
that live at levels considerably below the surface stratum and
plants
the great majority of deep-sea animals do not have gas-filled spaces
in their bodies. Since any cavities in these organisms are completely
filled with is caused
no mechanical deformation
fluids, by pressure
because the watery tissues are only slightly compressible.
changes
30 The Medium
Green plants are confined to relatively shallow subsurface depths be-
cause of their need for light. Many types of animals, on the other
hand, display a very great vertical range in their distribution.
Several species of invertebrates are found at depths extending from
near the surface in the littoral zone down to 4000 or even 5000 m m
in the abyssobenthic zone (Fig. 2.4). Although the individuals of
these bottom-living forms do not travel far, the species as a whole
have become adapted to this great vertical range over a period of
-
Neritic Oceanic
FIG. 2.4. The chief zones of the marine environment. The division between the
neriticand oceanic provinces occurs at the edge of the continental shelf where
depth is about 200 m. The lower limit of the archibenthic zone occurs between
800 and 1100 m. The littoral zone forms the upper part of the neritic benthic
zone and usually receives strong wave and current action and sufficient light for
plant growth. The depth of its lower limit is variable but is often in the neighbor-
hood of 40 to 60 m. These divisions also apply in a general way to lakes.
(Modified from The Oceans by Sverdrup et al., 1942, copyright Prentice-Hall,
Inc., N. Y.)
time. In certain other kinds of marine animals the individuals are

known to change level over considerable distances and hence are
able to withstand correspondingly great pressure changes.
Some
species of fish move downward as much as 400 during the m
day and
swim up again each night, thus subjecting them-
to their former level
selves twice daily to a pressure change of 40 atmospheres. Certain
small planktonic Crustacea similarly carry out diurnal vertical migra-
tions of 200 m
to possibly 600 m
in amplitude (Waterman, Nunne-
macher, Chace, and Clarke, 1939).

The foregoing is not to imply that most species of marine organisms
do not have definite vertical limits to their distribution. Many species
Pressure Increase with Depth 31
are confined to
relatively narrow zones, but these restrictions of range
are probably due
primarily to other factors, such as temperature,
light, or food. Nevertheless very great changes in pressure have been
shown experimentally to alter the rates of certain physiological reac-
tions; and a variety of invertebrate animals, fishes without swim

bladders, and bacteria are inactivated, or killed, when subjected in
pressure chambers to several hundred atmospheres, although the ef-
is much reduced if the
fect
temperature remains nearly constant
(
Zobell and Oppenheimer, 1950 ) Thus, although moderate pressure
.
changes do not ordinarily harm such organisms, great changes may

exert certain subtle influences on their life processes. Whether or
not such physiological action of pressure actually limits the vertical
range of aquatic organisms in their natural habitats has not yet been
ascertained.
For animals with air-filled cavities, such as fish possessing swim
bladders and diving birds and mammals, the rapid increase in pres-
sure with depth in the aquatic environment is a serious matter.
The swim bladder of a fish supplies buoyancy, and a fish with this air
cavity is similar in its flotation to the Cartesian diver of the physicists.
When moves downward, the bladder is compressed, and when
the fish
it moves toward the surface, the bladder
expands. Gas must be re-
moved from the swim bladder, or added to it, in order for the fish to
maintain control of its buoyancy equilibrium. If the fish moves up-
ward so fast that gas cannot be removed from the swim bladder at a
rate sufficient tocompensate for the reduction in pressure, the con-
tained gas will continue to expand and the fish will rise toward the
surface at an accelerating rate. If movement toward the surface con-
tinues, the swim bladder will eventually burst, and the expanding
gas, now in the body cavity, will force the stomach to protrude from
the mouth, the intestine from the anus, and the eyes from their
sockets. Presumably, under ordinary conditions, an early stage of in-
ternal volume change stimulates the fish to return to its former level.
The pressure factor thus definitely limits the vertical range of fish
with swim bladders as well as the speed with which they can move
from one depth to another (Jones, 1952).
For diving mammals and birds the problem of breathing is added
to that of the increased pressure. When a human diver descends into
the water in a flexible diving suit he must withstand the increased
to him through a hose or by means of an
pressure, but air is supplied
"aqualung." Pressure alone prevents the diver from descending more
than 100 m
or so. Whales, seals, and diving birds are forced to go
without a renewal of oxygen during the period of their dive. Whales
32 The Medium
apparently withstand the great pressures by allowing their lungs to

be completely flattened and the air that was in them to be forced
into the strong, boxlike larynx. How a mammal swimming vigorously
can get along for more than a few minutes without a renewal of its
oxygen supply has been a matter for speculation for generations.

Evidence obtained primarily from the study of seals indicates that
these diving mammals can store an increased amount of oxygen in
their tissues and that this reserved for the brain, heart, and
oxygen is
other vital organs by cutting off the circulation to other parts of the
body. The muscles build up an oxygen debt that is paid off when
the animal surfaces again ( Scholander, 1940 ) Too rapid ascent may
.
result in the formation of gas bubbles in the blood, but the fact that
the whale, unlike the human diver, has only one lungful of air during
the dive presumably reduces the danger from absorbed nitrogen.
The depth and duration of the whale's dive have been hotly con-
tested by captains of whaling ships and others for many years. It
seems probable that whales dive to 200 or 400 m regularly. Seemingly
indisputable evidence for an even greater dive was furnished by the
discovery of a sperm whale that had become entangled at a depth of
about 1000 m in a submarine cable running between two of the Carib-
bean Islands. At this depth the pressure is about 100 atmospheres
(or over 100 kg per sq cm), and evidently the mechanism of the whale
is
adapted to withstand pressures as great as this. Whales ordinarily
stay submerged for twenty minutes or so, but when harpooned they
may disappear from the surface for one to two hours. Such dives,
accompanied as they are by violent swimming, are a striking demon-

stration of the ability of these animals to remain active for long pe-
riods without renewal of oxygen.
SUPPORT AND RESISTANCE TO MOTION

The difference in density of the two elemental media, air and water,
also influences the degree to which they provide support and re-
sistance to motion. Since water has nearly the same density as proto-
plasm, whereas air is

very much less dense, water furnishes much
more buoyancy than air.
Effects on Structure and Size
Since terrestrial organisms are very much heavier than their sur-
rounding medium, they would tend to collapse from their own weight
if it were not for
special supporting structures. On land only very
Effects on Structure and Size 33
small organisms and such animals as earthworms and
slugs can main-
tain their
shape without skeletal material of some sort. The woody
tissue characteristic of the
higher plants provides rigidity against the
force of gravity. The bones and muscles of the land animals
larger
are similarly arranged primarily to (Thompson,
provide support
1942).
Generally speaking, the weight of an organism tends to increase as
the cube of its linear dimensions, but the
strength of supporting
columns increases only as the square of the dimensions. As a result
animals and plants are definitely limited as to size. Since land plants,
once established, do not require locomotion, they can have a much
larger amount of rigid supporting tissue than animals. Hence the
plant kingdom holds the record for size on land. The giant redwoods
of California (Fig. 1.1) attain heights well over 100 (record height: m
365 ft), and the trunks alone are estimated to weigh as much as 500
or 600 tons. In the animal kingdom few modern species attain a size
as great as 6 or 7 tons, although the dinosaurs of the past were some-
what larger. Possibly Brontosaurus would have tipped the scales at
30 or 40 tons, but Brontosaurus came to an unhappy end, no doubt
in part because of
its
ungainly size.
In the water environment, since all parts of the organism tend to be
buoyed up by the medium, supporting structures may be greatly re-

duced or entirely lacking. Woody tissue is needed for support by
few aquatic plants. When an elaborate skeleton is present in aquatic
animals, it usually occurs for purposes other than support. For many
crustaceans and mollusks the hard tissues serve primarily as pro-
tection; in other forms, such as fish, the skeleton is used chiefly for
the attachment of muscles of propulsion. Many aquatic organisms
such as the jellyfish have no skeleton at all. true that the jellyfish
It is
is a weak and
sluggish organism but the octopus and the giant squid
are decidedly vigorous, and yet the skeletons of the latter are reduced
to horny pens and a few cartilages in the head region. An octopus
kept in an aquarium at the Bermuda Biological Station was so suc-
cessful in getting out of his tank in spite of a weighted lid that he was
named Houdini.
In the plant kingdom even the algae can grow to tremendous sizes.
The giant kelp Nereocystis,

common off the west coast of the United
States, may grow to a length of more than 35 m, and Macrocystis is
reported to attain an even greater size. Such plants have no woody

tissue whatsoever, but the water buoys up the extended parts of the
Animals in the sea today attain sizes larger than ever
organism.
existed on land. By weighing the parts of a blue whale being cut up
34 The Medium
on the deck of a factory ship Hjort (1937) obtained the tabulated
values, which do not include the blood and viscera.
Muscles 56 . tons
Bones 23 .
Blubber 26.
Tongue 3 .
Heart 0.6
108.6
If a whale becomes stranded on the beach, the weight of its body
prevents breathing by crushing the lungs, and its great strength is of

little avail for
getting it back into the water since its muscles of pro-
pulsion are adapted exclusively for swimming in the open sea.
Effects on Locomotion through Medium
Differences in the viscosity, mobility, and inertia of air and water

have profound effects upon the resistance of the medium to the motion
of organisms through it. In general the resistance of water is very
much greater than that of air but the actual value depends upon the
size and shape of the organism, the viscosity of the medium, and the
speed of locomotion. The coefficient of viscosity of water is 60

times that of air at the same temperature. The result is that an im-
portant resistance to locomotion is felt at very much lower speeds in

water than in Animals whose living depends upon rapid swim-
air.
ming through water must be thoroughly streamlined. In the mack-

erel, forexample, not only is the body almost perfectly streamlined
but also the fins fold back into grooves and the surfaces of the eyes
conform exactly to the contour of the head.
Another result of this great difference in the resistance of air and
water is that really high speeds can be attained by animals only in the
air environment, and even at low speeds very much more effort is re-
quired to move through water than through air. The speed record
for the animal kingdom is probably held by the duck hawk, whose
flight has been clocked at 288 km per hr ( 180 miles per hr ) Several .
other species of birds can fly at speeds greater than 160 km per hr,
but no running animal can approach these velocities. The gazelle
and the antelope are credited with speeds of 96 km per hr, and the
cheetah can do 112 km per hr (70 miles per hr) over short distances.
These catlike animals are employed by the natives in Africa to bring
down antelope for them. The natives steal up as close as possible to
Passage of Medium through Organism 35
a herd of antelope in an old Ford and then release the cheetah for
the last short dash,
In the aquatic environment fish of the mackerel tribe are the fastest
swimmers and attain speeds as great as 48 km
per hr (30 miles per
hr). The flying fish has been reported to attain 56 km
per hr just
before its take-off. Anyone who has watched a flying fish, however,
will remember that in the last moments before the fish leaves the
water most of body is in the air, with only the
its tail sculling violently
an outboard motor.
in the surface like
Even the method of propulsion is controlled to a large extent by
the elemental difference in the nature of the air and water media.
Since the density, viscosity, and inertia of air are so low, most animals
cannot use the air alone for propulsion but must obtain a purchase
on the earth's surface. Only birds, insects, and a few other animals
can propel themselves wholly in air. In the aquatic environment, on
the other hand, the majority of animals swim in the free water, and
those for which speed is important do not use the substratum for ef-
fective locomotion. The lobster, for example, pokes around on its
walking legs,
but to make a sudden dash it uses swift strokes of its
tail, letting legs leave the bottom entirely.

its Because of the rela-
tively great inertia of the water medium, some animals, such as jelly
fish and scallops, can propel themselves in one direction by pumping
water an early version of "jet propulsion."

in the opposite direction
The squid can dart backwards with remarkable rapidity by ejecting
water from its siphon.
Passage of Medium through Organism

Sometimes the medium must move through the organism instead
of, or in addition to, the movement of the organism through the
medium. Because of the great mobility of air, this medium can move
in and out of the cavities of an animal or plant with relative ease, but
water does not circulate as freely. Special adaptations are required
to carry water to the tops of trees, including root pressure, transpira-
tion, and the tensile strength of fine water columns. The movement
of the water through the tracheids of the plant is relatively slow.
Even in aquatic plants direct water exchange by osmosis or colloidal
imbibition requires considerable physical force.
Although large expenditure of work is necessary, some aquatic ani-

mals do succeed in causing water to flow through relatively simple
respiratory or feeding
chambers. But water could not possibly be
pumped into and out of a finely branched system of tubules like the
36 The Medium
tracheae of the insects with sufficient speed to meet respiratory
needs. No doubt partly for this reason relatively few kinds of insects
have succeeded in establishing themselves in the aquatic environment.
Most of these remain in the water for only a portion of their lives
usually the larval stage. Insects living completely submerged possess
tracheal gills or other special devices for obtaining oxygen without
taking water into the tracheae. Many fresh-water species are adapted
to come In the marine environment Halobates
to the surface for air.
lives on the surface of the ocean as a "water strider," and several
dipterans inhabit shallow areas as larvae, but only one insect, the
midge Pontomyia natans, is known to complete its entire life cycle
submerged in sea water.
Existence of Plankton
The relatively high density of the water medium not only tends
to buoy up parts of the body but also in some instances supports the
whole body and thus allows certain organisms to float at various
depths in the free water. This fact makes possible the existence of
plankton plants and animals that live suspended in the ocean and
inland water bodies and that drift about either because they are non-
motile or because they are too small or too weak to swim effectively
against the currents (Figs. 2.5 and 2.6). The term plankton is de-
rived from a Greek word meaning "wanderer," and many organisms
in this category, or plankters, spend their whole lives drifting in
the water. Both animal and plant plankton is found in practically all
natural waters, frequently in enormous abundance and variety.
Other categories of life in the aquatic environment are the benthos,
which consists of the organisms living on or in the bottom material,
and the nekton, which is composed of the strong-swimming animals.
The benthos and the nekton have their counterparts on land, but the
permanent plankton represents an important category of life that is
totally absent from the air environment.
Certain planktonic animals and plants live permanently suspended
in the water by actual flotation. This method of support is not pos-
sible in the air. Pollen grains, seeds, and spores, commonly spoken
of as "floating" in the air, are not actually doing so, but are sinking
at a slow rateoften retarded by various feathery structures. In the
water actual flotation is
possible for forms that contain air cavities or
light materials such as fats or oils. The brown alga Sargassum is pro-
vided with gas-filled bladders, and the Portuguese man-of-war, a
siphonophore, has a pneumatic "sail."
Many fish
eggs float
by virtue
of droplets of oil. Some pelagic diatoms completely counterbalance
the weight of their siliceous shells
by means of a cell sap which is
lighter thanwater so that these organisms have no tendency to sink.
In addition to the truly floating forms many other kinds of plants
and animals sink so slowly that they are able to lead a planktonic
*-* . 1-
k _ I /
/
,
;;p** ;>
f^ 5 -9
:
.1
Bigelow, 1926
FIG. 2.5. Marine phytoplankton. Photomicrograph of several common genera of

oceanic diatoms. The largest cell is approximately 0.05 mm in diameter.
life. In the air environment the sinking rate of the smallest organ-
isms, such as bacterial spores, is
extremely slow, but eventually all
out.In the water, however, the sinking rate of some
particles settle
multicellular animals, is so retarded that
organisms, including many
a small amount of swimming allows the organisms to maintain their
position.
In other instances the amount of sinking may be inconse-
quential before the is

organism brought up to the surface again by
vertical currents.
38 The Medium
Sinking rate is also controlled in part by size and shape. It might
be said that one of the simplest ways of reducing sinking speed is to

be small because with reduction in volume the surface and hence
surface friction becomes relatively greater. Any departure from the
spherical shape increases the surface of an organism. Unicellular
forms like the pelagic Globigerina which are bristling with spines
Vigelow,
FIG. 2,6. Marine zooplankton, Photomicrograph of common copepods, chaetog-
naths, and medusae. The euphausiid "shrimp" is approximately 2.5 cm (1 in.)
long.
graphically illustrate this fact (Fig. 2.7). Most multicellular animals

that live a planktonic life also are liberally provided with long an-
tennae, spines, and bristles of a great variety of shapes. The effect

of these changes in the morphology of planktonic organisms, com-
bined with differences in density, in reducing the sinking rate is il-
lustrated in Table 1.
./wurray ana njon, 1x11, copyngni jnacminan & Co.

FIG. 2.7. Globigerina bulloides, a planktonic protozoan belonging to the order
Foraminifera. The largest sphere of the shell is about 0.5 mm in diameter.
TABLE 1
COMPARISON OF SINKING RATES OF THREE TYPES OF PLANKTON ORGANISMS

WITH THOSE OF SPHERICAL SAND AND SILT PARTICLES
Length or
Diameter Sinking Rate
(millimeters) (meters per day)
Sand grain 1.0 8600.
Copepod (Calanus) 3.0 576.
Silt particle 0.01 14.5
Diatom (Nitzschia) 0.020 0.050
Bacteria 0.001 0.132
Since the density of the water environment is closely similar to that

of organisms living in it, the reader will not be surprised to learn that
slight changes in the

former often have important consequences in
40 The Medium
the vertical distribution of plankton. Differences in the density of
fresh water and sea water at three temperatures within the biological
range are shown in Table 2. As a result of these density differences,
TABLE 2
SAMPLE VALUES FOB THE DENSITY OF WATER AT DIFFERENT TEMPERATURES

Fresh Water Sea Water (salinity 35%c)
Temperature C (g/cc) (g/cc)
0.999 1.0281
4 1.000 1.0278
SO 0.995 1.0217
relatively sharp stratification of water masses may come into being in

lakes and in oceanic areas. If other influences do not interfere,
warmer water tends to remain over colder water, and fresher water
tends to float on top of the more saline.
Murray and Hjort, 1911, copyright Macmiltan & Co.

FIG. 2.8. Calocalanus pavo, a tropical planktonic copepod. The length of the
crustacean's body is about 2.5 mm.
Although the differences in density shown in Table 2 may seem

very small, they are sufficient to influence profoundly the circulation
of water in lakes and in the ocean, and to make the difference be-
tween floating and sinking for planktonic organisms. Plankton some-
times tends to sink through the upper stratum of less dense water,
but to stop sinking when it reaches a stratum of higher density below.
In such a situation plankton and detritus accumulate and form a

"false bottom" that provides a potentially rich feeding zone.
Important changes in viscosity also occur as temperature is al-
tered. At 25C the viscosity of water is only half that at 0C. Since
viscosity is added
at higher temperatures the effect of the decreased
to that of the lower density, the rate of sinking tends to be greater.
Probably as an adaptation to this difference through natural selection

many planktonic organisms in warmer waters are more profusely pro-
vided with bristles and other feathery structures tending to retard
sinking (Fig. 2.8). Certain planktonic diatoms and dinoflagellates

also show a graded difference in the length of spines in tropical and
July Sept. Oct. Dec. Mar. Apr. June July
Redrawn from Wesenberg-Lund, 191'0
FIG. 2.9. Cyclomorphosis in Daphnia cucullata (upper) and Bosmina coregoni

(lower), showing seasonal changes in body form.
polar areas. Whether such geographical variants should be divided

into separate species may not be settled until controlled culturing ex-
periments are undertaken.

In some plankters the body form changes with the season of the
year. This phenomenon, known as cyclornorphosis, is strikingly ex-

hibited by the increased and length of spines
size of the "helmet"
in certain Cladocera during warm months
( Fig. 2.9 ) These changes
.
produce increase in body surface and were originally believed to be
entirely an adaptation to
the lower density and viscosity of warmer
water, but subsequent work has shown that factors other than tem-
perature also influence body form (Brooks, 1946).
In all such re-
lationships the ecologist must distinguish between actual causal fac-

tors and possible beneficial results. For a further discussion of
42 The Medium
plankton in relation to flotation the reader may refer to Sverdrup

et al. (1942) and Welch (1952).
TRANSPORTATION BY MEDIUM
Another significant action of the medium is its action in providing
transportation for plants and animals. Certain requirements such as
lightcan often reach the organism without the movement of either
the organism or the medium. Most necessities, however; are not
adequately provided for unless either the organism or the medium
moves. Put in simplest terms the needs for mobility are: (1) to
provide materials for metabolism and growth; (2) to remove waste
products; (3) to bring male and female elements together; (4) to
distribute progeny; (5) to avoid unfavorable conditions.
Either the organism must be able to forage and to distribute itself
some mobile agent must be available. Fortunately,

effectively, or else
both media are mobile, but their potentialities for providing transpor-
tation differ considerably. Air moves much more readily and much
water can carry heavier objects in suspension.
faster than water, but
The composition of air remains relatively constant, but the water
medium contains widely varying amounts of ecologically important
substances.
Movement of the air medium ranges from local wind currents of
beneficial nature to violent and destructive gales. Animals and
plants in some areas are subject to strong winds which vary in direc-
tion from day to day, whereas in other regions, such as the trade-
wind wind blows almost constantly from one quarter with
belts, the
corresponding unidirectional influence. Air movement also indirectly

affects other factors such as temperature, rainfall, and evaporation.
The circulation of the atmosphere controls the weather and provides
for the transport of moisture from the sea to the land.
Movement of the water medium similarly varies from small-scale,
sporadic circulation, generated by local wind and waves, to larger

and stronger currents. Plants and animals living in ponds and lakes
are regularly subject to wind-driven currents. In fresh-water streams
the one-way transport of water is a matter of vital concern to all the
inhabitants. Along the sea coast, on the other hand, the tidal cur-
rents are characteristically reversible or oscillatory. Farther offshore
the permanent currents are encountered, and these are parts of the
great current systems of the ocean which include a rotary circulation

in each of the ocean basins (Fig. 2.10). Two well-known oceanic
currents of great ecological significance in the northern hemisphere
43
44 The Medium
are the Gulf Stream, which carries water of tropical origin northward
along the east coast of the United States and thence northeastward
toward northern Europe, and the Japanese Current, which brings
relatively warm water to the Aleutians and the Alaskan coast.
Sessile Existence
Since both air and water are practically always on the move, the
possibility exists that animals and plants might live a lazy existence,
remaining in one place and letting the medium bring their needs to
them. The desirable features of such a tranquil life might appeal
to many of us. But since air can carry only the smallest and lightest
particles, no free-living animal on land can obtain sufficient solid food
by air transport. There are no completely sessile animals in the ter-
restrial environment. A few land animals, such as the spider and
the ant lion, lie in wait for the prey which flies or crawls to their
traps, and other forms,

such as the dung beetle and the wood borer,
live within their food material, but even for these animals some loco-
motion is
necessary.
The needs of land plants, on the other hand, are such that most of
them can be brought by mobile agents. The carbon dioxide and
oxygen exchange of the plant is readily taken care of by the movement

of the air. Mineral nutrients needed by the plant are also carried
to it, but in most situations these materials are brought by the soil
water so that we are really dealing with a special case of the mobility
of water. However, certain epiphytes, such as the bromeliads and
Spanish mosses, live in the crotches of trees or even on telephone wires
where absolutely all their needs reach them by means of air transport.
These "air plants" obtain mineral nutrients from rain water and from
dust particles which lodge in their crevices. For many species the
wind plays an essential role also in reproduction and in distribution.
In the water environment the transport by the medium of the needs
of both plants and animals is quite an ordinary occurrence. Most
multicellular plants in aquatic habitats lead a completely sessile exist-
ence, allowing the water to bring them the oxygen, carbon dioxide,
and food materials that they require. The algae absorb their nutrients
directly from the free water, but the
vascular plants generally obtain
these materials through their roots from the water in the mud.
Large numbers of aquatic animals enjoy a sedentary life. Many

groups of sessile animals, such as the sponges, coelenterates, bivalves,
and barnacles, are extremely abundant in the water environment, but
are entirely unrepresented on land. These forms can remain per-
Transport by Air 45
mancntly attached to the bottom, allowing food particles to be brought

to them by the water. Many of these animals are characterized by
radial symmetry; since food may be brought from any direction, a
radial arrangement of
feeding around the animal's mouth
appendages
is efficient. Only when locomotion is required is it particularly de-
sirable to have a head and a tail end with attendant bilateral sym-
metry. Colonial existence is also possible among sessile organisms
but would be very clumsy for forms that require active locomotion.
In aquatic plants and animals the male reproductive cells are carried
to the egg cells by water movement, or both are
discharged into the
surrounding medium where currents bring them together to accom-
plish fertilization. The young of sessile organisms are effectively re-
moved from the neighborhood of the adults during the larval stage.
The planktonic larvae, carried far and wide by the currents, ac-
complish the dispersal of the species and the colonization of new areas.
Many marine and fresh- water animals grow more effectively in a
current than in quiet water, and some forms can live only where the
water is moving rapidly. Certain caddis fly larvae ( Hydropsyche )
livingon stream bottoms construct funnel-shaped nets with openings
upstearm into which food particles drift. The larva of the mayfly
(Chirotenetes) braces itself on the bottom with its head upstream and
spreads hairy prothoracic legs like a net with the result that par-
its
ticles of food in the

flowing water are funneled into the animal's mouth
(Morgan, 1930). The term current demand has been used for the
dependence of certain forms, particularly stream forms, on a move-

ment of the water, but in sqme situations the precise reason that a
current is
necessary is not clear. The larva of the black fly (Sim-
ulium), for example, will not develop in quiet water even though
plenty of and other obvious needs are available.
oxygen
Distribution by Medium
If the needs are not brought to the organism, the organism must
go after them either by its own locomotion or by hooking a ride on
something else. The

transportation service of the environment is of
particular value to the seeds or larvae of sessile organisms and for
those motile organisms that are small or feeble and hence would be
their own steam.
very slow in getting around under
Transport by Air. No organism can live permanently floating in
the air, but this medium is often useful in the sporadic and intermittent
The smallest
transport of terrestrial organisms ( Wolfenbarger, 1946).
forms, such as the bacteria, can be transported effectively by even a
46 The Medium
slight movement of air. ZoBell calculated that if bacteria were re-

leased at a height of 33 m
when the wind velocity was as low as 16
km per hr (10 miles per hr), the microbes would be carried 4800 km
before they reached the ground. The heights to which small or-
ganisms may be carried by turbulence is illustrated by a record of
fungal spores collected on plates exposed from an airplane at an

altitude of 3600 m. The rapid transport by air of spores causing
diseases in both plants and animals represents an ever-present threat
to economically important populations. New strains of wheat rust,
for example, seem to be brought in by the wind almost as fast as
resistant types of wheat are developed.
Many of the earth's most abundant types of higher plants, such as

the conifers,depend largely upon wind
pollination. Pollen grains are
regularly transported hundreds of miles by the wind, and pollen has

been detected in the air over the ocean more than 1000 miles from
land. The
seeds of orchids and of certain other groups are so small
by the wind almost as effectively as pollen grains

that they are carried
and spores. Hairy structures, such as the familiar "parachutes" of
the dandelion, make possible the wind transport of larger seeds over
considerable distances, and wings, bladders, etc., such as those of the
maple and elm, enable many of the heavier seeds of these trees to be
blown far enough from the parent plants to avoid immediate com-
petition (Siggins, 1933). In the grasslands and desert regions yet
another bizarre method for transport is encountered in the tumble-
weed, the spherical upper portion of which breaks off and rolls for
miles before the wind, scattering the seeds as it goes.
Wind also plays a significant role in the distribution of animals, par-
ticularly of insects. The wings of flying insects provide the lift, and
even ordinary winds provide horizontal translocation which may carry
them very great distances (Gislen, 1948). Elton (1939) found cer-
tain aphids and flies alive over Spitzbergen after a wind drift over the
ocean of about 1300 km. Stronger winds carry flightless insects,
spiders, and other small invertebrates in either the active or the
encysted condition. Strong prevailing or seasonal winds in certain

situations tend to exert a regular influence on the distribution of in-
sects. Along the northern shore of the Gulf of Mexico, human in-
habitants are pestered by mosquitoes regularly blown many kilom-
eters inland from the salt marshes by the onshore breeze, and Garrett-
Jones (1950) reported mass wind-borne invasions of areas as much as

47 km from the mosquitoes' breeding places in Egypt. As compared
with oceanic currents, however, winds tend to be irregular, and in-
sects, as well as other flying land animals, can often migrate against
Transport by Air 47
the wind, as has been shown to be true for the monarch butterfly
(Williams et 1942). Consequently, regular transport by the
al,,
medium generally plays a far smaller part in the lives of land animals
than in the lives of the denizens of the sea.
Occasionally the geographical range of an insect pest is extended
by wind action. A classical example of such an occurrence is pre-
sented by the spread of the gypsy moth in New England. This
foreign insect escaped from cages in which experiments were being

conducted in the vicinity of Medfield, Mass., in 1869. Since the
female of the species is one might suppose that the insect
flightless,
would be confined immediate neighborhood of its point of
to the
introduction, or at least would spread very slowly. However, the

species has a special way of "thumbing a ride" on the wind. The
newly hatched caterpillars in their first instar are provided with espe-
cially long hairs. When
they crawl to the tops of trees and spin
long threads, they are soon blown off and are carried considerable
distances before they reach the ground. The caterpillars climb again
to the tree tops and the process is
repeated.By this means, to the
detriment of the oak forests, the gypsy moth was spread throughout
New England within a few years. It even succeeded in
crossing Cape
Cod Bay, a distance of about 40 km.
The lifting power of the wind during hurricanes and tornadoes is
well known and provides an exceptional opportunity for the transport
of larger animals and plants in unexpected directions. Although in
most regions hurricane winds are rare, over the centuries they may
nevertheless have made possible the introduction of new species to
islands and other locations which would ordinarily be inaccessible to
the forms concerned. If the new arrivals become established and if
they are predatory or infectious or if they compete successfully, though

passively, with the native species, they may completely upset
the eco-
logical adjustments of the existing community.

has long been recognized that the fauna of the Greater Antilles in
It
the West
Indies has been derived in the evolutionary sense principally
from the Central American fauna. No convincing evidence has been
brought forward, however, that land bridges ever connected these
islands with the mainland of Central America. How the amphibians,
rodents, snakes, and other species of this general size ever reached the
islands remained a mystery until Darlington (1938) pointed out the
possibility of transport by the hurricanes which so frequently cross

this area.
The exceptional lifting power of

certain atmospheric disturbances
is illustrated by the "rain of fishes" which occurred in 1947 at Marks-
48 The Medium
ville, La.,and is described by an eye witness (Bajkov, 1949). On the
morning of October 23, fish ranging between 5 and 23 cm in length
fell on the streets and in the
yards, mystifying the citizens of the town.
There were areas along the main street in which the abundance of fish
averaged one to every square meter. The fish belonged to the fresh-
water species native to the local ponds. Although no large wind
storm occurred, numerous small "devil duster" tornadoes had been
noticed in the area. At Marksville the majority of fish were dead
when picked up from the ground, but it is perfectly possible that
many of them would have survived had they fallen in water. Stock-
ing of remote ponds by dumping fish from airplanes has often been
successfully accomplished. Transport by local violent air currents
may therefore explain the introduction of fish into land-locked ponds
or lake systems not connected with other fresh-water areas in which
the species occurred.
Transport by Water. Since water has a much greater buoyant

effect than air, transport by the water medium is of vital concern to
many more kinds of plants and animals than transport by air. Trans-
port by currents plays an important role in distribution in both inland

waters and the sea. However, we find the fresh-water organisms do
not utilize this transportation system to nearly as great an extent as
marine organisms. This difference may be partially explained by the
fact that fresh water is less dense and hence less buoyant. Also, in-
land waters are less permanent. Since fresh-water bodies often dry
up, or freeze, their currents cannot always be relied upon for trans-
portation.
Correlated with the foregoing facts, we find that many types of
animals whose marine species have free-living larvae are represented
in fresh water by species with a much shorter larval life. In fresh
water there are many more species in which the eggs or young remain
attached to the adult, as in the copepod, Cyclops, and the crayfish, or
are retained within a brood pouch as in the Cladocera (Fig. 2.11).
If stream animals make use of the water transport system, they must
have some method for getting back upstream again. Some of the
devices serving this purpose are most intriguing. The larvae of
mussels are provided with special hooks with which they attach
themselves to the gills
of fish. Some
of the fish that are thus parasi-
tized eventually wander upstream where the maturing larvae drop off
and metamorphose into adult mussels. Other stream forms resort to
the formation of resistant spores that may be carried by birds or blown
by the wind.
Oysters and other benthonic animals remain established in tidal
riversand estuaries although the planktonic larvae might be expected

to be carried away since the net water movement is always toward
the sea ( Ketchum, 1951 ) In some situations the denser, more saline
.
water near the bottom tends to move predominantly into the estuary
whereas the greater flow in the surface strata is seaward. Observa-
tions in certain estuaries indicate that the older oyster larvae tend to
drop to the bottom on the ebb tide and to rise into the water on the
Single winter egg in ephippium
Redrawn from Needham and Lloyd, 1937

FIG. 2.11. Ceriodaphnia reticulata showing eggs developing in the brood pouch.
flood tide (Carriker, 1951). In this manner a sufficient number of
young oysters work way upstream to repopulate the upper

their
of the In barnacles the problem is solved by the fact
regions estuary.
that, although the younger larvae are found in the upper water layers,
the larvae approaching the setting stage tend to concentrate at deeper
levels where the net drift carries them up the estuary (Bousfield,
1954).
50 The Medium
In the ocean many more groups
are influenced by the transporting
action of the water. This affects not only the varied permanent
planktonic population but also the planktonic larvae of the benthos

and of the nekton. Species having planktonic larvae are much more
numerous in the marine environment than in fresh water. The plank-
tonic life of marine larvae also tends to
be longer, often with many
more stages. Certain euphausiid crustaceans, for example, have five
larval stages including about twenty moults before reaching the adult
condition.
So characteristic are the planktonic forms of many water masses
that they are sometimes used in tracing the currents. Members of the
permanent plankton which live in specific water masses must go
wherever the water goes, like Mary's lamb. Such species are termed
current indicators.Russell (1939) has shown, for example, that the
"Channel water" ordinarily occupying the English Channel is popu-
lated by a plankton community of which the chaetognath, Sagitta
is a characteristic member.
setosa, Occasionally, however, a mass of
"mixed oceanic and coastal water" moves into the Channel from the
region south of Ireland, and its presence is revealed by the abundance
of another species, Sagitta elegans. A
third species, Sagitta serrato-
dentata, is an indicator for the "pure oceanic water" which is found

to the west of the water mass tagged by S. elegans.
The Gulf weed Sargassum, which can be seen by any traveler
crossing the southern part of the North Atlantic ocean, is an indicator
for water of tropical origin. This plant begins life attached to the
bottom around certain of the West Indian Islands (Parr, 1939). It is
torn up by storms and drifts out to sea where it continues to grow
vegetatively for years. Masses of Sargassum clumped together gave

rise in the past to the legend of the Sargasso Sea where ships were
supposed to become hopelessly entangled in the seaweed. You will

remember that, in crossing the Atlantic, Columbus was encouraged
by the presence of drifting seaweed to keep on, believing that he had

seen an indisputable sign of the proximity of land. It is fortunate
that Columbus did not realize that Sargassum may be carried hun-
dreds or even thousands of miles from shore by the currents of the
ocean.
The matter by ocean currents should not be left without
of transport
mention of the classical example of the eel. For generations the
eels came from.
people of Europe wondered where Young elvers
ascended their rivers in the spring, and mature eels left their rivers
in the fall after several years' life in fresh water. No one knew where
or when the eels spawned. Finally, it was discovered that a small,
flat
planktonic organism, long thought to be another species entirely,
was the larva of the eel. The Danish marine biologist, Johannes
Schmidt, then set out to find the spawning ground from which the
larvae came. Little did he know, when he
began, that his search
would require many years of work and thousands of miles of explora-
tion before he answered the
question. By plotting the occurrence of
smaller and smaller larvae, Schmidt (1925) gradually traced the drift
of the larvae back to its
point of origin southeast of Bermuda (Fig.
2.12). Here the eels spawn apparently deep in the water, since no
40*
P
100* 90 80 7Q 60* 60 40 30 20
Redrawn from Schmidt, 1925

FIG. 2.12, Migrations of the eel. Dotted lines indicate drift of larvae from breed-
ing areas of American eel ( A ) and European eel ( E ) . Solid lines indicate return
migration of the adult eels.
one has seea them. The newly hatched larvae drift into the Gulf
Stream and after three years are carried to the European shore,
where they metamorphose into elvers and enter the rivers. The ma-
ture eels make the return trip under their own locomotion, although
how they find their way across two or three thousand miles of ocean
is a complete mystery.
52 The Medium
The American species of eel breeds in an area that overlaps the
spawning zone of the European eel. The Gulf Stream circulation
similarly plays an essential role in carrying its larvae northward.
Nevertheless, no record exists of European eels entering an American

American eels being found in Europe. The metamorphosis
river, or of
of the American eel takes place at the end of one year when it is
opposite the American shore. At this time it is ready to respond to
some influence which orients it toward the mouths of the rivers.
However, its
European cousin, mingled with it in the plankton, is not
yet responsive to shore influences and continues drifting with the Gulf
Stream water until it reaches the European coast. The eventual ar-
rival of the two species of eel on opposite sides of the Atlantic is a
spectacular illustration of the integration of the action of the environ-

ment with genetic differences in the timing of responses.
Harmful Transport
In our discussion of the favorable results of transport by the medium

in some natural situations we must not lose sight of the fact that this
influence, like most others, may also produce harmful effects under
different circumstances. Some terrestrial organisms benefit by the
distribution providedby the wind, but a great many others are carried
into regionswhere they cannot possibly survive. Many land animals
and plants are blown out to sea or over lakes, where they fall into
the water. Insects and birds permanently established in windy
regions have been forced to develop reactions that protect them from
being blown away. The insect fauna of islands and mountain
regions frequently includes a disproportionate number of wingless
species. Although in some circumscribed situations flightless forms

may have survived chiefly from lack of need of flight (Darlington,
1943), it is
probable that certain flying species have been positively
eliminated from many exposed islands by wind action.
Inhabitants of rapid streams are frequently swept down into slug-
gish rivers or into lakes in which conditions are unfavorable for them.
Fresh-water forms are carried into the ocean, where usually they are
quickly killed by the change in salinity. Similar harmful transport
occurs in the marine environment. All the Arctic organisms that are
carried southward by when the water
the Labrador Current are killed
in which they are living is mixed with warmer water in the vicinity
of the Grand Banks. All tropical phytoplankton and zooplankton
which are swept by eddies out of the northern edge of the Gulf Stream
succumb as a result of excessive cold.
Other examples of the destructive action of currents may be on a

smaller scale but of considerable economic importance. In many
coastal regions the planktonic larvae of cod, haddock, and other com-
mercially important fish are normally carried by currents from the

areas where they are spawned to offshore banks suitable for the feed-
ing of the juvenile stages when they take to the bottom. There is
evidence that occasionally unusual eddies carry the young stages
beyond the banks into the open ocean. When the developing fish
are ready for bottom life, they find no suitable bottom withia reach,
and whole populations may perish as a result. A practical applica-
tion for an understanding of these
ecological relations is involved
in the intelligent establishment of hatcheries both on the coast and
inland. Before large amounts of the taxpayer's money are spent,
some assurance should be gained that the young fish released will be
carried by the existing currents to areas suitable for their further
development.
ABRASIVE ACTION OF MEDIUM

The abrasive action of the medium and of the material carried
by it, sometimes referred to as "molar" action, is another aspect of the
ecological influence of the medium. If the medium is air, this action

means the mechanical force of the wind and the grinding action of
sand, dust, snow, and other materials driven by it. In the water, even
stronger abrasive action is produced by waves, currents, and particu-
larly by stones, sand, ice,
and the like, carried by the water.
Even the wind by itself can influence the growth form of plants in
exposed regions. The buttresses at the base of the ceiba trees in the
flatcountry of Cuba have been shown to develop to the greatest extent
in directions tending to support the trunk against the most frequent
winds (Fig. 2.13). In the Texas Panhandle it is said that on a normal
day a man can expectorate a mile and a quarter! The sand carried
by winds in such exposed areas produces an abrasive effect which can
be resisted only by plants with tough cuticle like the cacti and many
grasses. In mountain regions the amount of strong wind to which the
vegetation may be subjected is not always appreciated. For example,

on Mt. only 1920 in high, the weather station recorded
Washington,
a wind more than 120 km per hr (75 miles per hr) on
velocity of
85 days between October, 1940, and March, 1941. The average wind
speed for the period was 61 km per hr, with a maximum of 219 km
per hr. The all-time high for Mt. Washington was reached during
54 The Medium
Photo from J. H.Whh

FIG. 2.13. Buttresses at the base of a ceiba tree near Soledad, Cuba. The great
leverage action of the wind, which the buttresses help to resist, can be imagined
from the height and size of a similar tree seen in the left
background.
the early part of the hurricane of 1938 when the anemometer indicated
a wind velocity of 343 km per hr. The instrument then blew
away.
The continued pressure even moderate winds blowing
of
pre-
dominantly from one direction frequently produces a training action
on the branches of the trees. In addition, branches
whipped about
by the wind on the exposed side of the trees knock off one another's
growing buds. These effects, augmented by excessive evaporation on

the windward
side, often produce extremely asymmetrical growth,
"Flag-form" trees with the upper branches restricted to one side of

the trunk are commonly found on exposed mountain ridges. Groves
of trees in windy lowlands are often similarly wind-trained, the more
protected individuals growing many times higher than those on the
exposed side. The outline of such a grove presents a smoothly rising
contour (Fig. 2.14). Along the sea coast the harmful action of salt
V t
rS yppf*
P^r ,
1
/,'.,
,
>n
t
/^
.
'fpfi^^p^ ''*
'^j^\^fe^t^V':
FIG. 2.14. Live oaks on wind-swept shore at Morehead City, N. C. Wind action
has stunted the trees on the exposed side of the grove at the left.
spray is added to the other effects of the wind in progressively re-
stricting growth
toward the exposed beach and in causing a zonation
of the species present (Costing and Billings, 1942). At high eleva-
tions in the mountains low temperatures act with the severe winds to
the vegetation. The
produce the familiar dwarfing and gnarling of
combined effect of these ecological factors often limits the
growth of
trees to a sprawling mat only a few centimeters high near the summit,
whereas in the protected valleys trees of the same species grow to
heights of over 15 m.
In the water medium abrasive action is an even more serious in-
fluence with which the organisms must contend. Only strongly at-
56 The Medium
tached plants and those animals with streamlined forms and special
hooks, sucking discs, or other devices for clinging to the bottom can
exist in turbulent mountain streams (Nielsen, 1950; Welch, 1952,
Ch. 17). Currents of moderate velocity, such as are encountered in
the lower reaches of streams and along the shores of lakes and of the
ocean, produce a force that must be resisted or avoided by the animals
and plants attempting to maintain a foothold in such habitats. Mov-
to smother the
ing sand and silt continually threaten to abrade or
organisms present. A current with a speed of only 1.4 m
per sec
(2 /2 knots), for example, will move stones and gravel up to 2.5 cm in
l
diameter and thus would grind up all unprotected forms.

At the margin of lakes and particularly on the ocean beach the force
of breaking waves, and of the sand, gravel, or ice carried by the water,
produces an extremely serious abrasive action. Where exposure to
such molar action of waves is particularly severe, the shoreline is often
barren except for especially adapted species. However, in zones of
very considerable wave action, a surprisingly large number of species
of plants and animals have developed methods of maintaining them-
selves.
Devices for withstanding the serious molar action of the water
medium are extremely varied. Anyone who has caught and cleaned a
bass, a scup, or other fish inhabiting turbulent coastal waters knows
how much protection the tough skin and thick scales provide for the
surface of the animal. This resistant integument is in sharp contrast
to the tissue-thin skin of the deep-sea fish. These latter animals living
deep in midocean have no wave action to contend with, nor are they
subject to chafing against rocks or gravel by currents. The inverte-
brates of the shoreline as well as the fish are protected by a resistant
outer surface. The shells of mollusks and of crustaceans inhabiting
such situations are characteristically thick and hard.
During the course of evolution many species in the littoral zone
have developed holdfasts, cementing organs, suckers, etc., which pre-
vent their owners from being washed away. Instead of evolving
with more and more rigid outside surfaces, certain inhabitants of the
surf zone solved the problem in the opposite direction. The sea-
weeds, such as the kelp and the rock weeds, are composed of flexible
tissue with a leathery exterior. These forms give with the surge of
the waves rather than being built to resist their force. The form
and structure of these plants present a sharp contrast to the brittle
nature of the typical land vegetation. Imagine what would
happen
if
your flower garden were suddenly transplanted into the middle of
the breaker zone!
Sessile animals, of which there are many representatives in the tidal
zone, sometimes show a difference in growth form according to
whether the individuals are living in an exposed or a sheltered loca-
tion.
Sponges and tunicates, for example, grow with long pendulous
processes in quiet water, but, if exposed to strong currents and wave
action, individuals of the same species grow closely appressed to the
rocks (Wilson, 1951).
Certain physiological adaptations also exist for life where currents
are strong. Many motile animals exhibit a rheotaxis, that is, an
orientation of their locomotion with
respect to the direction of the
current. The rheotactic reactions of most stream fish is such that
they swim against the current and thus maintain their position in the
stream. Interestingly enough, in some species at least, this reaction
is mediated through the eyes. The fish are stimulated to turn and
swim until the image of the stream bottom no longer moves across
the retina. Other fish are stimulated by differential
pressure or
touch. Many stream animals react in such a way as to move out of
the current and thus reach quiet water or protected eddies.
Other reactions to the current are even more elaborate. Some
caddis-fly larvae, for example, alter the shape of their cases and the
materials of which they are built according to the strength of the
Modified from Dodds and Hisaw, 1925

FIG. 2.15. A series of caddis-fly larvae and their cases from quiet and swift
waters.
58 The Medium
current. Larvae living in quiet water build their cases of large and
irregular material (Fig. 2.15). With increasing strength of current
the cases become more and more streamlined and are built of finer
material. In the swiftest streams they are either completely stream-
lined or else done away with entirely.
The power of autotomy exhibited by some crabs may have arisen
partly in response to the dangers of life in turbulent areas. When

the leg of a crab has been caught or crushed between moving stones
it can be cast off, thus setting the animal free. After autotomy has
taken place, regeneration commences and a new leg grows from the
base of the old leg.
Emphasis has been placed in the preceding paragraphs on the
harmful action of fast-running water and breaking waves. There are
situations, however, in which extreme turbulence may be beneficial.
Some species of coral grow best where the pounding of the surf is
the heaviest, because waves remove sediment which in quiet waters
would accumulate on the coral polyps and tend to smother them.
Thus stimulated, coral colonies of this type rapidly produce "heads"
of coral rock at the exposed outer edge of the reef. Coral heads
reach such a size that they are eventually broken off by the force of
the waves. The form of the reef is the result of a balance between
the increased growth due to the turbulence of the surf and the loss
due to the breaking off of coral material. Here we have another il-
lustration *of the dynamic balance represented by natural communities
and their environments.
3
The Substratum
As a factor of the environment the substratum is only slightly less ele-

mental than the medium. The substratum is the surface upon which
the organism rests or moves, or the solid material within which it
lives in whole or in
part. Some ecologists have not distinguished be-
tween medium and substratum. Another possible source of con-
fusion is the use by bacteriologists of the term "substratum" for the
nutrient medium used for growing microorganisms. The important
point is between the concepts, and it seems most logical
to distinguish
to use "medium"exclusively for the material which immediately sur-
rounds the organism, and "substratum" only for the surfaces or solid
materials of the environment on which or within which the organism
lives.
SIGNIFICANCE OF THE SUBSTRATUM

The substratum is not inevitable as is the case with medium.
Every
organism has a medium, either air or water, but some organisms can
do without a substratum. In the aquatic environment the permanent
plankton and many pelagic fish have no substratum at any time, but
terrestrial organisms must have a substratum for at least part of their
lives since no animal or plant can live permanently suspended in the
air. Another general difference between the medium and the sub-
stratum is that, whereas the medium is rarely changed from air to
water, or vice versa, by the activity of the organism, the substratum
can be profoundly modified by many of the animals and plants which
live on it or in it.
Needs Provided by the Substratum
Fundamental needs of the organism which may be provided by

the substratum are purchase, attachment, shelter, and nourishment.
59
60 The Substratum
The limitation of growth or of distribution by the inadequacy of the
substratum in respect to these needs will be discussed in subsequent
sections of this chapter dealing with the different types of substratum.
The substratum may also have importance in various special ways.
For example, the hue and pattern of many animals protect them from
detection by enemies because they blend with the background ( Cott,
1940). The color and texture of the substratum are thus essential
considerations in the operation of protective coloration in nature.
The larvae of many sessile organisms will not continue their de-
velopment unless they find a suitable substratum. This fact has many
important practical applications as for instance in oyster culture. For
years oystermen have realized that clean, hard surfaces must be avail-
able in the spawning areas if the oyster larvae, or "spat," are to make
a successful "set" each year. To insure the presence of a suitable
substratum the oystermen dump overboard whole boatloads of empty
shells or other material at a time when ecological conditions are such
that the oysters of the region are about to spawn. This specially pro-
vided substratum is known as cultdi, and its presence in sufficient
abundance at the critical moment for the attachment of the larvae
is
necessary for successful oyster culture.
Attainment of the Substratum
The attainment of a proper substratum is crucially important in the

lives of most plants and animals, and special methods meeting this
need have developed during the course of evolution. One obvious

and common procedure is the broadcasting of such great numbers of
seeds, spores, or larvae in the attaching stage that some of them will
eventually "fall on fertile soil/' The majority of terrestrial plants

follow this method, and, although only an extremely small fraction
of the seeds are ordinarily carried to a substratum suitable for growth
to maturity, enough seedlings usually
become established to per-
petuate the species.

The tremendous numbers of animal larvae in the attaching stage
are often an index of the critical nature of the attaching process. A
illustration of the intensity of the reaction for the species to
graphic
attain a proper substratum was found in the study of the settling of
barnacle larvae in experiments conducted in Biscayne Bay, Florida.
Glass plates 20 x 25 cm in size were placed in the water each day, and
the number of barnacle cyprids which attached to them was deter-
mined. On one occasion the count showed that 3860 cyprids had
settled on one glass plate during the previous 24 hours! When it is
Reactions to the Substratum 61
realized that the same intensity of attachment for these fouling or-
ganisms extended throughout great areas of Biscayne Bay, the magni-
tude of the reaction is appreciated.
Reactions to the Substratum
Other animals and plants, rather than

relying on chance to reach a
suitable substratum,
actively seekit.
Climbing plants often exhibit
a tendency for the growing parts to
keep in contact with solid surfaces.
This differential growth in response to contact with a surface is termed
stereotropism. Thus, the tendrils or stems of climbing vines twine
around or adhere to objects with which they come into contact. In
other species, such as euonymus, the vine simply presses against the
solid surface without
any special structure for attachment. Root tips,
on the other hand, turn away from stones and other solid objects
which they may encounter as they grow through the soil. This
reaction might be thought of as a negative stereotropism.
Certain animals exhibit a locomotory orientation to surfaces, a
response known as thigmotaxis, by means of which they keep in con-

This reaction can be observed in
tact with, or avoid, solid objects.
many and
insects,among higher animals it is demonstrated
by rats
and house mice when they tend to keep in contact with a wall
(Fraenkel and Gunn, 1940).
Many worms and insects are stimulated to continue moving about
until their bodies are in contact with surfaces of the environment as
they would be when in burrows or under stones. In the laboratory
insects will come to rest between the surfaces of glass slides, and
this shows that the reaction is a positive response to touch
fact
rather than an avoidance of light. In nature, since there are no
transparent, solid objects, the organism would also be hidden from

view. If a caterpillar is placed on its back, it will immediately go
through righting reactions until its feet are again in contact with the
substratum. However, while the caterpillar is still on its back, if a
leaf is placed in contact with its feet, the animal makes no further
It will be perfectly content to remain upside
attempt to right itself.
down as long as its feet are firmly attached to a solid surface.

A great many animals obviously
exhibit the reverse reaction,
avoiding solid and ordinarily do so by the use of sight or

objects,
touch. Amost unusual method of avoiding collision in the dark is
now known to be employed by bats (Galambos and Griffin, 1942).
At night and in dark caves bats can fly rapidly about without running
into each other or into jutting rocks or other objects. The success
62 The Substratum
with which these animals can navigate in darkness was clearly demon-
strated on one occasion when a small
group undertook the exploration
of a cave inCuba under the guidance of a local plantation supervisor.
No sooner had the party passed through the entrance into the inky
blackness of the cave than hundreds of bats sprang into the air and
flew madly about. The air was thick with bats. At this point the
wife of one member of the group demurred. Not wishing to have the
lady of the party miss the trip, our guide offered her a dollar for each
bat that struck her while passing through the cave. Not a single
dollar had to be paid. Bats have been shown to emit supersonic
vibrations which are reflected from obstacles and are detectible by the
bats' ears. In this unique manner bats can avoid obstructions in their
paths during flight through dark forests and tortuous caves.
The Variety of Substrata
Many different substrata exist in the natural community. Almost

every object is a potential substratum. One often hears the state-
ment, "Nature is relentless." This saying springs from *he fact that, as
soon as any manmade object is abandoned, animals or plants attach
to it, bore into it, or grow over it. Thus nature is relentless in re-
ducing man's artifacts to natural conditions.

The most common substrata are the many derivatives of rock; these
will be considered shortly, but first let us take note of the great variety
of substances which can serve as substrata. It is not even necessary
for the substratum to be a hard surface. The surface film of water
serves as the substratum for a category of organisms called the neuston.
Many algae, certain higher plants such as the duck weed (Lemna)
water striders,and "whirligig beetles" which inhabit ponds are sup-
ported by the tension of surface film and use it as their regular sub-
stratum. Flatworms and pulmonate snails are able to employ the
underside of the water surface as a substratum. Under quiet condi-
tions the observer can see these forms progressing steadily along the
interface between the air and water. Mosquito larvae similarly
can attach to the underside of the surface film (Fig. 3.1). The par-
ticular ecological needs of the mosquito in this stage make it possible
for man
to exterminate the pest, locally at least, by applications of oil
or poison to the surface of the water. When undisturbed by public
health agents, however, the mosquito larva grows rapidly on food
which has accumulated at the water surface. Particles sinking
through the air accumulate on the water and other materials floating
up from deeper layers come to rest under the surface. Hence the
air-water interface presents an excellent
feeding ground. By pinching
the surface film with its
specially adapted mouth parts the larva ob-
tains the food material and causes more distant
particles to move
nearer the area of its In this way the larva is said to be
activity.
able to gather food from a circle
perhaps 30 cm in radius around its
point of attachment.
Anophelines Culicines
Anopheles Aedes Culex
g i
on water)
Water
surface
Fie. 3.1. Diagram of lite cycle of mosquito, showing the use of the water surface
as a substratum in the early stages. The larvae use the surface film not only for
support but also for a feeding ground.
Another special substratum of interest is wood. In the air environ-

ment "dry rot" fungi, termites, and other organisms find this material
a suitable substratum for their activities and for their nourishment.
Far from human habitation the destruction of dead timber by these

aids in the reduction of organic materials
plants and animals merely
64 The Substratum
to simpler substances. But when these species attack buildings, tele-
graph poles, and railroad ties unchecked,
they cause tremendous
damage, often within a short time. A knowledge of their ecological
relations then becomes a matter of vital economic concern. Wooden
structures under water similarly serve as the substratum for other
rotting fungi, for the "shipworm" Teredo, and for other forms, which
are consequently also of great economic importance. It is a curious
fact that, although the shipworms honeycomb a piece of wood, their
tubes rarely run into one another and practically never break through
the surface of the wood to the exterior. In some way the boring
shipworm is able to detect when it is

nearing the limit of its substratum
and stops itself from running out into the open.
Woods Hole Oceanographic Institution
FIG. 3.2. Mussels and other fouling organisms forming a crust 17 to 30 cm thick
and weighing up to 13 kg per 1000 cm 2 on the bottom of a bell buoy seen as it is
being hoisted out of the water off Cape Cod, Massachusetts.
The hulls of ships and the surfaces of underwater structures present

specialized substrata in the marine environment that are used as points
of attachment for many kinds of fouling organisms including
notably
algae, barnacles, mussels, and tubeworms. Even the smooth steel
bottoms of modern vessels and navigation buoys are rapidly attacked
by such plants and animals (Fig. 3.2). An understanding of the
ecology of fouling is
necessary for success in the very practical prob-
lem of preventing this attachment. In experiments with different
types of materials with which to coat submerged objects including
ship bottoms, it was found that no surface could be devised which
was so smooth, so slippery, or so soft that a barnacle cyprid could not
gain a foothold. Tests showed that fouling could be prevented only

by covering the ship's hull with paint that emitted copper or other ions
strongly toxic to the attaching stages, as illustrated in Fig. 3.3 (Woods
C. M. Weiss, Woods Hole Oceanographic Institution
FIG. 3.3. Bottom of boat showing reactions of fouling organisms to different anti-
Fouling has been prevented on port side by emission of ions
from
fouling paints.
cuprous oxide paint. On the starboard side fouling organisms
have attached be-
cause the emission from the metallic copper paint used on that side has been in-
hibited by the coupling action of the galvanized iron patch seen at right of
propeller.
Hole Oceanographic Institution, 1950). Without an effective anti-

is said to require 30 per cent more fuel to
fouling paint a battleship
maintain cruising speed within 6 months after launching, and its top
speed is seriously reduced.
The research program on the ecological
relations of fouling organisms and on poisoning methods undertaken in
American marine laboratories during World War II was reported to
have saved the United States Navy 10 per cent of its entire fuel bill.
The substrata used by plants and animals are not restricted to in-
66 The Substratum
animate objects. The surfaces of other organisms are also susceptible
to invasion by attaching organisms. Plants which grow on the ex-
ternal surfaces of other organisms but do not obtain nourishment
from them are called epiphytes. Similarly, animals which gain attach-
ment or shelter without using the tissues of the host organism as a
source of food are termed epizoans. If the attaching forms obtain
nourishment from the host, they are external parasites. The bro-
meliads and orchids perched on the branches of forest trees in the
tropics are typical epiphytes. The barnacles on the backs of whales
are examples of epizoans, although they are often erroneously referred
to as parasites.
When the substratum of an organism is another animal or plant,
new relations appear. The organism may find that its substratum
moves around, grows, or is destroyed. In such instances the inter-
dependencies between the epiphyte or epizoan and its host become
extremely complex. For example, the distribution of sessile rotifers
attached to the water plant Utricularia depends upon the relative rates
of the migration of the rotifers and the elongation of the stems of the
plant (Edmondson, 1946). Whenthe growth of the plant is rela-
tively slow, the rotifer population tends to become concentrated.

But, in the reverse situation, the rotifers become spread out with cor-
responding changes in
age the distribution of
population.the
Epizoans on the gulf weed Sargassum are similarly subject to the

vicissitudes of a living substratum. The floating gulf weed elongates
at its distal end and dies and breaks off at its basal end. A slow-
growing hydroid attached to the Sargassum might not be able to
maintainitself if its substratum grew out from under it.
ROCK, SAND, AND MUD IN AQUATIC ENVIRONMENT

Influence of the Aquatic Substrata
Although many different materials can serve as substrata in the

water environment, by far the most common are rock and its deriva-
tives. Whether the substratum consists of smooth rock, loose stones,
sand, or mud has a profound effect on the distribution of aquatic or-
ganisms and on the regulation of their growth. Different textures,

various degrees of stability of the material, and a great variation in
the nutrient content have an important selective action.
In shallow water the difference in fauna and flora on a rocky bot-
tom, on a sandy beach, or in soft mud can easily be studied by anyone
visiting the shore.
On the sea coast, for example, a rock substratum
Influence of the Aquatic Substrata 67
will characteristically support a rich growth of brown, green, and red
algae attached by "holdfasts*' and a wide variety of snails, mussels,

sea anemones, starfish, and many other invertebrates secured by suck-
ing or cementing devices. Some of these animals indirectly
derive
nourishment as well as attachment from the rock substratum. The
snails, for example, scrape off and eat the slime that forms on all
underwater surfaces as well as the bacteria and algae contained in the

slime. In shifting sand or gravel few species except rapidly burrow-
ing animals can maintain themselves; but on and in firm sand, espe-
cially when
mixed with mud, a distinctive and rich population of
mollusks, worms, and crustaceans will be found, provided that other
environmental factors are favorable. On a mud bottom where the
in abun-
water is
quieter, rooted plants like the eel grass often grow
dance, and sea cucumbers, brittle stars, sea urchins, and a different
selection of worms are among the common inhabitants. However, if
is formed in the
oxygen has been depleted and hydrogen sulphide
mud habitat, the benthic population will be greatly reduced. Further
Wood, Hoi, Uc<ano grap*<c insn.unon

Photo mad. with Ewin g undersea camera by D. M. Owen,
at a of about 100 m on Georges Bank off
FIG. 3.4. Ripples on the sea floor depth
Massachusetts. The numerous brittle stars and sand dollars visible in the 10 nr
to maintain themselves
on the shifting surface.
area shown are able
68 The Substratum
examples of the control of bottom fauna and flora by the substratum

in coastal water are given by Yonge (1949) and by Pratt (1953), and
in fresh water by Krecker and Lancaster (1933) and by Wilson
(1939).
Farther offshore conditions cannot be so easily observed, but the
modern quantitative dredge and the underwater camera have shown
that the same selective action is
being exerted by the nature of the
bottom Samples dredged from mud and from coarse shell-
material.
gravel at locations only a few miles apart in the English Channel were
shown by Wilson ( 1951 ) to contain strikingly different animal types.
Such differences influence fish populations feeding upon these ben-
thonic species. Studies made with the underwater camera have the
advantage that they reveal the nature of the bottom material and the
organisms living on it in their undisturbed condition. The dimen-
sions of bottom features such as ripples, and the spatial distribution of
the inhabitants may then be examined quantitatively (Fig. 3.4). In
the abyssobenthic zone of the ocean the bottom material usually con-
sists of a soft mud, and here only those animals with long legs, broad
bases, or other special adaptation can move about without being

smothered. At a depth of 1% miles the candid underwater camera
caught a 60-cm "sea cucumber" as it cruised across the muddy bottom
Fie. 3.5. A holothurian ("sea cucumber") moving over the mud at a depth of
2600 m ( 1]X> miles ) on the floor of the open ocean off the coast of New York.
Note the imprint of the double row of tube feet.
Build-Up of the Substratum 69
(Fig. 3.5). These holothurians, and other types of bottom animals,

pass the mud through their intestines and extract the organic matter
from it. For such forms the nutritive value of the substratum may
have an even greater ecological importance than its
physical nature.
Breakdown of the Substratum
We have seen the various ways in which the nature of the sub-
stratum in the aquatic environment limits the growth and distribution
of organisms living on it or in it.
Turning the ecological picture the
other way around, we find that the presence of organisms often has a
profound influence on the substratum. Even solid rock can be bored

into and broken down by animals living on it. In the intertidal zone
of the Oregon coast sea urchins have carved craters for themselves in
the sandstone. So abundant are the sea urchins and their craters that
in many places the shelving rocks present a honeycombed effect.
Even more remarkable is the
ability of bivalve molluscs of the family
Pholadidae to drill into
gneissic rock and into concrete. Some of the
pillars of thecauseway to Key West

have been pockmarked by these
animals. Empty shells which form the major portion of the sub-
stratum in some regions are broken down by the sponge Cliona. The
basal part of this sponge produces an acid secretion that hastens the
disintegration of the shells. In some situations the complete destruc-

tion of an organism's substratum by its own activity has led to its
undoing, for without a suitable substratum it is eliminated from the

habitat.
Build-Up of the Substratum

In addition to the breaking down of solid materials many aquatic
organisms play an important part in building up their substratum.
Calcareous algae and many types of coral animals cause calcium
carbonate to be deposited in and around their tissues. As a result
limestone formations of various sorts are brought into being. Anyone
who has visited the tropical ocean is well aware of the vast extent of
reef -building activity (Fig. 7.12). In fresh water certain organisms
similarly cause
the precipitation of calcium carbonate. The marl
in this way often come to be the chief com-
deposits that are formed
ponent of the substratum in ponds and lakes.
In some situations the new substratum produced by the activity of

consists of the surfaces of the organisms
living agents essentially
themselves. Occasionally a few mussels become attached to individ-
70 The Substratum
ual stones scattered over a clam flat of fine sand or mud. During the
next season young mussels may attach to the shells of the old mussels.
Subsequent "sets" gain foothold on the surfaces of the second genera-

tion until gradually a mat of mussels spreads across the area that had
previously been covered by soft material quite unsuitable for the
growth of a mussel population. Mussel mats of this sort have some-
times been a serious economic concern because they have smothered
valuable populations of clams beneath.
In other situations the action of the living part of the ecological
system is that of binding together loose particles of the bottom material

so that a firm substratum is
provided in place of the shifting sand or
mud previously existing in the area. The eel grass that was formerly
prevalent along our Atlantic coast played this important role in many
Murray and Hjort, 1911, copyright Macmillan & Co.

FIG. 3.6. A washed sample of
pteropod ooze from the Indian Ocean showing the
conical and angular shells of pteropods, the rounded tests of Foraminifera, and the
remains of other types of plankton, enlarged about 10 X .
Influence of the Land Substrata 71
bays and estuaries. After the eel grass was killed by a disease in the
early 1930*s, large amounts of bottom material were washed away from
many sections of the shore. As the unprotected sand and mud was
scoured out by the tides, a great
many other plants and animals that
had been living in the area were destroyed. Eventually the com-
plexion of the whole ecological community became altered ( Stauffer,
1937).
Another way in which organisms can modify the substratum is
through contributing their own remains. The sand on the famous
Pink Beach in Bermuda is composed chiefly of coral fragments. In
the deep sea some of the bottom oozes consist mainly of the skeletons
of planktonic organisms (
Sverdrup, Johnson, and Fleming, 1942, Ch.
20). Globigerina ooze formed by the accumulation of the shells of a
genus of Foraminifera and pteropod ooze similarly composed of the
shells of gastropods are
examples of calcareous oozes (Fig. 3.6). Two
important types of siliceous deposits are formed by the accumulation
of the hard parts of radiolarians and diatoms, Radio-
respectively.
larian ooze is found in certain tropical waters, and diatom ooze is
limited to colder seas.
Aquatic organisms also act reciprocally on their own substratum

by adding organic material to it. In a peat bog the substratum is
practically 100 per cent organic matter resulting from the accumula-
tion of vegetable material (Welch, 1952, Ch. 16). In inland waters
and in the sea, fragments of dead organisms reach the bottom as an
organic detritus. Seaweeds and shore animals that have died and
been broken up by the waves contribute abundantly to this material,
but farther from shore the plant and animal plankton are largely re-
sponsible for the particulate organic material in the water. This
detritus tends to settle out where the current has been sufficiently
reduced. The excreta of worms, lamellibranchs, and various Crus-
tacea have been reported to form as much as 40 per cent of the fine
material in the mud of the Clyde Estuary. All these organic sub-
stances gradually become incorporated into the substratum and serve

as sources of nutriment for the mud-eating benthic animals (Twen-
hofel, 1939).
ROCK, SAND, AND SOIL IN TERRESTRIAL ENVIRONMENT

Influence of the Land Substrata
In the land environment soil is by far the most important substratum,

but rock and such as plants and plant products also
special materials
72 The Substratum
serve as ecologically significant substrata. As rock breaks down it
produces areas of stony ground, gravel, and sand; and with the ad-
mixture of organic matter soil is formed from the parent rock sub-
stance. Each of these materials has
its influence in
controlling the
growth and distributionof plants and animals. As in the aquatic en-
vironment the land substrata provide purchase, shelter, attachment,
and nourishment in varying degree according to circumstances.
Land Surfaces and Animals. Physical differences in the land
surfaces are correlated with special adaptations of animals inhabiting
them. On rocky terrains and in regions with hard, open ground the
running speed of animals is improved by the possession of small re-
sistant feet usually with a reduced number of toes, as in the deer, ante-
lope,and ostrich. Animals living in areas of soft sand, marsh, or

snow are characterized by spread-out feet, like those of the camel, or
of toes that present a large surface, like those of wading birds and the
snow-shoe rabbit. Sand-dwelling lizards and insects similarly are
enabled to move over loose sand by toes or legs widened by lateral
scales or hairs. Other animals have feet especially adapted for climb-
ing trees (squirrels), or for clinging to branches or leaves (tree frogs),
or for dealing with other special substrata.
In contrast to the species requiring rapid locomotion over the land
surface is a
large group of animals that burrow into the substratum.
Many rodents, some birds and reptiles, and a great many insects, as
well as other types of invertebrates, are built for effective digging in
the ground. Certain species such as the mole, the earthworm, and
many spend most of their lives underground. All these bur-

insects
rowing forms are limited to regions in which suitable soil conditions

exist. For example, a tongue of soil running across the Florida pan-
handle that is too dry for the burrowing of crayfish acts as an ecological
barrier separating certain west Florida species of crayfish from species
limited to areas farther to the east (Hobbs, 1942).
The chemical composition of the soil substratum affects animals
both directly and indirectly through their food. Land snails with
calcareous shells are especially abundant on soils rich in lime, but
the abundance of the shell-less slugs is not affected in this way. The
one species of Helix was found to weigh 35 per cent of the total
shell of
weight of the snail in limestone regions but only about 20 per cent of
the total weight in areas with soils
poor in lime (Hesse, Allee, and
Schmidt, 1951, Ch. 20). The bones of mammals likewise are heavier
on limestone soils; this especially true of deer that annually must
is
grow new antlers sometimes weighing as much as 7 kg. It is no acci-

dent that strongly built race horses are raised in the bluegrass pas-
Soil and Its Action on Plants 73
on the limestone soils of
Kentucky. Many herbivorous mam-
tures
mals require an abundant
supply of salt (NaCl) beeause sufficient
sodium must be taken in to maintain a
proper ionic balance with the
large amount of potassium contained in their plant food. If an ade-
quate amount of salt is not available in the halophytes of salt meadows,
ruminants are forced to travel to "salt licks," or are excluded from the
region entirely.
The amount of organic matter in the soil is of vital concern to
earthworms and other small invertebrates that use this material as a
source of food. A great number and
variety of small insects and
spiders, of still smaller nematodes, and of
microscopic Protozoa, which
live
permanently in the soil, also depend upon this organic matter
for their nutrition. Thus the microfauna also is controlled by the
chemical qualities of the soil, as well as by its compactness, dryness,
and other physical characteristics.
Soil and Its Action on Plants. An adequate discussion of the
nature of the time
soil, its
changes and space, and its influence on
in
plants, and indirectly on animals through its effect on vegetation,

would require a whole book in itself. Nothing more than an intro-
duction to the subject can be given here. For a more extensive treat-
ment of soil itself the reader is referred to Lyon, Buckman, and Brady
(1952), Kellogg (1941), and the Yearbooks of Agriculture issued by

the U. S.
Department of Agriculture. Excellent chapters on the
ecological relations of soils in relation to plants are to be found in
Costing (1948) and Daubenmire (1947).

Besides its
ecological importance as a substratum, soil has im-
measurably great economic importance. Fortunately we are rapidly

becoming aware of the critical value of the productive capacity of
soil as a support for civilization. Wolf anger (
1950 ) has stated that
"the soil of a nation is its most valuable material heritage." The cru-
cial need for immediate soil conservation in almost all countries of the
world has been ably pointed out by Osborn (1948), Vogt (1948),
and others. A
thorough understanding of the ecological relationships
involved is essential for the intelligent use of our existing soils, for the
and degradation, and for the restoration
prevention of further
soil loss
of the fertility of worn-out soils.
Soil represents an extremely complex matrix consisting of minerals

derived from the parent rock of the area, organic matter of local origin,
and substances carried in by various agents. The physical nature of
a soil depends first of all upon its texture and structure. Texture is
determined by the size of the constituent particles, and structure is de-
pendent upon the aggregation

of these particles in the undisturbed
74 The Substratum
soil into
grains, clumps,
and flakes. Soil particles are classified ac-
cording to size in the accompanying table.
Sand 1 00-0 05 mm
. . in diameter
Silt 0.05-0.002
Clay < 0.002
In a good loam all three of the categories are well represented.

The type of structure into which the particles are arranged affects
profoundly the porosity of the soil. It also controls the amount of
surface which is presented on the one hand to the air and water
moving through the soil, and on the other to the hairs of roots grow-
ing in the soil. The relative proportions of the soil constituents and
of the air and water present are extremely variable. In an average
good soil about half the volume is
commonly represented by pore
space of which half may be occupied by air and half by water. The
solid material of such a soil may consist of 95 per cent mineral par-
ticles and 5 per cent organic matter. In tropical soils, however,
organic matter may be less than 1 per cent, and in peaty soil it may
approach 100 per cent of the dry material. In addition to differences
in texture and structure, soils vary physically in the type of layering
that they develop as they mature under biological and climatic in-
be discussed in the next section.
fluences as will
The chemical nature of soils is even more diverse and variable.
Upon the disintegration of the parent rocks the whole spectrum of
minerals present becomes available for incorporation into the soil.
Added to these are a wide variety of organic substances derived from
animals and plants and other materials introduced from the air and
ground water. Further chemical changes take place within the soil
as climatic and biological agents work on it. As a consequence, soils
and soil water differ widely in chemical composition, organic content,
and total salinity, as well as in degree of acidity, oxidation-reduction
potential, and other physicochemical characteristics. Some of these
features of the soil are intimately interrelated with the physical char-
acteristics. For example, the smallest particles involved in the texture
of a soil are colloids, and their behavior and reactions are also in-
volved in the chemistry and physical chemistry of the The abun-
soil.
dance and type of the colloids present affect the amount of water
retained by the soil and its availability to plants. At the same time
the colloids influence the chemical composition of the soil water.
In the present section we are concerned with soil primarily in its
physical nature as a substratum for land organisms. The foregoing

brief sketch of soil has indicated the extremely complicated nature
Soil and Its Action on Plants 75
of this substratum and the degree which
to its
physical characteristics
are bound up with chemical and
its
physicochemical features.
its
Certain of the latter will be further discussed in relation to other
ecological factors considered in later chapters.

When gross differences in the land substrata exist, factors limiting
plant growth can frequently be distinguished. On a solid rock sub-
stratum lichens and certain mosses are
characteristically the only
plants that can survive. In situations with a coarse, shifting sub-
stratum, like a sand dune or a gravel slide, the vegetation is limited to
specially adapted forms. Dune grasses with their network of hori-
zontal rhizomes, and certain other
plants, such as Paronychia, with
strong and extensive root systems are among the few plants that can
maintain a foothold. When the soil is
very hard owing to a high silt
and clay content or to the development of a hardpan, the roots of many
species cannot penetrate. At the other extreme the presence of very
soft soils prevents the establishment of
plants that require firm anchor-
age. Many evidences of this relation were seen in New England
after the hurricane of 1938 when whole
groves of trees on loose soil
were uprooted but neighboring groups of the same species on hard
ground remained standing after the storm.
In other situations a gross difference in such factors as the chemical
composition, moisture, or temperature of the soil may be distinguish-

able as the prime influence controlling the vegetation, and examples
of these will be considered later in the appropriate chapters. Too
oft^n for the peace of mind of the ecologist, however, very compli-
cated or subtle differences occur in the soil, and frequently two or
more interdependent factors appear to act mutually in limiting plant
growth. More detailed treatments of the ecology of soils, such as

those referred to earlier in this section, should be consulted for a
further discussion and examples of situations of this type. In some
habitats investigators disagree not only as to which of the soil influ-
ences is determining the composition of the vegetation but

critical in
also even as to whether the climatic factors are not more important
than the edaphic (
soil )
factors.
These ecological relations in the soil also frequently illustrate the

an increase in one factor may some-
principle of partial equivalence:
times partially make up for a deficiency in another factor (Allee et al,
1949, Ch. 16). The lack of moisture in a sandy soil may
be compen-
sated for to some extent by a greater rainfall in certain localities, or,
for which a given region is generally too
conversely, plant species
hui&d may find the effective moisture conditions sufficiently reduced
in a local area with a sandy substratum.
76 The Substratum
Action of Organisms on Soil
Having seen the many ways in which the nature of the land sub-
stratum may influence the lives of organisms, we now may inquire to
what extent the action is reversed. The fact is soon revealed that
animals and plants play a very important part in modifying their sub-
stratum on land just as they do in water. This activity on the part
of terrestrial organisms is particularly striking in relation to the forma-
tion and development of soil. It has been truly said that if it were not
for organisms there would be no soil at least none of biological im-
portance. The soil is an outstanding example of the result of the
organism and the environment acting as a reciprocating system.
Abundance of Organisms in Soil. The great abundance of organ-
isms which live wholly within the soil and the far-reaching extent of
the underground parts of organisms are not always appreciated.
Anyone who spades up a garden or transplants a shrub should be
impressed with the number of roots and the bulk of the root systems
of even small plants. The roots of the typical plant are so finely
divided and subdivided into rootlets and root hairs that a tremendous
surface provided for the exchange between the organism and its
is
surroundings (cf Weaver, 1947). Roots are frequently sufficiently

abundant produce a continuous mat or network extending several
to
feet into the soil. The root system of a maize plant may extend over
1 m laterally and 2% m
deep. The roots of 17-year old apple trees
were found to have occupied all the soil between rows 10 m apart
and to have grown to a depth of 10% m (Weaver and Clements,
1938).
In the animal kingdom the number of species that burrow through
the and thus influence it is also very large. Burrowing rodents
soil
and moles of one kind or another exist almost everywhere, and they
are much more abundant than is generally realized. Although in
some instances the actual number of the larger forms may not be
impressive, their digging activities may be remarkably extensive.
Prairie-dog burrows more than 4 m deep have been reported. In
certain parts of California systematic trapping has shown that as
many as 50 mice inhabit each hectare (2% acres) under normal con-
Periodically, as we shall
ditions. see later, the rodent population
tends to increase greatly in numbers. Even when the population of
mice and other burrowing forms is at low ebb, a considerable in-
fluence on the soil may be produced in the course of a year. In addi-
tion to a variety of mammals, many kinds of reptiles and amphibians
Soil Formation 77
as well as a few species of birds least a part of their lives
spend at
burrowing in the soil.
Of
present in the soil are much
smaller animal forms, the numbers
greater (Chapman, 1931, Ch. 18). Earthworms have been estimated

at hundreds of thousands
per hectare, and their burrows may extend to
depths greater than 2 m. Insects, especially in the larval stages, are
very numerous in the upper centimeters of the soil (Salt et ah, 1948).
In some regions population densities of several million soil insects
per hectare have been found, Spiders, tardigrades, millipedes, and
isopods are also abundant inhabitants of the land substratum. An
extensive study in Illinois showed that the invertebrates of the soil
reached an average summer maximum of 3300 per sq m-or roughly
one animal under every 3 sq cm of surface. Since many of these
forms are short lived and populations succeed one another in the soil,
the study indicated that at least one or two invertebrate animals had
existed during the year under
every square centimeter of soil surface.
In mineral soils of Jutland nematodes have been found to range in
number from 175,000 to 20,000,000 per square meter (Nielsen, 1949).
The numbers of microorganisms in the soil are, of course, much
greater and produce a profound effect on the substratum. Protozoa
may hundreds of thousands per gram of soil,
exist in concentrations of
The mycelia of fungi penetrate the soil wherever suitable conditions

are found, and bacteria are extremely abundant almost everywhere.
In raw humus as many as 20,000 bacteria per gram are a common
occurrence. In rich loam the bacteria population may rise to 50 or
even 100 million cells
per gram of soil (Waksman, 1932). All these
denizens of the soil from sizable burrowing animals to the smallest
microorganisms add their influence to that of the underground parts
of plants in modifying the substratum.
Soil Formation. Soil is formed by the combined action of several
agents. First may be mentioned the process of fragmentation, the

mechanical breakdown of rock material into smaller pieces. The
process is carried forward partly by geological agents, including
especially the freezing
and thawing of moisture in the ground. The
action of roots in splitting
rocks is also important and represents the
biological part of the process.

No matter how small the rock par-
ticles be, however, plants cannot obtain nutriment from the ma-
may
terial until the minerals are rendered soluble. The second step in
soil formation, and one that goes on simultaneously, is termed cor-
rosion and includes the chemical processes of oxidation, reduction,

and others. These processes go
hydration, hydrolysis, carbonation,
forward and soil materials go into solution under the influence of rain
78 The Substratum
after percolates through the ground as soil water. But the action of
it
pure rain water would be extremely slow. Root secretions added to

ground water, including notably carbonic acid, cause the rock mate-
rials to
go into solution much more rapidly. If you look closely at a
rock covered by lichens, you can see the results of the combined action
of mechanical breakdown and corrosion due to plant secretions.
The third factor taking part in the process of soil formation is the
addition of organic matter. Plants contribute their deciduous parts
at regular intervals, and when each plant dies it adds its whole
body to the soil. Since one thinks of plants as growing by withdraw-
ing material from the soil, one might ask how any net gain would
result from the death of the
plant and the return of this material to
the substratum. The answer is that the plant builds additional
materials from the air and the ground water into its tissues. The
carbohydrate synthesized by the plant is formed from carbon dioxide

and water absorbed from the environment. Certain bacteria, such
as Azotobacter and Clostridiwn, living freely in the soil, are able to
take nitrogen from the air and to use it in their constructive growth.
Other nitrogen-fixing bacteria live in nodules on the roots of legu-
minous plants to which they pass on the nitrogenous compounds that
they have manufactured. Thus, more is added to the soil by the
activity of the vegetation than is removed, and, what is more im-
portant, the new material is in a very different chemical form. The
inorganic substances taken up from the soil and from the atmosphere
are converted into complex organic compounds by the growth proc-
esses of the plant. When the plant dies, these organic substances
become incorporated into the soil and many of them decompose only
slowly.
Animals living on and in the soil their excreta regularly and
add
contribute their own bodies to the substratum when they die. Bur-
rowing animals of all sorts mix into the soil the organic remains that
have been added to the top of the ground. Rodents and many kinds
of insects play an important role in this regard, as well as the pro-
verbial earthworm. A visit to a deciduous forest where earthworms
are abundant will provide an opportunity for seeing the effectiveness
of this animal in tilling the soil. If you remove from the surface of
the ground the leaves that accumulated in the last few months, you
find few leaf remnants from previous years. All the older leaves
have been eaten by the earthworms, and their faeces have largely been
discharged at subsurface levels. When constructing new burrows,
earthworms deposit from the deeper levels upon the
their casts of soil
surface. This action of the earthworms in mixing the upper layers in
Humus and the Colloidal Complex 79
the deciduous forest helps to
produce an entirely different soil profile
from that found under coniferous trees. Because of the lack of the
earthworm population in the typical coniferous forest as well as the
slower rate of decay the fallen needles tend to accumulate
year after
year as successive layers on the surface of the ground.
We have seen that living agents aid in the breakdown of the parent
rock material, take part in the vertical
mixing and the horizontal dis-
tribution of soil substances, and add
organic matter to the soil. The
by-products of animals and plants, and their own bodies when they
die, are the only source of organic compounds for the soil. These
organic substances provide necessary soil components, modify the
soil into
many different types, and make possible the growth of a
varied fauna and flora that would not otherwise be able to exist.
The organic matter contributed to the soil contains a larger amount

of energy than the inorganic substances from which they were formed.
Living organisms therefore provide both potential and kinetic energy

as well as materials in helping to build the soil.
Humus and the Colloidal Complex. Soils are far more than piles
of material derived from rocks and biological sources, Soils have
organization. Structure, layering, and other aspects of soil

arrange-
ment are influenced by the animals and plants present, and, recipro-
cally,
the activities of the soil organisms are frequently controlled by
the organization of the soil material. One way in which animal life
and vegetation through their contribution
affect soil organization is
to the colloidal complex. The

partially decomposed organic matter
components is known as humus, and this
added to the soil by living
material combines with the finest clay particles to form the colloidal
complex. Often referred to as the "heart and soul" of the soil, the
colloidal complex plays many essential roles in its
dynamic activity
(Waksman, 1936). In the first place the presence of colloids de-
rived from humus and other sources influences the water-holding
soil and the rates at which air and ground water can
capacity of the
circulate through it. Water tends to move too freely through sandy
soil. The addition of humus to such soil tends to bind the grains to-
gether, to
reduce pore size, and to increase the amount of water held.
A contrasting situation is found
in soil that is too dense because of
an excessive amount of clay. Increasing the amount of humus present

has the effect here of separating the soil material into clumps, and
thus allowing better aeration, increased percolation, and easier root
In both instances the structure of the soil has been im-
penetration.
proved.
The colloidal complex also acts as a source of plant nutrients and
80 The Substratum
as a particularly desirable type of storehouse since materials are re-
leased from it only gradually. As will be discussed more fully in
Chapter the critical plant nutrients of the soil consist of such in-
8,
as
organic materials as nitrate, phosphate, potassium, and calcium,
well as of certain organic substances. These are derived in part from
the humus itself and in
part from the breakdown of the mineral
com-
ponents of the soil.
Many of the nutrient materials are held by the
colloidal complex in loose chemical combination or physical adsorp-
tion on the surfaces of particles. The reactions involved in the de-
composition of soil components and in the association of nutrient mate-
rials with colloids are extremely complicated, and the reader should
turn to a treatment such as Lyon, Buckman, and Brady ( 1952 ) for a
further discussion. The general
point emphasized here is that
the
colloidal the
fulfills function of for the
complex important providing
slow release of nutrient materials in such a way that they can be
absorbed by plant roots as needed.
The between the gradual delivery of nitrogen from the
difference
supply in the organic matter and the rapid exhaustion of soluble nitro-
gen that is freely mobile in the soil has no doubt been observed by the
reader for his own lawn or garden. When nitrogen fertilizers are
added to the soil, they tend to be rapidly dissolved. In this condition
the nutrient salts may be quickly leached away by rain and ground
water, or they may produce a "flash" growth of the plants present.
More desirable for the growth- of cultivated plants, as well as for vege-
tation in general, is the slow availability of nitrogen from the organic
colloidalcomplex and from the decomposing organic matter, although
nitrogen fertilizers are often beneficial if used properly.
The Soil Profile. A broader aspect of the organization of the soil
and another on which animals and plants exert a profound influence

is the soil
profile.
If
you look closely at a fresh vertical section
through the soil, as exposed in an excavation or a road cut, you will
see a succession of layers, or horizons as they are called, that together
form the soil profile (Fig. 3.7). This arrangement of the soil mate-
rial in the result of the action of the living
layers is
components and
the climatic influences of the region on the original parent material.
In some regions this master organization of the land substratum has
been in the process of formation for thousands of The nature
years.
of the soil profile is of crucial concern in respect both to the natural
vegetation and to commercial crops. We

should think
accordingly
twice before allowing agricultural procedures that may permanently
destroy the established layering of the soil.
The depth and composition of each horizon of the soil
profile differ
The Soil Profile 81
Photo U. S. Soil Conservation Service
FIG. 3.7. Vertical cut through fine sandy loam supporting blue-stem grass at Red
Plains Experiment Station, Guthrie, Oklahoma.
A Horizon, 0-1.0 (0-30 cm): grayish brown fine sandy loam.

ft
B Horizon, 1.0-2.5 (30-75 cm): reddish sandy clay.

ft
C Horizon 2.5-3.2 ft ( 75-95 cm ) weathered parent material.

:
D Horizon below 3.2 ft: laminated sandstone and shale.
from place to place and change with time in the same region
greatly
as the soil matures. A varying number of subdivisions of each hori-
zon are recognized with the result that a com-

in different situations
of the soil profile becomes very complex, In many

plete analysis
soils some of the subdivisions are entirely unrepresented, and in other
soils strata of considerable thickness may be uniformly mixed, The

general nature of the principal horizons and subdivisions are shown
diagrammatically in Fig.
3.8.
82 The Substratum
The process involved in producing and maintaining
a typical soil
profile are indicated schematically in Fig. 3.9.

Here is shown one
Horizons
I r i
Litter of undecomposed organic debris

Organic debris partly decomposed or matted
High content of organic matter

mixed with minerals
Zone of maximum leaching
Transitional
True soil
Zone of concentration of transported

fine particles
Weathered or unconsolidated parent material
Rock or unmodified parent material
FIG. 3.8, The soil

profile with its
principal subdivisions shown by a vertical sec-
tion through the soil.
member of a stand of trees that has grown in the area for a long time
and that has produced in the soil the characteristic layering. Per-
haps the most important of the climatic agents is the moisture factor,
and this involves both rainfall and evaporation. Where the land is
sloping a portion of the rain runs off over the surface, sometimes wash-
materials or even some of the soil itself. The rain
ing away organic
that enters the ground percolates through the pore spaces carrying
fine
particles
and dissolved salts with it. In some places the ground
water drains completely through the soil into the strata beneath, with
the result that the dissolved materials may be lost from the soil.
With excessive rainfall valuable constituents may thus be leached
from the upper horizons. Other portions of the ground water enter
the roots of the vegetation and are carried upward again. Water
vapor moves through the pore spaces and evaporates from the sur-
s
1
2
-H ^
rt S
CO
83
84 The Substratum
face. The circulation of soil air provides for the transport of oxygen
and carbon dioxide necessary for or resulting from the metabolism
of the plant roots, soil animals, and microorganisms.
The diagram also indicates the photosynthesis of the vegetation and
the transfer of organic substances formed by the foliage to the roots
deep in the soil. Organic compounds in great variety are elaborated

by the growth of associated plants and animals. As described earlier,
the Deciduous parts of plants, the excrement of animals, and finally
the bodies of all these organisms, when they die, are added to the
soil. These organic remains accumulate on the surface, or are carried
to deeper levels by the soil fauna, where they begin the slow process
of decomposition.
This interplay of biological and climatic agents working on the
parent material results in the production of the soil profile as indi-
cated in the diagram. The A horizon is the scene of the major
biological activity, and here organic matter chiefly accumulates.

Rain entering this layer and percolating through tends to leach out
the soluble salts. The A horizon is
usually darker in color and lighter
in texture than the B horizon below it, from which it is often sharply
distinguished. The B horizon is characterized by less intense biologi-
cal activity. Fine particles tend to accumulate here and mineral

salts are often concentrated in this layer. The material of the B
horizon usually bright in color and densely compact. The B hori-
is
zon tends to grade rather indistinguishably into the C horizon below

it where there is little or no biological activity. As the soil matures,
the A and B horizons tend to deepen until equilibrium conditions are
reached.
Theexact nature of the mature soil profile depends upon the par-
ticular balance reached between the climatic factors, the biological
agents, the contour of the land, and the type of the parent rock in
the given area. The many possible variations and complications
should be borne in mind in a consideration of the simplified ex-
It should be
ample outlined above. amply clear, however, that the
condition of the soil is a result of the elaborate interaction between
organisms and their environment. The dynamic viewpoint of the

modern ecologist therefore, especially appropriate in the further
is,
investigation of the problems of the soil that have been merely touched
upon here. As one further illustration let us consider briefly the in-
teraction between the vegetation and the substratum that underlies
the differentiation of two great categories of soils.
Soils are classified into and
Soil-Group Divisions. groups, in the
United States soil groups fall into two great divisions: the
pedocals
Soil-Group Divisions 85
and the pedalfers, in the terminology used by Wolf anger (1950) and
shown in Fig. 3.10 or the aridic and the humid soils, in the some-
what different classification used by Lyon, Buckman, and Brady
(1952). The pedocal division is composed of incompletely leached
soils found characteristically in the arid Great Plains of the West.
The slight rainfall of these regions does not saturate the soil to a depth
sufficient to reach the water table deep in the ground, and lime tends
to be deposited in the B horizon. When evaporation from the soil
begins, the ground water is drawn upward again
toward the surface,
carrying some of the solubles with it. The water in
this kind of soil
has been said to be "hung from the top, like Monday's wash!" The
roots of the grasses and shrubs that are characteristic of the pedocal
soils absorb the water and the contained salts and also carry them to-
ward the surface. Grasses particularly tend to absorb

a considerable
amount of calcium and to restore this material to the upper layers.
This process aided by the low growth habit of these plants and the
is
great development of rhizomes

and other structures near the surface
PEDALFERS
PEDOCALS PEDALFERS
I
BLACKERTHS GRAY BROWNERTHS
]
DARK-BROWNERTHS RED-AND-YELLOWERTHS
|
BROWNERTHS PRAIRYERTHS
1 GRAYERTHS PODZOLS
[i;,'i,'i;ij|
(JNDIFFERENTIATED HIGHLANDS
| ,]
SANDHILLS
FIG. 3.10. The soil groups of the United States and their division into Pedocals
and Pedalfers. (Modified from Wolf anger, 1950, in Conservation of Natural Re-
sources, reprinted with permission of John Wiley and Sons. )
86 The Substratum
of the ground. For these various reasons calcium tends to accumulate
in the upper layers of the soil, and from this fact the name "ped-o-cal"
is derived. The retention of calcium carbonate and magnesium car-
bonate helps to prevent pedocals from becoming acid.
An
outstanding example of the interaction of climatic, edaphic, and
biological agents in the development of a pedocal soil is furnished by
the chernozems, or blackerths, that occur from the Dakotas south-
ward. The climate is less arid than it is farther west, and a
good
grass cover is typically present. The action of the ground water and
of the
vegetation in retaining salts and other nutrient materials in the
upper layers and in maintaining a neutral or slightly alkaline condi-
tion is indicated
schematically in Fig. 3.11. Organic matter tends to
accumulate in the A horizon, resulting in the
development of the rich,
dark soil condition familiar in the region. Chernozems are conse-
quently especially valuable for grazing or for farming because when
leftundisturbed, or properly managed, the ecological processes pres-
ent tend to perpetuate soil
fertility.
The pedalfer division consists of soil groups found
principally in
the more humid regions of the eastern half of the United States.
Here the rainfall is heavier with usually more than 75 cm falling per
year. The solubles of the soil tend to be carried
beyond the reach of
the roots large amount of water percolating through the upper
by the
horizons. Much of this water filtering through the soil reaches the
water table and drains off valuable nutrients. The roots of the typical
forest vegetation
growing on such soils extract salts but do not remove
relatively as much calcium as do the grasses. The annual leaf-fall is
on the surface, and some of the organic matter
resulting from decay
is carried the run-off, particularly in
away by This loss
hilly regions.
of nutrient materials is in marked contrast to the situation with the
pedocals and the grass vegetation of the prairies.

In the north-central and northeastern
parts of the United States
the increased rainfall and lower
temperatures tend to accentuate the
processes just described. Soils of the podzol group, which occur in
these regions, represent an extreme of the conditions
development
found in the pedalfer division 3.12).
(Fig. In the podzols the
solubles are rapidly lost
by leaching. The A horizon becomes acid
because of the accumulation of leaves, needles, and other
organic
debris on and in the surface strata. Calcium is dissolved from the
upper layers and is often almost completely extracted from the soil.
Aluminum and iron are similarly carried down but tend to be
precipi-
tated again as silicates in the less acid B horizon, and may cause the
formation of hard-pan at this level. The tendency toward relative
accumulation of aluminum and iron generally in this soil-group divi-
sion provides the derivation for the term
"ped-al-fer."
In the region where the pedalfers exist there is generally sufficient
rainfall for agriculture, but the soil tends to lose its nutrients. In the
extreme podzol type the interaction of the climate and vegetation is
such that solubles are rapidly carried away and the soil becomes
progressively poorer and more acid, whereas in the chernozem soil
fertility
tends to be perpetuated. The contrast between the cherno-
COVER OF
GRASSES
AND HERBS
Organic matter
mixed with
mineral soil Soluble
calcium magnesium
nitrogen silica
TRUE SOIL
potassium iron
(Neutral to <
slightly
aluminum etc.
alkaline)
Lime (CaCo 3 )
accumulation
DRY SOIL
MATERIAL
1.5m-
FIG. 3.11. Schematic vertical section through a soil of the chernozem group as
representative of the pedocal soil-group

division, The subdivisions of the soil
profileare indicated in relation to the vegetation and the movement of soil water.
The depth scale is
representative, but varies greatly.
88 The Substratum
zem soil as a representative of the pedocal and the podzol as a rep-
resentative of the pedalfer illustrates the effectiveness with which the
activity of organisms coupled with differences in climate

can modify
the substratum. The soil in its
development and its
organization is
FOREST COVER
Raw litter
A wn
Decomposing <*>
lBr L
litter
Siliceous
mineral soil
TRUE
(Acid
reaction)
Organic material
Compacted aluminum iron
zone ited
WEATHERED
ROCK
Ground water level
(fluctuates) Calcium
2m nitrogen
potassium etc.
eached from soil
W OUftV 7*
FIG. 3.12. Schematic vertical section through a forest soil ofthe podzol group as
representative of the pedalfer soil-group division. The subdivisions of the soil
profile are indicated in relation to the vegetation and the movement of soil water.
The depth scale is representative, but varies greatly.
thus seen to be an
outstanding example of a system formed by the
interaction of organism and environment.
The review of the substratum as an ecological factor given in this
chapter has revealed the great variety of surfaces and solid materials
on and in which animals and plants live. The manner in which the
nature of the substratum controls the distribution and the growth of
different species has been indicated. We have also stressed the fact
that the activities of living organisms and the material of their bodies
after death may profoundly alter the substratum in both the aquatic
and the terrestrial environments. Sometimes this process itself brings
about the further progressive changes in the fauna and flora known as
ecological succession, considered in Chapter 12. Most of all, our
present discussion focuses attention upon the reciprocally dependent
relation existing between the living community and the substratum.
The nature of the mud on the bottom of a pond or of the soil on the
surface of the land is
partly the result of the biological influences and
is at the same time partly responsible for their existence.
The physical conditions of the water or the land originally allow
certain plants and animals to exist in a given area. These living
agents may then modify the substratum, whereupon the substratum
may be further affected by the "climatic" factors, perhaps resulting
in additional changes in the fauna and flora. When we visit the area
after these activities have gone on for a long time, and are still going
on, it is not easy to distinguish cause and effect. Nevertheless, with

a good grasp of the relations summarized in this chapter we are better
equipped to determine the specific action of climatic, physiographic,

and biological influences in bringing about the existing conditions.
4
Water
Water plays several different roles in the ecological relations of plants

and animals. In Chapter 2 we considered the physical characteristics
of water in its mechanical action as a medium. In this chapter we
shall discuss water as a substance taking part in the living complex,
and in subsequent chapters we shall see how water may modify the
action of other factors.
As a material entering the organism water is important as a neces-
sary and abundant constituent of protoplasm, and the plant or animal
body as a whole generally contains a large percentage of water
sometimes 90 per cent or more. Water is essential also as one mate-
rial
taking part in the photosynthetic reaction, through which energy
becomes available either directly or indirectly to all living beings.
Water is necessary as a solvent for food and as an agent for the chem-
ical transformation of the materials within the body. Plants obtain
mineral nutrients from the after they are in aqueous solution.
soil
Although most animals take in their food in solid form, this material
must be dissolved before it can be absorbed by the blood and tissues.
The intestines could be stuffed full of solid food and yet the animal
would starve if no water were available for its digestion. Further-
more, water serves as a vehicle for transport or circulation within the
bodies of organisms. Minerals are carried up the stem of the plant
by the transpiration stream. Water is the principal constituent of
the circulatory, excretory, and reproductive fluids of animals, and it
is
necessary as a transfer agent at respiratory and olfactory surfaces.
Water also acts as a regulator of temperature for plants and animals.
For these essential purposes the proper concentration of water must
be maintained inside the organism, and, at the same time, the transfer
of water must be suitably regulated. If the organism could live
hermetically sealed in a capsule, the retention of the necessary amount

of water would be easy, but exchange of this material between the
90
organism and the outside world is necessary. The crucial need for a
proper water balance is demonstrated by the consequences of even
small water losses in some instances. During starvation, for example,
man may lose as much as 40 per cent of his body weight including
half of the proteins and nearly all the
glycogen and fat without serious
danger, but if 10 per cent of the water content of the human body
is lost, serious disorders result. If as much as 20 per cent of the
water is lost, death follows.
The concentration of water divides the environment into aquatic
and terrestrial habitats. At first sight, one might say that more than
enough water exists in the aquatic environment and that water is a
problem only on land. Closer scrutiny shows, however, that water
is
tending to enter or to leave the organism too fast in almost every
situation. In certain terrestrial habitats the water supply may be
excessive for organisms, as in some tropical rain forests where
many
the air is often 100 per cent saturated and the ground is completely
permeated. A visitor to such a habitat sees moisture condensing on

every surface and hears the steady dripping of water from the
vegetation.
In the ocean in spite of the thousands of cubic miles of sea water
a scarcity of water would nevertheless exist for many plants and
animals! For these organisms the concentration of water is too low
relative tothe abundance of salt for proper osmotic equilibrium.
Fresh-water environments, viewed similarly in relation to osmotic
balance, tend to contain too much water. This contrast between
marine and inland aquatic situations is, in a sense, analogous to the
difference between dry and humid climates on land. The mainte-
nance of the proper water balance is thus a problem that must be
considered in all habitats for all types of organisms.
WATER PROBLEM IN THE AQUATIC ENVIRONMENT

Since the water in natural environments contains a varying amount
of dissolved materials and usually a different amount from the fluids
of the organism, the resulting differences in osmotic pressure raise a
in regard to water exchange. In order to determine the
problem
or to leave the organism we must know
tendency of water to enter
something of the composition of the surrounding medium.
Composition of Natural Waters

for the amounts of the more numerous ions
Representative values
found in sea water and in the water of ponds and lakes are shown in
92 Water
Table 3. A larger amount of each of the common ions occurs in sea
water than in typical fresh water. Hard fresh water contains more
dissolved salts than soft fresh water, particularly with respect to the
calcium and the carbonate ions. A comparison of the relative abun-
dance of ions in each type of water may be made by recalculations in
which sodium is given a value of 100 in each case ( Table 4 ) In sea .
water chloride and sodium are the first and second most abundant
ions, respectively, whereas in typical hard fresh water carbonate is
the most abundant with calcium second. In typical soft fresh water
the calcium and carbonate ions are relatively less concentrated than
sodium and chloride.
TABLE 3
COMPOSITION OF SOME TYPICAL NATURAL WATERS

(After Baldwin, 1948)
Total
Na K Ca Mg Cl S0 4 CO 3 (g/liter)
Soft fresh
water 0.016 0.010 0.019 0.007 0.012 0.065
Hard fresh
water 0.021 0.016 0.065 0.014 0.041 0.025 0.119 0.30
Sea water 10.7 0.39 0.42 1.31 19.3 2.69 0.07334.9
TABLE 4
RELATIVE ABUNDANCE OF IONS IN NATURAL WATERS
Samples of sea water from different localities are surprisingly con-

stant in respect to the relative abundance of the major constituents.
So much is this the case that for many purposes it is not necessary to
measure the concentration of more than one ion. The standard pro-
cedure for determining the salinity of sea water is to find the amount
of chloride ion present by titration and then to calculate the total
salinity and the concentration

of the other ions from the known ratios.
The individual salts occurring in fresh water by contrast, vary widely
often with far-reaching consequences that will be considered later.
At the moment our concern is with the maintenance of the proper
osmotic balance between the inside and the outside of the organism.
Since osmotic pressure is determined by the total concentration of
molecules and ions in solution, values for the aquatic medium will
vary according to the total salt content of the water body (Table 5).
Dissolved materials in soft fresh water vary in amount from prac-
tically zero to about 65 parts per million. Lakes and ponds with
hard water exhibit a variety of higher values for salt content, but
the value given in the table, 300 ppm, is representative. Salt lakes
may contain exceptionally large amounts of dissolved substances.

In contrast to the extreme of inland
variability in the salt content
waters, the open ocean tends to remain highly uniform in salinity
over huge areas (Fig. 4.1), A fish could swim up the middle of the
Atlantic Ocean, for example, from the Cape of Good Hope almost
50
50 60 60 50 40*
FIG. 4.1. The surface salinity of the Atlantic Ocean.

94 Water
to Iceland without encountering a salinity, lower than 35% or higher
than 37%o (parts per thousand). Ocean areas with unusually high
the Red Sea, exhibit
evaporation, like the Mediterranean Sea and
salinities that run up to 40%c. On the other hand, the salinity of the
Baltic Sea is reduced to 8% or less due to the large inflow of fresh
water.
In river estuaries and in coastal areas receiving large amounts of
run-off, salinity may vary over the whole range from nearly fresh
water to full sea water. In smaller habitats, such as rock pools filled
by ocean spray, the salinity may be typical of ordinary sea water on

one occasion, may be greatly increased by evaporation a few days
later, and may be reduced almost to zero by rainfall on another day.
However, for the marine environment as a whole, such situations are

definitely exceptional.
The great length and breadth of the ocean
maintains a nearly constant salinity of about 35 to 36% G .
TABLE 5
TOTAL SALT CONTENT OF SOME TYPICAL NATURAL WATERS

Soft fresh water < 65 .
ppm 065 fa
. 0065
.
%
Hard fresh water 300.0 0.300 0.03
Devils Lake, N. D. 11,000.0 11.0 1.1
Great Salt Lake 170,000.0 170.0 17.0
Baltic Sea 8.0 0.8
Ocean water 35 . 3.5
Red Sea 40.0 4.0
TABLE 6
TYPICAL VALUES FOR OSMOTIC PRESSURE EXPRESSED As FREEZING POINT

DEPRESSION (=A)
AWater AInvertebrates ATeleost Fishes
Mediterranean 2.14 2.2 0.8-1.0
Coastal Atlantic 1.79 1.8 0.7-0.8
Fresh water .03 0.1-0.8 0.5-0.6
The osmotic pressureof a molar solution of a non-electrolyte is 22
atmospheres or 1700 cm of mercury. The freezing point of such a

solution is depressed to 1.84C, and this fact provides a convenient
method for measuring osmotic pressure. In a solution of electrolytes
with the same number of molecules the osmotic pressure would be
higher because of the dissociation. Sea water is a 0.55 molar solu-

tion, and, since the salts are dissociated, sea water has an osmotic
pressure almost equal to that of a molar solution of a non-electrolyte.

Methods of Meeting Osmotic Problems 95
The salt content of the water in alkali soils may be even greater.
Animals and plants living in sea water or in saline soils are therefore
surrounded by liquids having a very high osmotic pressure.
Methods of Meeting Osmotic Problem

If the osmotic pressure inside the organism is the same as that out-
side, relatively little
experienced in adjusting water
difficulty is
equilibrium. the osmotic pressure of the fluids of an animal or

If
plant departs greatly from that of the outside medium, a problem re-
sults in maintaining the
proper water balance. If an animal moves
into an area of very different salinity, a further adjustment will be
required.
Sample values for the depression of the freezing point (Table 6)
reveal the fact that because of the higher salinity the osmotic pressure
of Mediterranean water higher than that of Atlantic water.
is The
latter is
very much higher than a typical value for fresh water. The
osmotic pressure of the internal fluids of plants and of invertebrates is
generally equal or higher than, the medium in which they are

to,
living. Among elasmobranch fishes the osmotic pressure is main-

tained at different levels but is always higher than the surrounding
water due to the retention of urea in the blood.
The osmotic pressure of teleost fishes tends to remain within a

relativelynarrow range for both fresh- water and marine species. In
fresh water the teleost is osmotically superior to its medium, whereas
osmotically inferior in salt water (Fig. 4.2).

it is In the sea a strong
exists for the fish to lose water from its tissues to the sur-
tendency
roundings. A brief calculation will indicate the magnitude of the
osmotic force with which such an animal must contend in order to
maintain its water balance. If the osmotic pressure of the fish is
represented by A = 0.7 and that of the surrounding medium
by A = 1.8, the magnitude of the pressure corresponding to this

difference of A 1.1 is:
X 1700 = 1015 cm Hg
1.84
An osmotic pressure of 1015 cm Hg, or 13 atmospheres, is tending to

extract water from the fish's tissues! For the teleost, therefore, and
for other animals and plants with low internal osmotic pressures sea
water has the being physiologically dry. Fresh-water ani-
effect of
mals and plants that migrate, or are carried, into the sea are indeed
entering an arid climate.
96 Water
The easiest for an organism to deal with the osmotic pressure
way
of its environment is to establish an internal osmotic concentration of
the same magnitude. Even better, if the tissue fluids can be main-
tained at a slightly higher pressure, then there will be a tendency for
the needed water to enter the organism from the surrounding medium.
This is the general method followed by plants and by invertebrate
animals.
A Water
FIG. 4.2. Schematic diagram of osmotic pressure differences between elasmo-
branch and teleost fishes and their fresh-water (left) and marine (right) media.
The degree of shading indicates the relative values of osmotic pressure. ( Modi-
fied from H. W. Smith, 1936).
Most fresh-water and terrestrial plants can maintain an internal

osmotic pressure ranging up to 2 atmospheres but they find saline
habitats with a higher osmotic pressure physiologically too dry and
uninhabitable. However, specially adapted plants, the halophytes,
Limiting Effects of Salinity 97
are able to tolerate higher salinities and to absorb water
by virtue
of the fact that the osmotic
pressure of their tissue fluids is
excep-
tionally high. Halophytes sometimes exhibit internal osmotic pres-
sures as great as 35 or even 40
atmospheres. Such plants inhabit the
margins of the ocean and of salt lakes and can grow in saline soils
with salt contents considerably greater than sea water. Although the
exact relationship is not understood, it is of interest to note that
halophytes, living in a physiologically dry habitat, exhibit succulence,

heavy' cutin, and other xeromorphic characteristics familiar in the
plants of the physically dry desert.
In fresh water the tissue fluids of invertebrate animals have a con-
siderable osmotic superiority because of the typically low salt content
of ponds and lakes (Krogh, 1939). The chief problem here is a
matter of getting rid of the excess water which enters through the
membranes. In marine invertebrates the osmotic pressure of the body
fluids is ordinarily only slightly,
if
any, higher than that of the sur-
rounding sea water. When invertebrates living in estuaries move into
fresher water, they must exert more osmotic work to eliminate the
larger amount of water that tends to enter (Baldwin, 1948). The

teleost fish must actively regulate water transfer both in the sea and
in inland waters to prevent excessive loss of water in the former and
excessive intake of water in the latter. A few animals can tolerate
exceptionally high salinities as is

exemplified by a fish that inhabits
Japanese rock pools with salinity of 60%o and insect larvae that live in
water of 42 to 62%o at Dry Tortugas Island (Pearse, 1950).
Limiting Effects of Salinity
The extent to which the various species can endure changes in

osmotic pressure influences their activity and range. Organisms with
a low tolerance for differences in salinity are known as stenohaline
the open ocean or to
forms. Species that are limited exclusively to
fresh water are thus classed as stenohaline. Others may be limited
to a narrow range of salinity of an intermediate value, but this is
relatively rare. Plants and animals that are able to tolerate a wide
range of salinities are termed euryhaline. Thus organisms inhabiting
an estuary and fish that migrate back and forth from fresh to salt
water are euryhaline. Such fish as the salmon and the eel are able
to regulate their water balance in either a hypertonic or hypotonic
medium.
It should be noted that no sharp line or numerical limit distinguishes
98 Water
stenohaline and euryhaline forms these terms are entirely relative
The terms also do not bring out the important difference in the effect
of time of adaptation. Some species can withstand wide differences
in salinity if
they become adapted slowly, but are unable to tolerate
rapidly changing salinity. Relatively few species can endure both a
wide range and a rapid change in salinity. Interestingly enough,
some euryhaline organisms may grow best in intermediate salinities,
or even require them for a part of their life cycles, although our
knowledge is scanty in this area, The division rate of certain species

of phytoplankton has been shown in laboratory cultures to be twice
as great at a salinity of about 20% than at salinities of about W%o
or 30%o (Braarud, 1951). We
also know that the American oyster
can live in full sea water and can tolerate fresh water for short
periods, but observations indicate that the larvae of this species will
not settle at salinities above 32% nor below 5.6%<j. The optimum
salinity for settlement in this estuarine animal is 16 to I8.6%c. Chang-

ing salinities in estuaries may control the growth, reproduction, and
distribution of more of the inhabitants than we now realize.
Limits of distribution are sometimes sharply determined by salinity
toleration. The vegetation around the margins of salt lakes and in
the spray zone near the ocean shore exhibits a characteristic grada-
from those most tolerant of salt to those least tolerant.
tion of species
In San Francisco Bay the slight difference in tolerance between two
species of pile-boring mollusks was of great economic importance.
In the upper regions of the Bay wharves and other wooden structures
had been built without any special protection against shipwonns be-
cause the local species of the genus Bankia was unable to grow in
salinities below 10%o which frequently occurred in the area. In 1913
another species of wood-boring mollusk, belonging to the genus
Teredo, was introduced into San Francisco Bay, and this species
could tolerate salinities as low as 6% Spreading rapidly, Teredo
.
attacked the wooden structures of the area and within a few years
had caused destruction amounting to more than $25,000,000.
Many interesting problems in relation to salinity tolerance remain

unanswered. In most cases we do not know why one species can
tolerate a rapid or extensivechange medium
in the salt content of its
and another cannot. Why the euryhaline species have not come to
dominate the open ocean as well as the coastal regions is unknown.
Another question of interest is how oceanic birds and mammals can
balance their water budgets without a supply of fresh water. Many
of these forms eat marine invertebrates whose salt content is nearly
Swamps and Temporary Pools 99
as great as an equal volume of sea water. It may be that the water
contained in the tissues of the food and the metabolic water are suffi-
cient for the needs of marine birds and mammals (
Clarke and Bishop,
1948), or these animals may have some special adaptation for excreting
excess salts.
AMPHIBIOUS SITUATIONS
It is difficult
enough for animals and plants to move from the sea to
fresh water or vice versa, but, when
aquatic organisms attempt to in-
vade dry land, they are flopping out of the frying pan into the fire as
far as the water problem is concerned. Nevertheless life on land
does have certain advantages. Plants usually find more light,
better
anchorage, less and more concentrated nutrients.

abrasive action,
Animals may make use of the more abundant vegetation for food
and shelter, the greater oxygen supply, and the possibility of more
rapid movement. Against these advantages there is one signal dis-
advantagethe scarcity of water. Success in colonizing dry land has
been dependent on securing and retaining sufficient water. This
could be easily arranged, perhaps, if the organism could seal itself up,
but, as already stressed, surfaces must be left open for exchange with
the external world. Of immediate importance is the problem of how
to feed and to respire in air without drying up. Representatives of
relatively few phyla have succeeded
in
becoming wholly independent
of the aquatic habitat. However, some plants and animals have come
part way out of the water, or come out for short periods, and we shall
consider these first.
Swamps and Temporary Pools
Certain organisms have hit upon methods of using the advantages

of both the airand the water environment. The emersed vegetation,
for example, which grows in swamps or on the margins of lakes has
its roots in the water and its
upper portions in the air. In this position
the roots find plenty of water available and extract nutrients from the
mud, while the leaves are in the best position for receiving light from
the sun and carbon dioxide from the atmosphere. However, many
in a partially submerged condition be-
plants are not able to survive
cause of inadequate direct supply of oxygen for the roots. Plants
that are adapted to existence entirely under water, or with their roots
in water or in saturated soil, are known as hydrophytes.
100 Water
Amphibians are similarly able to utilize both air and water environ-
ments. species can leave the water to forage on land, and,
if
Many
their pools dry up, they can migrate to other bodies of water. Such
animals are not limited by land barriers as most fish are. Yet am-
phibians must re-enter the water or visit damp places at intervals to
keep and most species need water for reproduction.
their skins moist,
Organisms living in swamps and pools must be able to deal with

periods of drought when they occur. Physiological adaptations mak-
ing this possible have been evolved sometimes without any accom-
panying special morphological structures. Certain species of fish in
India can live in wet grass for as long as 60 hours, and other animals
burrow in the mud where they remain in a dormant condition during
the dry period. Water mites have been shown to be able to live
under debris for intervals up to 6 months after their pools have dried.
The African lung fish, on the other hand, constructs a special mud
cocoon and curls up inside for the duration of the dry spell. The
fish secretes about its body an impervious sheath which prevents
the loss of water from its tissues, and is thus enabled to survive drought
for more than two years. Other specialized structures for tiding over
dry periods are the spores and seeds of water plants, and the "resistant"
eggs and cysts of various aquatic animals ranging from the Protozoa
to the Crustacea.
Another adaptation for meeting the water problem in temporary
pools is that of speeding up development and taking quick advantage
of water when it is available. In certain animals living in this type
of habitat the larval stage is accelerated; in others parthenogenesis
has been adopted. If a single female cladoceran such as the water
Daphnia, finds
flea, a temporary pool, it does not have to hunt
itself in
around for a male. can go ahead and reproduce rapidly by par-

It
thenogenesis. The young animals mature and continue reproducing

parthenogenetically until in a relatively short period a large population
has built up. Later, however, if the pool tends to dry up, the adverse
conditions cause the appearance of males in the population. The
females now produce a different type of egg, one which requires
fertilization. Only one of these "winter" eggs is formed by each
female and the brood pouch is modified into an ephippium (Fig.
2-11). After the death and disintegration of the female, the ephip-
pium remains as a resistant egg case within which the egg can survive
prolonged periods of drought and even freezing. In this condition
the egg may be blown into another pond or carried in dried mud by
animals to another region. When water is again supplied to it, the
egg hatches out and is able to start the cycle over again.
Tidal Zone 101
Tidal Zone
Another situation that might be classified as amphibious is the

Plants and animals
living on the seashore between
tidal zone. tide
marks are subject to alternating droughts and floods twice each day.
Organisms living in the center of the tidal zone must be able to be
aquatic for about hours when the tide is in, terrestrial for about
6%
6% hours when the tide has ebbed, and so on, day in and day out.
The tidal zone is thus a
very difficult habitat, and also a very compli-
cated one because of differences in the timing and height of tide.
Since low tide and high tide occur about 1 hour later each day, the
intertidal area comes to be exposed at all hours of the day and night.
During the first and last quarters of the moon neap tides occur, i.e.,
tides of low amplitude, but during periods of new moon and full moon
the tides rise higher and fall lower and are known as spring tides. In
some such as the west coast of the United States, one set of
localities,
high and low tides each day has a much greater amplitude than the
other set. The height of the tide differs greatly from locality to lo-
cality,and the duration and depth of inundation varies according to

the level within the tidal zone at which a given organism exists. For
example, an animal or a plant attached near the upper extreme limit of

the tide will be covered by sea water for only a brief period about
every two weeks, whereas an individual living near the lower extreme
tidal limit will receive only a correspondingly short exposure to the air.
The tidal zone comprises the upper portion of the littoral zone, as
indicated in Fig. 2.4.
Despite these severe conditions a dense population of a very wide

variety of plants and animals characteristically inhabits the tidal
zone. The fauna and flora consist chiefly of sessile forms, such as the
common seaweeds, barnacles, mussels, and clams. Those plants and
animals that cannot move out of the tidal area must endure the full
effect of the differences in the two media. In addition, certain active
forms forage here under favorable conditions and then retreat. At
low tide animals come from the land and feed on the material that
has been exposed. Shore birds are familiar in this area, but less
observed are rats, mice, mink, skunks, and other small mammals that
feed chiefly at night. During high tide the reverse situation obtains.
Fishes, crabs, and other active animals from deeper water move in for
a few hours and then retire. The alternating visitations of these forms
remind one of Box and Cox in the famous operetta.
It is of interest to inquire why the tidal zone should be so densely
102 Water
populated since the habitat presents such great difficulties for exist-
ence. In the first place all species tend to spread; marine animals
and plants are pressing on their boundaries, and terrestrial forms arc
likewise attempting to invade regions at the margin of their habitats.
These species extend their ranges just as far as
they can endure the
conditions. Furthermore, the tidal zone has certain advantages over
conditions in deep water. Shallow water has more light, oxygen, and
and certain predators are excluded.
circulation; A great amount of or-
ganic matter occurs in this zone, and much of this forms a valuable
food for the inhabitants. A large part of the organic material exists
as detritus resulting from the breakdown of seaweeds and the dis-
integration products of dead animals. In addition many kinds of

smaller planktonic organisms abound in the shallow water and add
to the food supply. The adaptations of the intertidal organisms and
the control of their activities by the environment will be discussed
briefly here;more comprehensive treatment of this special habitat will

be found in such books as Yonge (1949) and Wilson (1951).
Practically all the permanent inhabitants of the tidal zone are

aquatic organisms, and accordingly their chief problem is dealing with
the adverse conditions of the air medium when the tide is out. Many
physiological and morphological adaptations are displayed for with-
standing the recurring periods of water shortage. Many of the
simpler organisms can endure a considerable drying of their tissues

without permanent injury. Sea anemones, for example, have been
kept out of water 18 days; at the end of this period the animals looked
like dried raisins but became active again when replaced in sea water.
Attached forms with shells like the barnacles, mussels, and oysters
simply close up shop during the period of low tide, and many can
remain sealed up in this way for unbelievably long periods without
injury. If
kept at a low temperature oysters may be stored out of
water for 3 months or more and will resume normal activity when
replaced in the sea. Snails and starfish move into sheltered crevices
when the tide is out; worms, clams, and other burrowing forms dig
deeper in the mud and wait for the tide to return.
Certain intertidal forms seem to benefit by periods out of water. A
practical application of this fact is made use of by many oyster

growers who prefer to raise oysters in situations where they are ex-
posed to the air for at least a portion of each day. The oystermen
believe that as a result the muscles which close the shells are strength-
ened. When the oysters are shipped to market, the shells are held
more tightly together
thus retaining the fluids within the mantle
Tidal Zone 103
cavity. These oysters are said to be better able to "hold their
liquor!"
Actively swimming or crawling animals may be able to go without
food for the period when the tide is out, but they cannot remain
active and do without a
supply of oxygen for this length of time.
There is no lack of oxygen in the air, of course the problem is to
get it without allowing the respiratory membranes to dry out. The
gills
of many of these animals are placed in partially enclosed cavities
of the body where oxygen may reach them but the loss of water from
the respiratory surfaces is retarded. A

striking gradation in the
anatomical adaptations of crabs in the tidal zone has been described
by Pearse (1950) for the North Carolina shore. He called attention

to the fact that species living higher up on the shoreline show a re-
duction in the number of their gills and particularly in the ratio of
gill
surface to the size of the whole animal (Fig. 4.3).
Pearse, 1936
FIG. 4.3. Reduction in number of gills

and increase in ratio of body to gill
volume in species of crabs living at progressively higher levels in the littoral zone.
Differences in ability to withstand the progressively more difficult

conditions from the low-tide mark to the high-tide mark have resulted
in a zonation of the plant and animal species. On rocky shores this
zonation is often so pronounced that it can be seen at a distance when
104 Water
the tide is In the situation shown in Fig. 4.4 the sharp upper
out.
limit for thegrowth of the rock weeds is easily apparent. At a higher
level the longer period of air exposure and of lack of sea water im-
posed a clearly defined upper boundary upon the barnacle population.
Photo by R. W. Miner
FIG. 4.4. Rocky shore at low tide showing stratum of sea weeds (Laminarian
Zone) and of barnacles (Balanoid Zone).
Stephenson and Stephenson (1949) have found that on rocky shores

in many parts of the world three fundamental zones may be recog-
nized: the Littorina Zone, characterized by snails; the Balanoid Zone,
characterized by barnacles; and the Laminarian Zone, characterized
by laminarian seaweeds. The relation of these zones to the tide marks

is shown in the
diagram.
High Spring Tide
TIDAL ZONE
Low Spring Tide

Tidal Zone 105
The upper part of the Littorina Zone extends into the spray zone, or
supralittoral zone, where the salt-resistant outposts of the land fauna
and flora are to be found as well as those marine organisms most
capable of enduring desiccation (Boyce, 1954). The Laminarian
Zone extends for varying distances below low-tide mark.
v,-:^Wp:$^^ %%:*&
^&^'&
'i$fcM?^^^
y/vv
Aftr :'Jv ,>':'? *,**,*i*tY
;
oiA**,*t* *%** <r ".. >\H-
Stephenson and Stephenson, 1949, copyright Cambridge University Press

FIG. 4.5. Diagram to illustrate the three fundamental /ones of lile between tide
marks on a rocky shore. Balanoid showing'/one (A) barnacles only, (B)
barnacles inhibited by sweep of seaweeds, and (C) alâl subdivisions typically
encountered on British coasts.
the main
Frequently various subdivisions are superimposed on
zonation of rocky shores (Fig. 4.5). The vertical limits of the various
subzones of fucoid algae and of associated invertebrates are deter-
mined not only by relative tolerance in respect to the water factor
but also by type of rock, wave action, and relative success in invasion,
competition, and resistance
to predation.
106 Water
On muddy and sandy shores zonation of plants and animals be-
tween tide marks similarly occurs, but the observer will have to dig
to ascertain the full ramifications of the tidal
relationships since many
of the inhabitants are burrowers. In addition, microscopic examina-
tion of samples of bottom material reveal a zonation of the microor-
ganisms in the tidal zone. The distribution of copepods dwelling in a

sandy beach of Cape Cod will serve as an illustration ( Fig. 4.6 ) . The
Paraleptastacus
brevicaudatus
Psammoleptastacus
arenaridus
-1.5 1.5 3.0 4.5 6.0 7.5 10.5
Horizontal distance above low-tide mark, m

Pennak, 1939
FIG, 4.6. Abundance and distribution of sand-dwelling copepods across the tidal
zone at Nobska Beach, Woods Hole.
different species present were so nearly alike in general appearance

that they could be
distinguished only by careful study. Neverthe-
less, one species was found to live only below low tide where it is never
subjected to the drying action of air. Another form was most abun-
dant midway between the tide marks. A
third species was most
numerous just below the high-tide line. Furthermore, when samples
of sand from different levels beneath the surface were studied, the
numerical abundance of the copepods was found to vary with depth
in the sand as well as with position across the tidal zone. At the low-
tide mark the largest number of animals was found in the upper 4 cm,
whereas at the high-tide mark the largest population occurred at a
depth of 12 to 16 cm. In the tidal zone, as in other amphibious situa-
Water Problem in Terrestrial Environment 107
tions, the water factor thus exerts a major controlling influence on

the minute organisms as well as on the more conspicuous species.
WATER PROBLEM IN THE TERRESTRIAL ENVIRONMENT

When we turn to dry land as an environment, the water problem
becomes extremely acute. Everyone is familiar with the quick death
which awaits aquatic organisms brought out on land. Many terres-
trial forms can endure the land environment only if they remain in
damp places. This is true for a great many species of plants and
also for animals without special protection. A
salamander that has
escaped from its terrarium and run across the
dry floor will soon be
dead if it is not rescued and put back in a humid atmosphere.
Earthworms frequently crawl out on a concrete sidewalk during a
spring rain. If the sun comes out and dries the sidewalk, many of
the animals will be killed before they can find their way back to the
moist ground.
The successful invasion of the dry land is dependent upon the
possession of methods for securing and retaining sufficient water and

at the same time allowing adequate exchange with the environment.
Terrestrial plants and animals as well as aquatic forms must maintain
a proper water balance between water income and loss. Great varia-
tion exists in the moisture content of land organisms and also in the
amount of water intake and output that is necessary to provide for
metabolic processes. Insects subsisting on dry food probably hold
the record for the minimum water exchange. At the other end of the
scale are many types of higher plants that are extravagant water
users. Some species of plants absorb and transpire more than 2000
grams of water for every gram of dry matter produced by assimila-
tion. An oak tree may transpire as much as 570 liters of water in a
single day.
How many taxonomic groups have succeeded in colonizing dry
land? Actually, only a very few of the prominent kinds of plants and
animals have accomplished this. In the plant kingdom it is chiefly
the vascular forms that are able to live in really dry places, but some
low-growing bryophytes, algae, and fungiparticularly the rock lich-

ensform noteworthy additions to the list.
Among the vertebrates,
only the reptiles, birds, and mammals can live

freely exposed to dry
air.
Although several phyla of invertebrates are reported in the land
fauna, only the insects, spiders, and snails occur in important numbers
in dry habitats. By contrast, there are a great many plant and ani-
mal groups that are exclusively aquatic.
108 Water
Occurrence of Water in Land Environment 109
Occurrence of Water in Land Environment

The onlyultimate source of water for the terrestrial environment
is condensation chiefly in the form of rain. If precipitation were
evenly distributed, it would cover the earth to a uniform depth of
1 m
each year. Quite the contrary is the case. A glance at Fig. 4.7
will reveal the irregularity in the
geographical distribution of rainfall.
In the desert areas the rainfall is less than 25 cm
per year and is in-
adequate for most organisms. In the black areas of the chart an
excessive rainfall of 200 cm or more is reported each year. The mean
annual rainfall on the south side of the Himalayas is recorded as 1232
cm. At the other extreme is Iquique, Chile, with an average precipi-
tation of 0.125 cm
per year. Among the Olympic Mountains in the
state of Washington the annual rainfall totals as much as 380 cm on
thewindward side of some ridges and as little as 25 cm on the lee-
ward side.
Seasonal variations in the distribution of rain may be of even
greater importance than the geographical aspect. In many regions
the division of the year into a rainy season and a dry season is of more
ecological significance than the change in temperature between sum-

mer and winter. In the climatographs shown in Fig. 4.8 two situations
are contrasted. In the Chicago region representing a temperate cli-
mate, an average of 5 to 9 cm of rain falls in every month of the year,
but the average monthly temperature varies from about in 4C
the winter to about 22C in the summer. In the tropical climate of
Barro Colorado Island the average monthly temperature changes by
not more than 1 or 2C
throughout the year, whereas the precipitation
drops to less than 3 cm per month in the dry season and rises to more
than 40 cm per month in the wet season.
Wherever the amount of precipitation varies greatly during the
course of the year, the time of occurrence of rain in relation to the
temperature cycle has great effect on the vegetation. Only those

species will thrive whose varying needs for water in the different life
of available moisture.
stages are satisfied by the seasonal distribution
In manyof the grass-covered or forested areas of the temperate zone
the major portion of rain occurs in the summer, but in other regions,
of which southern California is an
example, most of the precipitation
occurs in the winter. In the latter situation we find a special type
of vegetation with broad evergreen leaves known as the sclerophyllous
forest. In the prairie provinces of Canada the average annual precipi-
tation of 50 cm would not be sufficient to support agriculture if the
110 Water
rain were evenly distributed through the year. However, the major
portion of the rainfall occurs during the spring growing season, and
also at a time before the rate of evaporation has become excessive,
with the result that the available precipitation is used to best advan-
tage. The existing combination of ecological conditions thus permits
western Canada to be an important wheat-growing region.
The foregoing discussion brings into relief the fact that in consider-
ing moisture conditions on land the actual amount of the precipita-

tion is only part of the story. The other part, and often the more
5.1 10.2 15.2 20.3 25.4 30.5 35.6 40.6 cm

C
-26.7
1 23456 7 8 9 101112 1314 15 16 17 in.
Average monthly rainfall
FIG. 4.8. Average rainfall and temperature for each month of the year as indi-
cated by the numerals.
Lower temperate climate data for Chicago, Illinois.

figure:
Upper figure: tropical climatedata for Barro Colorado Island,
Panama Canal Zone. (Reprinted with permis-
and Schmidt, Ecological
sion from Hesse, Allee,
Animal Geography, 1951, John Wiley and Sons,
New York.)
Moisture in the Soil 111
important part, is the loss of water. Of the

upon the
rain that falls
surface of the soil, and another
part runs off immediately, part sinks in,
portion is lost by evaporation. The amount of moisture at any one

time and place depends upon the relative rates of the supply of water
to the soil and evaporation from itanother
example of an ecological
factor whose value depends upon an equilibrium. In evaluating the
water factor in the terrestrial environment both supply and loss
processes must be taken into account.
Moisture in the Soil. Since the terrestrial environment is so varied,
it not surprising that the water factor is very complex, and the
is
moisture in the soil and in the air will first be considered separately.
When rain water enters the soil, it fills the spaces between the par-
ticles. The volume that can be filled is known as the pore space and
commonly varies between 60 per cent for heavy soils and 40 per cent
for light soils. When the rain ceases, a certain portion of the water
in the soil soon drains out, and this is known as the gravitational water.
The portion of the soil water that is held by capillary forces around
and between the particles is the capillary water. That part of the
pore space which is not filled by gravitational or capillary water may
be occupied by water vapor. Another form of soil water, termed the
hijgroscopic water, occurs as an extremely thin film on the soil grains
but cannot move as a liquid. A small portion of the soil water is
this
chemically bound with soil materials and is known as combined water.

The total amount of capillary, hygroscopic, and combined water plus
the water vapor constitutes the field capacity and is the maximum
amount of water that the soil can hold after the gravitational water
has drained away (Fig. 4.9).
The maximum potential supply of water available for the vegetation
is
represented by the full field
capacity of the soilf As plants draw
on the soil water, they gradually reduce the amount held in capillary
spaces. Below a certain moisture content plants tend to wilt in the
hottest part of the day. When the capillary water has been still
further depleted, a point reached at which the plants will not re-
is
cover from wilting until more water is added to the soil, regardless
of other environmental conditions. At this point absorption of water
by the plants has become too slow to replace the water lost by
transpiration. The moisture then remaining in the soil is designated
as the permanent wilting percentage, or the Since
wilting^coefficient. _
little difference has been found in the abundance oi' soil water when
various species growing in the same soil,

wilting occurs for plants of
the permanent wilting percentage is primarily a characteristic of the
soil, and as such has great ecological significance. The amount of
112 Water
water between the full field
capacity, as a maximum, and the wilting
coefficient, as a minimum, represents the available water. As indi-
cated in Fig. 4.9, a considerable amount of water may still remain in
the soil after the wilting coefficient has been reached. A fraction of
the capillary water, the hygroscopic water, and the combined water,
as well as the water vapor, cannot be obtained by the plant, and these
constitute the non-available water.
SOIL WATER
Gravitational
Pore space-
Available
Capillary
Field capacity
Wilting coefficient
Hygroscopic Non-available
Combined
Fig. 4,9. Generalized diagram of the forms of soil water. The relative propor-
tions and the value of the wilting coefficient differ according to the nature of
the soil.
f;
Soils vary considerably in the relative proportions of the different

categories of soil water. In some instances the amount of non-avail-
able water may actually be larger than the available water. The
fact that a portion of the water in the soil is unavailable to land or-
ganisms and that water in the sea is unavailable to aquatic organisms

not adapted to its osmotic pressure contributes further to the general
concept of the universality of the water problem. From the ecologi-
cal point of view the actual amount of water present in any habitat is
not as^ important as its availability.

Moisture in the Air. For many terrestrial organisms the moisture
in the soil is chiefly important as constituting the principal source of
water, whereas the moisture in the air is mainly significant as con-

trolling the loss of
water. For this reason the absolute humidity, or
total amount of water hi the air. is

generally of far less ecological
consequence, than the relative humidity. The ecologist therefore
focuses his attention on the relative humidity or the amount of mois-
ture in the air as a percentage of the amount which the air could hold
at saturation at the existing temperature. Since the capacity of air
for water vapor increases with temperature, the relative humidity of
the atmosphere is reduced in any situation in which an increase of
temperature occurs without an accompanying increase in the total

moisture content of the air.
In the terrestrial environment the ecological effect of the water

factor is consequently strongly influenced by the temperature factor.
Of two regions having the same rainfall the warmer is the drier in
the ecological sense (Fig. 4.10). The climate of a locality with a
10 16 21 27
in. cm
0F 10 20 30 40 50 60 70 80
Mean annual temperature
Modified from McDougatt, 1925
FIG. 4.10. Schematic representation of the influence of rainfall and temperature
on climate.
114 Water
mean annual precipitation of 50 cm would be characterized as humid
if the mean annual
temperature were 7C or less. On the other
hand, another locality with the same rainfall would be regarded as
having a semiarid climate if the annual temperature were 21C or
more. The contrast in climate between the Canadian prairie and
the Mexican desert, both with about 50 cm of rain but with very
different temperature conditions, is an excellent illustration of this
principle.
The geographical
variation in relative humidity is very great. Rela-
tive humidities of 80 to 100 per cent characterize the tropical rain
forest. Regions reporting values of less than 50 per cent are regarded
as
having dry climates, and those with values of less than 20 per cent
arc extremely arid. It is of interest to note in
passing that in cold
winter weather the relative humidity inside our houses is rarely higher
than 35 per cent.
At any one locality the relative humidity may remain relatively con-
stant for long periods of time or may vary widely. On many oceanic
islands the humidity is very nearly the same throughout the year.
In other localities, characterized by wet and dry seasons, the humidity
fluctuates widely from one part of the year to another. In certain
situations as on the plains and in desert regions considerable changes
in moisture content may occur during the course of each day. Records
made in a short-grass prairie in the United States showed a variation
from a relative humidity of less than 30 per cent in the early after-
noon to more than 95 per cent in the middle of the night (Fig. 4.11).
Obviously animals and plants living in such habitats must be equipped

to withstand these rapid and extensive changes in the water factor,
and their lives must be attuned to them. In deserts where daytime
humidities are extremely low and evaporation is excessive, the greater
relative humidity at night helps to relieve the critical moisture con-
dition. Many desert animals take advantage of this situation by
going abroad only during hours of darkness; in some desert plants the
stomata open only at night when transpiration loss is at a minimum.
The foregoing has been a brief description of the amount of moisture
in the air in terms of its relative humidity. As already implied, the
chief ecological significance of the relative humidity is its effect on
the rate of water loss. Terrestrial animals and plants lose water
directly to the air by evaporation and transpiration. The water

supply in their substratum is also reduced by direct evaporation from
the soil and indirectly by the transpiration of the vegetation. Evap-
oration takes place very rapidly from soil because of the great surface
area presented by the fine particles and causes the drying of the upper
layers. Water is lost from the deeper layers chiefly by absorption by

roots and attendant transpiration.
The magnitude of water loss by transpiration is often not appre-
ciated. Perhaps some idea can be obtained from the fact that a
hectare (2% acres) of mature oak trees will transpire as much as
25,000 liters of water per day. This volume would be equivalent to
a layer of water 0.25 cm deep over the whole area. In other words,
if
tra'nspiration continued at this rate for 10 days, a rainfall of at least
2.5 cm would be required to One might think that
restore the water.
the maximum rate of water would occur from saturated soil or
loss
from a pond. Actually the rate of water loss per unit area from the
ground due to the transpiration of the vegetation may be nearly twice
as great as that from a free water surface due to evaporation.
40C
100
12 18 6 12 18 6 12 18
Thursday Friday Saturday

of temperature (broken line) and relative
FIG. 4.11. Hygrothermograph record
in the short-grass plains of central United States during 3
humidity (solid line)
days in early July. ( By permission from Plant Ecology by

Weaver and Clements,
1938, McGraw-Hill Book Co.)
Three atmospheric conditions that greatly modify the rates of evap-

oration and transpiration are the saturation deficit, the temperature,
and the wind velocity. The saturation deficit is the difference be-
tween the actual vapor pressure and the maximum possible vapor
pressure at the existing temperature.
The saturation deficit thus gives
more information of ecological significance than the relative humidity
116 Water
alone since the saturation deficit provides a measure of the capacity
of the air to take up additional moisture. An increase in the satura-
tion deficit produces a rise in evaporation rate.
An increase in temperature similarly speeds up the evaporation
process. Agood indication of the magnitude of this influence is ob-
tained from the amounts by which reservoirs are lowered in regions
of different temperatures. During one year the water level in a
reservoir in Ontario dropped 38 cm, a reservoir in California lost 2.4
m, and one in Egypt was lowered 3.6 m through evaporation alone.
The distribution of plants often reveals the influence of temperature
on direct and indirect water loss. On mountain slopes vegetation
zones are found at different altitudes according to the exposure to the
heat of the sun. Moisture-demanding species are restricted to
higher levels on the side of the mountain toward the equator than they
are on the cooler opposite side. The variation in the amount of rain
required for a good growth of short grass furnishes another illustra-

tion of the effect of temperature. In Montana a precipitation of
about 35 cm is sufficient, but in northwestern Texas a rainfall of 53
cm is needed to produce the same amount of grass. The explanation
of this fact is
easily apparent when it is realized that evaporation in
the 6 summer months Montana amounts to 84 cm, whereas in
ifl
Texas it averages 137 cm (Weaver and Clements, 1938).

The wind velocity also exerts a major effect on the loss of water.
With a gentle zephyr of only 8 km per hr the transpiration of plants
is increased 20
per cent over that which they exhibited in still air. A
wind velocity of 16 km per hr increases the transpiration by 35 per
cent and one of 24 km per hr increases it by 50 per cent. The desic-
cating action of warm dry winds sometimes prevents the invasion of

windward mountain slopes by vegetation that grows perfectly well on
the leeward sides of the same slopes where water loss is much re-
duced. The harmful effect of excessive evaporation caused by the
wind may also be seen in one's own garden. Here winter killing is
occasionally due not low temperatures, but to a combination

to very
of wind and high temperatures! If a warm, dry wind blows for too
long a period when the ground is frozen, the

plants may lose water
faster than their roots can obtain it and
they will succumb as a result.
Our discussion of moisture in the soil and in the air has indicated
that living organisms particularly the higher plants may exert a pro-
found reciprocal influence on the water conditions in the terrestrial
environment. Water
a highly modifiable factor in land habitats,
is
and living elements may act either to augment or to diminish the

amount of moisture present. Vegetation tends to catch the rain, so
Microclimates 117
that, if the precipitation comes in short showers, the rain may never
reach the soil. The plant cover also tends to reduce the water con-
tent of the deeper layers of the soil as a consequence of its transpira-
tion, but at the same time it adds moisture to the air. On the other
hand, the presence of forest vegetation favors the reduction of evap-
oration at levels near the ground because it lowers the temperature,
slows down the wind, and sometimes causes increased condensation.
The amount of water present at any point in a land habitat and the
rates of gain and loss are thus seen to be the result of the equilibrium
of processes both climatic and biological. The interrelations of the
water factor and the vegetation are frequently very complicated and
are the subject of elaborate studies beyond the scope of this book.
For further information the reader is referred to such treatments as
Daubenmire (1947, Ch. 3). In addition to presenting the general
picture, the present discussion reveals the interdependency of vegeta-

tion and moisture as another outstanding example of the organism and
the environment acting as a reciprocating system.
Microclimates
In the foregoing discussion of the variation in water loss under

various conditions of humidity, temperature, and wind the values
given were often those characteristic of extensive areas. Very fre-

quently, however, the values of these factors in the immediate sur-
roundings of the organism differ markedly from the regional values.
In other words, the immediate climate of the organism is often sharply
differentfrom the average climate of the region as reported by stand-
ard meteorological records. The realization of this fact has led to
the development by ecologists of the concept of the microclimate. In
considering the effect of climatic influences on plants and animals it is

absolutely essential to measure the environmental factors as they ac-
tually reach the organism. The effective climate, upon which the
lifeof the organism depends, is its microclimate. The characteristics
of the microclimate may, or may not, be similar to the regional climate
(Geiger, 1950).
Local variations in climatic factors of concern to many organisms
may involve distances of several meters in the surroundings, but the
microclimates of forms living in the soil, in rock crevices, under the
bark of trees, or in similar confined habitats often have dimensions
measured in fractions of a centimeter. The concept of microclimates
is
particularly important in relation to moisture because relative
humidity varies widely within short distances in any irregular habitat.

118 Water
In swampy woodland, for example, measurements have shown that
a high rate of evaporation characteristically exists near the tops of
the trees. At half this height the evaporation rate was reduced to
30 per cent in one instance. Lower down, near the damp soil, the
animals and plants were subject to an evaporation rate of only 7 per
cent of that near the treetops. We are consequently not surprised to
find that this verticalchange in moisture, as well as vertical gradients
in light and in other factors, contributes toward the establishment
of a characteristic vertical layering by species of the vegetation and
sometimes also of the arboreal animal life in forests. Among animals
the difference in evaporation rate with distance above the v ground is
of particular importance to soft-bodied insects. Many of these are
known to move higher in the vegetation at night, but during the day
they are forced by excessive evaporation to retreat to lower levels.
Contrasting values for the rate of water loss in two microclimates

were reported in an interesting experiment performed in the Middle
West. Two sets of green ash seedlings were planted in pots; one set
was placed in the midst of a sumac thicket, and the other set was put
out on the open prairie only a short distance away. At the end of 8
days the first group of seedlings was found to have lost an average of
0.38 grams of water per sq cm of leaf surface. The seedlings set out
in the adjoining prairie had lost 0.88 grams of water per
sq cm, or
more than twice as much during the same period.
A traditional method of finding the approximate direction of north
in the woods is
by observing on which side of tree trunks the growth
of green "moss" is thickest. In the North Temperate Zone in habitats
in which the atmospheric moisture is near the critical
point, algae and
other non-vascular plants are characteristically more abundant on the
north surfaces of tree trunks, because the higher evaporation due to
insolation effectively retards growth on the south side. However, in
situations differences in exposure to wind may
many outweigh the
effect of thesun in causing excessive water loss. Thus, although the
"north side" rule does not always hold, direct measurements show
that the growth of epiphytic moss is a very sensitive indicator of micro-
climate, and the striking differences in the growth of algae on op-
posite sides of tree trunks can be seen by anyone who walks through
the woods. An average value for the moisture condition of the forest
would give no inkling of these small-scale ecological differences of
crucial importance to the species concerned.
The burrows of desert animals furnish another example of a micro-
climate in which the physical conditions are radically different from
those of the region as a whole. When the kangaroo rat retreats
Meeting Water Problem on Land 119
within its it breathes air two to five times more humid than
burrow,
that outside. Measurements indicate that, if this animal remained
out of its burrow
continuously, the rate of evaporation from its lungs
would exceed the rate at which it could obtain water, and the re-
sulting water loss would eventually cause death. Temperatures in
desert burrows have been found to be as low as 28C at the same time
surpassed 71C (cf Table 9).

that values for the surface of the soil
Such critical differences in
temperature and in other factors in micro-
climates will be considered further in
subsequent chapters.
Meeting Water Problem on Land

Land plants can live only where sufficient water reaches them either
through the air or through the soil. Some species can get along with
relatively small amounts of water, and these plants, inhabiting deserts
and other arid regions, are termed
xerophytes. Plants eke out an
existence on a reduced water
supply in a variety of ways. Certain
annual plants are adapted to maintain themselves in desert areas
by
remaining in the seed stage during the dry season and then passing
rapidly through their life cycles when a rainy period occurs. Other
desert plants, like the cacti, store water in succulent
organs in suffi-
cient quantities to tide them over
long intervals of drought. In addi-
tion, various species of non-succulent perennials, such as the creosote
bush, sage brush, and many grasses are included among the xerophytes
because they possess unusual ability to endure long periods of per-
manent wiltingin some instances running into years. Lichens and
some mosses similarly can survive a surprising degree of
drying out.
Plants with water relations intermediate between those of the xero-
phytes and the hydrophytes are classified as mesophytes.

Since land animals can move about,
they can actively seek pools
of water from which to drink or can obtain water in the form of snow
or dew. Many are able to endure long periods between visits to the
water supply. Some terrestrial animals, such as
frogs and toads,
have the ability to absorb water through their skins from
damp sur-
roundings, whereas others find sufficient liquid water in the tissue
fluids of their food. Another method of solving the problem of
securing water in dry land habitats, and one employed widely by the
insects, is the use of metabolic water, that is, the water resulting from
the chemical breakdown of food materials. The clothes moth and
the meal worm have the ability to obtain metabolic water in sii%sient
quantity to enable them to live their whole lives without any addes$
to free water in the environment.
Experiments indicate that the
120 Water
kangaroo rat of the Arizona desert may similarly secure the whole of
its water
supply as a metabolic by-product of its food (Schmidt-
Nielsen, 1950).
methods obtaining water are means
Just as important as for for
resisting desiccation. Many of the same general types of adaptations
having this effect have appeared in the course of evolution amongst

both plants and animals. First and most obvious is the possession
of a more or less impervious covering for the body. The cuticle of
plants, the skin of birds and mammals, the exoskeleton of insects, and
the mucous secretion of mollusks are
examples. Another means for
avoiding the excessive loss of water is the reduction of body surface.
The leafless plants of the desert are an instance of this
plan carried
to an extreme. Other plants fold their leaves or turn them edgewise
to the
scorching heat of the sun. Many terrestrial invertebrates
bury part or all of their bodies to reduce evaporation. It is well
known that the termites which attack our houses are killed by exposure
but these pests can cross exposed foundations by construct-
to dry air,
ing runways within which protection from excessive evaporation is af-

forded. In arid regions
many of the smaller animals are active only
during the night when drying is less intense.
Water may also be saved by reducing the
loss in respiratory and
excretory systems. Respiratory organs are often enclosed within a

cavity as has already been mentioned in relation to the gills of land
crabs. Lungs are commonly located well within the body with only
a small opening to the exterior. The branching tubular tracheal sys-
tem of the insects is another example, and in many forms the size of
the external openings of the tracheae may be reduced by the spiracles.
In a similar way evaporation from the internal cavities of the plant
leaf is restricted by the guard cells of the stomata.
Reduction in the amount of liquid needed for excretion is a physio-
logical adaptation that also aids the body in conserving water. Many
animals with little available moisture in their habitats and in their
food produce dry faeces. Furthermore, in the interest of saving
water, it is desirable for land dwellers to dispose of their nitrogenous
wastes with the minimum quantity of liquid urine. Mammals ac-
complish something in this direction by reabsorbing part of the water

in the urine before it is excreted. The ultimate attainment in this
matter reached by those birds, reptiles, and insects that are able to
is
excrete nitrogenous wastes as solid uric acid practically without

water loss.
If lack of water becomes still more acute, it

may be necessary for
the organism to go into a dormant condition in which its need for
Influence of Moisture on Growth and Distribution 121
water and its loss of water will be reduced. In the tropics many
trees shed their leaves during the dry season. Other plants pass
periods of drought as seeds or as spores. Many lower animals form
cysts of one sort or another. Terrestrial mollusks close their shells
with a thin epiphragm of secreted mucus, and more highly organized
animals, such as the ground squirrels, go into a state of torpor in
their burrows. Dormancy of this sort occurring under conditions of
heat and drought is referred to as aestivation, and is found character-
istically
and tropical communities. Using various combina-
in desert
tions of the methods for procuring and conserving water discussed

above, plants and animals in arid habitats may live for months or
years without rain or access to free water.
Influence of Moisture on Growth and Distribution
The growth of plants is often more directly and obviously affected
by the water factor on land than is that of animals. The availability of

moisture influences not only the rate of plant growth but also the
growth form. Since the roots of most land species will not grow into
saturated soil nor into soil devoid of available water, plants will be
shallow rooted either if the water table is
high or, in the contrasting
situation, if the soil water
limited to the uppermost layer.
is In the
latter case growth will be confined to the rainy season. The growth
of plants whose roots extend to the permanently moist subsoil are
largely independent of rain periodicity.

Competition for water and the horizontal spread of roots often con-
trol the spacing of plants in regions where water shortage is critical.
The extent of the root system in a typical cactus is shown in Fig. 4.12.
The be spread widely just beneath the surface of the
roots are seen to
soil where they can absorb whatever rain falls before it evaporates.
This cactus will capture all the available moisture within the area
shown and prevent any other perennial plant from gaining foothold
in close proximity. Desert vegetation is often spaced out in a strik-
ingly regular pattern as a result of this intense root warfare for water.
The water factor also controls the geographical distribution of
plants on both small and large scales. Such trees as willows and cot-
ton woods are characteristically limited to the moist banks of water
courses because their seeds will not survive more than a few days
unless the soil upon which they fall is wet. The seeds of other species
can remain dormant completely dry condition for months or years
in a
but will germinate only when sufficient water is available in the soil
immediately surrounding them. In other instances the critical period
122 Water
for the survival of a
plant in relation to its water need may occur
during adult life. Plants differ greatly in the amounts of water that
they must absorb from the soil and transpire. Those with a high
water requirement, or transpiration ratio, are limited to habitats where
the supply of moisture is
adequate.
FIG. 4.12. Top view of surface roots of cactus plant shown on a 30-cm grid.
Almost the entire absorbing system of roots occurred in the upper 2 to 10 cm of
soil.
(By permission from Plant Ecology by Weaver and Clements, 1938, Mc-
Graw-Hill Book Co.).
Differential ability to withstand

drought will also play its role in
If the soil moisture
controlling geographical distribution. drops be-
low the permanent wilting most annuals and other tender
coefficient,
plants will die within a matter of hours or days unless the soil water
is renewed. On the other hand, succulents and plants that can
aestivate are able to withstand varying periods of drought extending
into years with some species. Thus the duration of drought periods
is of crucial importance in determining whether a region may be in-
habited by different plant types.
Local differences in the moisture factor were clearly shown to in-
fluence the species composition of the forest vegetation in a study
made in Indiana (Potzger, 1939). A series of observations in a line
running from north to south over a ridge revealed the fact that soil
moisture was 20 per cent higher on the north side of the ridge than
on the south side. Moreover, the rate of evaporation was more than
twice as great at stations on the south slope than at those on the north
slope. Quadrat counts of the number of trees on the two sides of the
ridge gave the tabulated results. Although other ecological factors
Species North Slope South Slope

Maple 202 31
Beech 112
White oak 80
Black oak 70
Hickory 4 44
undoubtedly played some part, moisture conditions were chiefly re-

sponsible for the very pronounced differences in the occurrence of
the various species on the two sides of the ridge.
Another example of the control of distribution by the moisture fac-
tor, and one on a much larger scale, is found in the regional occurrence
of the natural vegetation in the United States. A glance at Fig. 4.13
will reveal the fact that in the central part of the country, leaving
out of account the irregularities due to the mountains, the zones of
principal vegetation types tend to run north and south. In the eastern
part of the continent the typical natural vegetation is forest (

in areas
where man has not interfered). West of the forest zone is a belt of
tall-grass prairie, and beyond that, running
from Montana and North
Dakota down to Texas is a zone of short-grass rangeland. Still farther
westward lies the desert belt. These fundamental zones are indicated
schematically in Figure 4.14.
This zonation is obviously not controlled primarily by temperature
since the temperature belts are seen to run generally east and west,
124 Water
'"_''.""
'
Modified from the Graphic Summary of American Agriculture, U. S. Department of Agriculture

FIG. 4.13. The distribution of the natural vegetation in the United States.
if we
again out of account the mountain and coastal regions
leave
(Fig, 5.14). Similarly, no good correlation will be found between

the occurrence of these principal zones of vegetation and the annual
precipitation taken by As already explained, regional dif-

itself.
ferences in evaporation must be considered, and water loss- in

also
turn depends in part on temperature. Both the supply and the loss
.. ,.
FIG. 4.14. Schematic representation of the four fundamental vegetational zones

in central United States (from This is Our World by Sears, 1937, Univ. of
1
Oklahoma Press).
aspects of moisture conditions taken into account by calculat-

may be
ing precipitation-evaporation ratios for the "frostless season," i.e.,
the
growing season for plants between the last spring frost and the first
autumn frost. The pattern of the distribution of these ratios in the

United States (Fig. 4.15) shows a general agreement with the oc-
currence of the major vegetation zones. The natural forest vegeta-
tion of the eastern part of the country occurs where the precipitation-
evaporation ratios are high. In the Southeast and Northeast, the P-E
ratio exceeds 100 per cent; that is, more rain falls during the course
of the average frostless season than evaporates. Incidentally, this
fact accounts in part for the excessive amount of soil leaching in these
regions. In the Middle West precipitation-evaporation ratios be-

tween 80 and 100 per cent exist. Values below 100 per cent indicate
that the potential rate of water loss is greater than the rate of water
supply. The line representing a P-E ratio of 60 per cent runs almost
due south from the Dakotas to the southernmost point of Texas. It
will be noted that this line agrees closely with the division between
the tall-grass prairies and the short-grass rangelands. The 60 per cent
line also approximates the position of the division between the pedo-
cals and the peclalfers discussed earlier. P-E ratios .of less than 20
cent are found in the southwestern states, and their occurrence
per
corresponds to the distribution of the desert vegetation.
In view of the modifying influence on plant distribution of such
tt ftt
-,
Modified from Livingston and Shreve, 1921

FIG. 4.15. The precipitation-evaporation ratio indicated as a percentage for the
average frostless season in the United States.
126 Water
factors as soil type and temperature and of the interdependence of
these factors with moisture it is clear that no simple relation between
the type of vegetation and any single physical influence can be ex-
pected. However, the broad correspondence between the zones of

vegetation and the availability of moisture will suffice to emphasize
the major role of the water factor in the region under discussion in
controlling both the soil and the plant life that develops with it.
The application of the P-E ratios to these distributional problems has
again brought into relief the fact that the balance between supply
and loss is the most crucial aspect of the moisture factor for the vege-
tation.
The water factor on land also seriously affects the growth and dis-
tribution of animals either directly or indirectly. Since the range of
land animals is
profoundly influenced by the vegetation, moisture
often exerts its
greatest effect on animals indirectly through its con-
trol of the plants. For the higher vertebrates temperature is
generally
a more important direct environmental influence than moisture.
However, it has long been known that races of birds and mammals
in warm humid regions tend to be darker in color than races inhabit-
ing the cooler and drier parts of the geographical range of the species.
This generality is known as Glogers rule, but many exceptions exist
and the relative effects of heat and moisture are not known. There
isevidence that humidity acts more through the color of the soil as a
background for the animals than in a direct way. The implications of
Gloger's rule will be discussed in more detail below in relation to
temperature and light.
For amphibians and for insects and other terrestrial invertebrates
moisture often of great direct importance. Many insects exhibit
is
critical dependence upon humidity conditions with sharp limits of

tolerance. For example, the "silverfish" (Lepisma saccharina), a
common household insect pest, finds optimum conditions for repro-
duction at relative humidities of 85 to 90 per cent. The newly
hatched nymphs die if they are subjected to relative humidities of less
than 70 per cent, and they also succumb if their surfaces once become
wet. The
relationship of inserts to moisture is frequently very com-
plex and varies greatly in different species and sometimes even in
different parts of the life cycle of the same species (cf. Chapman,
1931, Ch. 4). In the development of the flour beetle Tribolium the
larval stage is accelerated by an increase in relative humidity, but the
duration of the egg and pupa stages is unaffected by wide changes
in this factor (Fig. 4.16). Moisture produces entirely different ef-
fects upon the survival of this insect. In the larval stage increasing
humidities tend to improve survival, but the egg and pupa stages are
affected to only a slight extent
by relative humidities up to 70 per
cent. However, at humidities above 80 per cent the viability of eggs
and pupae drops very sharply.
When become adverse,
the moisture conditions of the environment
the animals concerned either die or must migrate longer or shorter
distances to more favorable locations. Droughts greatly reduce the
occurrence of some species temporarily, or for long periods, by deci-
mating the population. Locomotor activity, as well as feeding and

other reactions, are controlled by humidity in certain animals, espe-
cially insects. It is known that in some instances increased

activity
without orientation results from a change in moisture conditions; in
Fig. 4.16. Duration (upper) and survival (lower) of the indicated stages of
Tribolium confusum at different relative humidities under constant temperature
of 27C (by permission from Animal Ecology by Chapman, 1931, McGraw-Hill
Book Co.).
128 Water
other instances animals are able to respond clirectionally to a humidity
gradient. However, the whole problem of control of locomotion by
moisture in land animals calls for further investigation.
The discussion of water relations in this chapter emphasizes the
universality of the water problem. Plants and animals in aquatic,
amphibious, and terrestrial situations are faced with the difficulty of
providing for water exchange and at the same time maintaining the
proper water balance. The amount of water present in the environ-
ment and its transfer into and out of the organism are results of an
equilibrium of forces. In the terrestrial environment the organisms
themselves, especially the plants, can affect the amount of moisture in
their surroundings. The total amount of water is not of as great
significance as its
availability. In some situations the environment
tends to extract water from the plant or animal at an alarming rate.
If the organisms cannot obtain water fast
enough and retain it in suf-
ficient quantity, the lack of this essential material becomes a critical
limiting factor.
5
Temperature
Temperature perhaps the most commonly familiar ecological fac-

is
tor. The
great variation in heat conditions and their general influence
are evident to everyone. In contrast to many ecological factors tem-
perature may be measured with relative ease. Temperature is a uni-

versal influence andfrequently a limiting factor for the growth or
is
distribution of animals and plants. Even when an organism is dor-

mant, the chemical processes going on in its body are controlled by
the existing temperature. This factor is also important indirectly as
modifying the effects of other ecological agents.

Temperature is the intensity aspect of heat energy. The capacity
aspect of this form of energy will also be considered in this chapter.
Differences in the heat capacity of the media, of the various types of
substrata, and of the bodies of organisms themselves are significant
in
controlling susceptibility to temperature change. The amount of
heat in an ice-covered lake may change considerably without any
alteration of temperature, but in most natural situations increase or
decrease in heat also affects the temperature. Both aspects of heat
energy must be kept mind, but temperature has the more predom-
in
inant direct influence on the lives of organisms.
DISTRIBUTION OF TEMPERATURE
Extremes of Temperature and of Tolerance
In considering heat conditions in -natural environments the extremes

of temperature may first be reviewed. In the open water of the
aquatic environment the temperature cannot drop below the freezing
point. This means that the ternperature of the water in ponds is
never lower than 0C
and in the ocean never lower than about
129
130 Temperature
2.5C. The maximum temperature in marine environments of any
size is
probably represented by records of 36C in the Persian Gulf.
In tide pools of the littoral zone and in shallow bodies of fresh water
temperatures may go higher,
On land the record for the lowest temperature is held by a locality
in the north interior of Siberia where the thermometer was read at
70C ( 93.6F) in 1947. Temperatures almost as low were re-
ported for a northern outpost in the Yukon territory. At the other
end of the scale air temperatures ranging above 60C (140F) are
recorded in desert areas. Desert soils have been found to rise as high
C when exposed to the noonday sun. The water in hot springs
as 84 9
and geysers may approach 100C, and even higher temperatures occur
sporadically in the very special situations presented by volcanic areas.
Are these environmental temperatures beyond the ranges that can
be tolerated by animals and plants? Birds and mammals are warm-
blooded animals, or homoiotherms, that maintain their own constant
internal temperature, and their tissues are insulated from the heat or
cold of the outside world. All other animals are cold-blooded forms
or poikilotherms. The tissues of these animals and of all plants, which
are also poikilothermous, tend to approach the temperature of their
immediate surroundings and to vary with external thermal conditions.
No organism can continue to live in an active condition at tempera-
tures below those at which its tissues freeze. As we shall see below,
the freezing points of living tissues differ widely, but no poikilother-
mous animal or plant can actually grow at continuing temperatures
lower than a few degrees below 0C. At the other end of the thermal
scale we find certain blue-green algae and thermophilous bacteria
growing happily in hot springs, such as those in Yellowstone Park, at

80 to 88C! However, relatively few poikilotherms can live perma-
nently at temperatures above about 45C, and Brues (1927, 1939)
found no reliable records for poikilothermous animals above 52C.
It appears from the foregoing that the whole range of temperature
in the sea is within the limits of tolerance of many plants and animals.
In contrast, land temperatures may be far below or far above the
temperatures that can be withstood by organisms in an active condi-

tion. Since the range of heat conditions sustained successfully by in-
dividual species is
very much less, temperature extremes present a
problem on both land and sea. The homoiothermous animals form a
special case since the thermal range that they can tolerate is generally
much greater than that of other animals and of plants, but even this
group has definite limits of toleration. We
shall be concerned, then,
with the limiting action of extreme temperatures, as well as with the
Changes in Time 131
controlling action of intermediate temperatures for all kinds of plants

and animals.
Changes in
Temperature
Changes in Time. Let us consider first the temperature changes to

which an animal or plant may be exposed if it remains in the same
place. Time changes in temperature are controlled by various astro-
nomical and climatic cycles. Within a few hours the sun
may move
so as to the of a form from
change exposure sessile direct sunlight to
shade. On land a very great difference in the heat received may oc-
cur within a short period of time. In the water environment the
change from sun to shade produces only a minor effect less than
0.1C at a depth of 5 m.
Similarly, the diurnal fluctuation in temperature is very much
damped in the aquatic environment. In a body of water of any con-
siderable size differences between day and night are commonly less
than 1C. The maximum diurnal change in the ocean is about 4C,
and with increasing depth the amplitude is reduced. Probably no
diurnal temperature change is detectible below a depth of 15 m.
Time of day
8 12 16 20 24 4 8 12 16 20 24 4 8 12 16 20 24
Modified from Weaver and Clements, 1938

FIG. 5.1. Portion of a soil thermogram made in June on the prairie near Lincoln,
Nebraska, showing the temperature fluctuations at depths of (A) 7.6 cm and
(B) 30 cm.
The temperature of the air near the surface of the land is sometimes
17C higher in the daytime than at night, and in desert localities this
difference may be as much as 40C. still A
greater diurnal range in
temperature reported for the surface of the soil in desert areas. In
is
every situation the amplitude of the change from day to night in the
soil is reduced with depth. The soil thermogram shown in Fig. 5.1
illustrates this point and also reveals the lag in the time of occurrence
of the maximum and minimum daily temperatures. At a depth of
132 Temperature
only 30 cm the highest temperature of the day occurred near midnight
and the lowest temperature occurred near noon!
In the tidal zone changes in temperature are the result of a com-
bination of the differences in the amount of heat delivered from the
sun and of differences in the temperature of the air and of the water.
AH organisms living between tide marks are subject to rapidly chang-
ing temperatures that may test their tolerance to the limit.
At one
locality on the coast of Maine the temperature of the mud exposed at
low tide was observed to rise to 38C under the midday summer sun.
A short time later plants and animals living in this area were flooded
by the incoming tide with a temperature of 10C. In the winter

when the tide is out these organisms might be exposed to tempera-
tures as low as 25C. The occurrence of these extreme and rapid
fluctuations in temperature adds to the rigorous conditions of the tidal
zone.
Plants and animals are also concerned with seasonal cycles of tem-
perature. In both tropical and polar seas the intensity of heat does
not vary by more than about 5C
throughout the year. In the tem-
perate seas, however, changes of 10 to 15C from summer to winter
are common, and sometimes seasonal differences of 23C or more are
observed. At increasing depths in the water these differences become
less. Measurements off Portsmouth, New Hampshire, where there
was a 15-degree change in the surface temperature showed no meas-
urable variation from summer to winter below a depth of 140 m.
Probably more than 95 per cent of the oceanic environment exhibits

no seasonal change in temperature that is of significance to living or-
ganisms. In lakes and ponds seasonal fluctuations are generally
greater than those found in the sea. Springs, however, form a very
special fresh-water habitat in which the temperature may vary by only
a degree or two the year around. A typical example of the thermal
cycle in a lake showing the reduced amplitude in the deeper layers is
presented in Fig. 5.2.

On land the seasonal changes in temperature are familiar to every-
one and are almost always of ecological significance. The greatest
variations are found in continental areas in the temperate zones with
and near the coasts. At St. Paul,
lesser seasonal ranges in the tropics
Minnesota, the average difference between summer and winter is

33.7C. Possible extremes in range are even greater. From Tibet
comes a report of temperatures varying by 77C from 37C in the
winter to -f 40C in the summer. In contrast to the foregoing are
certain localities where temperatures vary only slightly throughout
r
(
tS
-
a
s
'H
I.E
5 .a
tlS
U3
I
I
-S
0)
CT5
f g
E
S
no
o
134 Temperature
the year. At Quito, Ecuador, seasonal
for example, the average
change is
only 0.5C.
Horizontal Changes. The changes in temperature from place to
place over the surface of the globe run the whole gamut of extreme
values that have been reviewed. Average temperatures on land are
too variable for simple generalization, but a value of 32C may serve
as an approximation for
tropical regions and one of polar 12C for
regions. For detailed information a textbook on climatology should
be consulted, such as Kendrew (1949). In the sea average tempera-
tures run from about 30C in the 1.5C in the Arctic
tropics to about
and Antarctic Oceans.
Vertical Changes. The temperature of the air varies widely in a
vertical direction according to local conditions, but a decrease of
about 1Cfor every 150 m
of altitude is generally found. On Mt.
Washington in New
Hampshire the tabulated average values were
obtained for observations from 1933 to 1940.
Feb. July Year

Pinkham Notch (Altitude 610 m) - 8.5C 17.1C 4.2C
Summit (Altitude 1915 m) -14.9 9.9 -2.8
Difference: 6.4C 7.2C 7.0C
The unequal heating of the air horizontally and vertically causes

movements of the atmosphere. These are manifested as local winds,
trade winds, and storms with consequent further influence on tem-
perature, precipitation, and other ecological factors.

Air is heated at all levels
by the solar radiation that it absorbs and
similarly may cool at but the greatest amount of heating
all levels,
and cooling takes place at the bottom. This effect is due to the great
transparency of the air and the fact that the surface of the earth is
heated faster during the day and cools by radiation more rapidly dur-
ing the night than the atmosphere. The air in immediate contact with
the earth therefore changes temperature more rapidly than strata at
greater altitudes. The easy mobility of the atmosphere also contrib-

utes to the thermal variations of the air environment. When air
overlying heated land masses becomes warm,

it tends to rise and to
leave the earth's surface, but air chilled by contact with cold earth is
heavier and tends to accumulate in hollows of the landscape. In
these respects air stands in distinct contrast to the water environment
in which heating and cooling take place principally at the upper
surface.
The of the energy received from the sun at the
spectral distribution
vertical U flanges 135
earth's surface is shown in
Fig. 5.3. It will be seen that about half of
the solar radiation in the infrared
region and this represents a di-
is
rect supply of heat. The visible and

ultraviolet portions of the sun's
emission will also produce a
heating effect after they have become
absorbed. The sample calculations in Table 7 illustrate the relative
absorption of the various parts of the spectrum by air and water.
3000 5000 7000 9000 11,000 13,000 15,000 17,000 19,000 A

Wavelength
Data from Fowle, 1927
earth's surface, showing the
Fie. 5.3. Spectral distribution of solar energy at the
ultraviolet (UV), visible (divided into color components), and infrared portions.
TABLE 7
COMPARISON OF RELATIVE ABSORPTION OF SUN'S RADIATION BY AIR AND

WATER
Visible Light Infrared Radiation
at 5500 A at 8000 A
Percentage Percentage Percentage Percentage
of initial absorbed of initial absorbed
value by each value by each
remaining stratum remaining stratum
Outside atmosphere 100 100
Mt. Whitney, California 93 97

9
(summit 4420 m)
Sea level 75 88
5 98
2 m below surface 71 2
70
30 m below surface 17
(clearest ocean water)

136 Temperature
The data show that the atmosphere absorbs radiation of short wave-
is most
length faster than that of long wavelength, whereas water
to radiation in the central of the visible region.
In
transparent part
all
parts of the is absorbed
spectrum radiation by water at far higher
rates than by air, and the contrast is in the
particularly great
infrared. The absorption by a few meters of water is enormously
greater than by several thousand meters of air. In general we
can
say that in comparison with the atmosphere the water medium is al-
most opaque to the sun's radiation.
Because of the differences in mobility and transparency of the
two media we find that the vertical temperature changes in the water
environment are controlled differently from those in the air. Radia-
tion from the sun isthe only important source of heat for natural
bodies of water. Tests have shown that no significant exchange of
heat takes place between the water and the mud of the bottom. Since
solar radiation is absorbed in the upper strata of water, water bodies
receive heat only at their upper surfaces. Cooling of water can take
place through radiation, evaporation, or the melting of ice. All of

these processes occur at the surface in natural water bodies. Gen-
erally speaking, the seas and the inland waters gain and lose their
heat primarily at the top.
It is not
surprising to find as a consequence of the foregoing that in
deep bodies of water the major changes in temperature are limited
to the surface strata. In some situations these changes are superim-
posed on a deeper, permanent temperature structure. In the tem-

perate and tropical regions of the sea a permanent thermal gradient
has been produced between the mixed layer near the surface and the
deep layer filling the bottom of ocean basins. The depth and extent
of the zone of relatively rapid temperature change, known as the
permanent thermocline layer, differ in the various parts of the ocean

but an illustrative example is
given in Fig. 5.4 for water in the great
central eddy of the Atlantic Ocean. From roughly 2000 m down
to the bottom the water is everywhere just about 3C. Since the
average depth of the ocean is approximately 4000 m, there is ob-
viously a very large amount of space in which the temperature is uni-
form month in and month out, year in and year out. If you were a
fish restricted to 3C, you could nonetheless travel over more than
60 per cent of the globe without being exposed to a significantly dif-
ferent temperature.
Although essentially all the incident radiation is received at the

surface of natural water bodies, we know that some heat is
eventually
transferred to deeper layers. How does the heat get there? Radia-
tion from one water layer to another is Conduction
negligible.
through quiet water is similarly very slow. If all the water in Lake
Constance with a depth of 100 m were cooled to and then the 0C
uppermost layer heated to 30C, more than 100 would be re-years
quired for a measurable amount of heat to reach the bottom by radia-
tion and conduction alone. It is obvious that heat can be transferred
Temperature,C
8 12 16
Deep layer
2000 L.
Modified from Iselin, 1936
FIG. 5.4.Permanent vertical temperature structure the deep water of the
in
eentral North Atlantic. The seasonal changes in upper layers are indicated; (A)
April, (B) August, (C) December, (D) February.
to lower levels only by the actual movement of the water, that is, by
vertical circulation.
We are further convinced when we examine situa-
of this conclusion
tions where been prevented. When the tem-
vertical circulation has
perature of the bottom water of a pond has been reduced to and 4C

138 Temperature
ice has formed across the surface, no by the wind can
vertical stirring
take place. Although the air temperature may go far below zero,
heat can be lost from the deeper layers of the pond only very slowly
because of the insulating action of the ice and the upper strata of
water. This is the chief reason why most ponds do not freeze to the
bottom during the winter.
has been
Stirring is also inhibited when a sharp density gradient
Such a
developed between the upper strata and the deeper layers.
condition arises in situations in which the surface water is warmer or
less saline, and hence
lighter, than the deeper water.
Under these
circumstances the vertical stability of the water tends to resist any
efforts on the part of the wind and waves to produce stirring, and heat
exchanges are consequently limited to the surface. When no impor-

tant density gradient exists, vertical stirring can become effective and
heat will be carried down. This condition commonly occurs during
the springand during the autumn in waters of temperate regions,
Under some circumstances the density of the surface stratum may be-
come actually greater than the underlying layers resulting in a gravita-
tional overturn of the water.
Seasonal Changes in Vertical Temperature Structure. The typical
temperature cycle in a lake during the course of a year is indicated
diagrammatically in Fig. 5.5. Since the maximum density of fresh
water occurs at 4C, any warmer or colder water will float on top of
water of this temperature. If the underlying water has a temperature
lower than 4C as it
may in the C
early spring, or higher than 4 C, as it
may in the autumn, then a top-heavy condition will
develop.
Archimedian forces aided by strong winds will then bring about the
spring overturn and the fall overturn. Winter stagnation occurs when
the lake is ice covered and summer found after a dis-
stratification is
tinct density gradient hasbeen produced by the warming of the sur-

face layers. The layer of rapid vertical temperature change is termed
the thermocline and in lakes is defined as having a thermal gradient
of at least 1C
per m. In lakes the wind-stirred and largely homog-
enous water layer above the thermocline is known as the epilirnnion
and the relatively stagnant water mass below the thermocline as the
hypolimnion. The very considerable changes in temperature and the
occurrence of stagnation and of circulation which they control exert
a profound effect upon the biota of the lake as will be discussed in
later sections. For further details on the thermal cycles of lakes the
reader should consult a textbook of limnology such as Welch ( 1952 )
orRuttner (1953).
A similar seasonal cycle of temperature occurs in the ocean but the
circumstances are somewhat different. For a fuller treatment of the
subject reference should be made to a textbook of oceanography such
as Sverdrup 1942). In the open ocean the seasonal changes are
et al. (
superimposed upon the permanent thermal structure as relatively

minor fluctuations (Fig. 5.4), but in coastal regions the water may be
Winter
-> Spring overturn
stagnation
Ice cover
20
Summer _ -+~ Fall overturn

stratification
22
Epilimnion
21
Thermocline
9
Hypo-
limnion
20
Fie. 5.5. Diagrammatic deep lake of the temperate region,
vertical section of a
showing the seasonal cycle of temperature (by permission from Limnology by

Welch, 1952, McGraw-Hill Book Co.)
for the seasonal effects to reach the bottom (Fig.

sufficiently shallow
5.6). In this type of situation the
thorough stirring of the
water
uniform temperature from top to bottom
during the winter produces a
with a minimum value in February for the North Temperate Zone. As
spring comes on, the larger supply of solar heat, often combined with
lowered salinity
due to increased run-off, causes the surface water to
become and tends to stabilize the upper layers. This effect is
lighter
self-accelerating. By August the relatively thin stratum at the top has
attained its maximum temperature. Below this layer a sharp ther-
140 Temperature
mocline leads to the deeper water which is still quite cold. With the
onset of autumn solar radiation has become reduced and the winds
have been stronger with the result that the surface waters are stirred
downward, and the thermocline is shifted deeper and eventually de-
stroyed. By November the whole water column has become mixed
at an intermediate temperature and remains uniform as it cools to its
minimum temperature during the winter.
Temperature, C
2 4 6 8 10 12 14 16 18 20 22 24
Fie. 5.6. Seasonal changes in the vertical distribution of temperature in coastal

water off New York, showing the positions of the thermocline ('/') (Modified from
Clarke, 1940.)
The radiation received from the sun reaches a maximum in

June in
the North Temperate Zone, but owing to the lag in the heating effect
the highest temperatures on land ordinarily occur in July. At the
surface of water the maximum heat of the season is not experienced
until August, and in subsurface layers the thermal peak is even more
delayed. If you imagine yourself a worm living in the bottom mud
in the situation
represented by Fig. 5,6, "summer" for you would not
begin until November! In similar fashion the seasons are often com-
pletely reversed in deep lakes.
This review of the thermal conditions has shown that in the ocean
and in lakes there is no temperature too high and no temperature too
low for active life of some kind. Temperature changes are much
less and much slower than in the terrestrial environment. Further-
more, organisms can usually get out of excessively high or low tem-
peratures by a short journey into deeper water. In these respects life
in the water is much easier than life on land. The results of this
Biological Action of Temperature 141
situation over the ages has been that many oceanic plants and animals
have become adapted to a
relatively stable temperature.
The disadvantage of this
easy life in the water is that when an un-
usual change in temperature occurs dire results
may follow. Many
instances are known in which
slight temperature changes have caused
mass mortality in the aquatic environment. A
spectacular example
occurred in 1925 Peru (Murphy, 1926). In the spring
off the coast of
of that year the cool Humboldt Current, which flows northward along
the coast, apparently moved slightly offshore, allowing the warm
countercurrent, El Nino, to flow farther south than usual, raising the
temperature 5 or 6C above normal. The result was that the plankton
and a great many of the local fishes were killed. Larger fishes, de-
pendent upon these smaller forms of life for food, soon afterward
succumbed and washed up on the shore in windrows. In addition
huge numbers of fish-eating birds died of starvation. Many of these
birds inhabited coastal islands where their droppings produced guano
deposits of extreme value as fertilizer. Guano production was

stopped, and the coast line for miles was strewn with the carcasses of
the dead birds. Torrential rains caused by the slight change in ocean
temperature washed away tons of guano that had required generations

to accumulate. The heavy rainfall also devastated the neighboring
ordinarily arid land areas, disrupting the natural fauna and flora, and
destroying crops, roads, and buildings. The ecological and economic
repercussions of the oceanic change thus extended to the shore and
inland. A few months later the ocean currents returned to their nor-
mal course, and after a period of years the fauna and flora were grad-
ually restored. This cataclysmic destruction of ocean life off Peru
well illustrates the far-reaching consequences of a slight temperature
change in a situation where the plants and animals have become ad-
justed to relatively uniform conditions.
BIOLOGICAL ACTION OF TEMPERATURE

What are the ecological consequences on land and in the water of
the thermal conditions that have just been reviewed? Environmental
temperature exerts a direct action on the tissues of poikilothermous

organisms, that is, on those whose body temperatures vary with the
surroundings. The heat exchanges of homoiothermous animals are
also affected by external thermal conditions, but the insides of these
animals are thermostatically maintained at a nearly constant, and
relatively high, temperature.

142 Temperature
Extreme Temperatures
Minimum Temperatures. We may consider first the danger to the

tissues.
organism of the freezing, or the actual congealing, of its living
Freezing is likely to produce mechanical harm in the
form of the rup-
ture of cell walls and the stoppage of circulation. When ice crystals
are formed, they also withdraw water from neighboring areas, pro-
ducing a condition similar to desiccation in the surrounding

cells. No
tissue will freeze at 0C. The concentration of solutes in body fluids
and tissue fluids causes a depression of the freezing point with the
result that even a tem-
unprotected protoplasm will not freeze until
perature a few degrees below zero is reached. For this reason many
can be active at 0C; will germinate on
tissues
rye seeds, for example,
ice.
In situations where the temperature drops considerably below zero

the danger of freezing is ever
present, Some forms have special
adaptations for lowering the freezing point of their tissues still farther
so that congealing does not take place until a very low temperature is
reached. In addition a few species are able to withstand actual freez-
ing for short periods of time. The green alga, Chlorella, while in
active vegetative condition, has been frozen at 182C for 1 hour
without harm. At subzero temperatures ice crystals are known to
occur in the needles and wood of the Norway spruce, and their pres-
ence evidently produces no serious injury to the tree. Among the
higher animals the Alaskan blackfish has the ability to recover normal
activity after being frozen for periods of 40 minutes at temperatures
down to 20C.
Many plants and animals are killed by temperatures that are too low
for them but are nevertheless far above values at which tissues would
actually freeze. Similarly, at the other end of the scale we find many
instances of death from excessively high temperatures but at values
below those producing heat coagulation. Lethal extremes vary greatly
from species to species, and the organism as a whole
may be killed by
a degree of chilling insufficient to cause direct
damage to individual
protoplasmic structures. Temperatures that are too low for some
species may be favorable, or even too high, for others. The English
daisy grows and flowers best when subjected to night temperatures
below 10C and dies if kept for long periods above 20C. In contrast
the African violet is killed
by long exposure to night temperatures
below 10C and grows best and flowers when the thermometer stands
Maximum Temperatures 143
above 20C (Went, 1950). Many do not venture south
Arctic fish
into waters as warm as 10C, but this same temperature would be far
too cold for most tropical fish.
If the tissues do not freeze,
why are these animals and plants killed
by moderately low temperatures? Most chemical reactions are slowed
down by lowering temperature and eventually stop. The cessation
of any one of the vital processes will cause the death of the
organism.
However, the various processes going on in the body come to a stop
at various points on the
temperature scale. No one biological zero
exists for all reactions even within the same individual. Under some
circumstances the organism may when all the vital processes
die are
still The explanation may be that the different reactions
going. are
slowed down by different amounts with the result that mutually de-
pendent processes get out of adjustment and cause the death of the
organism.
The minimum temperature for the organism as a whole is thus de-
termined by the most susceptible of the vital processes. The lowest
temperature at which the organism can live indefinitely in an active
state is termed the minimum
effective temperature. After a further
reduction of temperature the organism goes into chill coma. If too
long a period does not elapse before the organism is again warmed, it
will become active once more. The lowest temperature at which
survival is
possible is called the minimum survival temperature. The
actual value of this temperature depends upon the period of exposure.
For the eggs and larvae of the fruit fly, Ceratitis capitata,
example,
were killed at 7C
only after an exposure of 7 weeks, but death was
caused after 3 weeks at 4C
or after 2 weeks at 1C. Accordingly, a
statement of the minimum survival temperature for an animal or a

the period of exposure is also stated.
plant has no meaning unless
Maximum Temperatures. At the other end of the temperature
scale the same sort of situation exists. The maximum effective tem-
perature the greatest intensity of heat at which the species can live
is
indefinitely in the
active state. The effective temperature range within
which the organism can carry on its active life and beyond which death
eventually results extends
between the maximum and minimum effec-
tive temperatures. At higher temperatures the organism goes into
heat coma but will recover if restored before too long to cooler con-
ditions. For the maximum survival temperature the period of the ex-
heat conditions must be given.
posure of the organism to the specific
All these thermal relations are illustrated for the house fly in Table 8.
144 Temperature
Methods of Meeting Temperature Extremes

A surprising number of plants and animals can withstand consider-
able thermal fluctuation but do not possess any special mechanism for
meeting the problem of temperature extremes. When the ther-

mometer rises too high or drops too low, these animals and plants
must either "take it" or die. This is the situation for most sessile
aquat|c invertebrates, for many other invertebrates, for many plants,
and for some higher animals. As we have
seen, getting along without a
in
special temperature adaptation is
easily possible for many organisms
the aquatic environment. On land, however, north and south of the
killing frost line, and in areas exposed to intense heat, something must
be done to survive periods when the temperature exceeds the effec-
tive range within which active
life can be maintained.
Morphological and Physiological Adaptations. During the course

of evolution many specialized structures have been developed that
relate to the temperature problem. Certain animals and plants pro-
duce spores, cysts, eggs, pupae, or seeds that are capable of resisting
thermal extremes. Each of these structures represents a whole organ-
ism. In other species special parts of the fully developed organism
are resistant to extremes of heat and cold. Such is the situation with
the hardy roots and stems of many perennial plants. For example,
the tops of grass plants freeze off in winter. Life remains in the
stolons and roots of the grass, and from these organs new leaves are
produced when warm weather again returns.
TABLE 8
TEMPERATURE RELATIONS OF THE HOUSE FLY, Musca domestica
Death 4fl.5C in few minutes Maximum survival temperature

Heat coma 44 6 .
Excess! ve acti vi ty 40.1

Rapid movement 27.9 \ Maximum effective temperature
I
> Effective temperature range
lo
{23 I
Feeble movement 10.8 / Minimum effective temperature

Begins moving 6.7
Chill coma 6 .
Death 5.0 in 40 minutes Minimum survival temperature

8.0 in 20 minutes Minimum survival temperature
12.0 in 5 minutes Minimum survival temperature
In some species physiological changes take place in the tissues that
prevent freezing. Osmotic concentration is increased, and water is

"bound" in colloid form with the result that the
freezing point is de-
below values
pressed ordinarily experienced. If
you walk through a
field of winter
rye on a day when the temperature is well below 0C,
you will notice that the leaves and stems are flexible and not brittle as
they would be if the tissues were frozen. The completion of these

hardening processes takes time, and the actual high or low tempera-
tures that an animal or a plant can withstand often
depend upon the
interval available for acclimatization.
Evergreen trees and many
other types of plants similarly become "frost
hardy" during the winter.
Everyone is familiar with the
ability of crocuses, snowdrops, and other
spring flowers to withstand frost and even to push up through a layer
of snow. On one occasion a fully opened crocus blossom was ob-
served unharmed after a night during which the air temperature had
dropped to 16C. Various species of alpine plants exhibit a se-

quence flowering in or near snowbanks according to their
in their
differing tolerance of low temperatures. Many insects are similarly

able to form bound water in their tissues and thus avoid the dangers
of freezing.
Another method by which extremes of temperature are endured
involves the removal of water from the tissues.Dried seeds, spores,
and cysts avoid freezing because no liquid remains that can freeze.
Dry seeds have germinated successfully even after exposure for 3
weeks to liquid air (about 190C). In another experiment nema-
todes and tardigrades are reported to have recovered after chilling for
several hours at an even lower temperature. At the other extreme
tardigrades in a resistant condition have endured immersion in boiling
water for short periods of time, and dried cysts of the ciliate, Colpoda
cuculluSy have survived after exposure to dry heat at 100C for 3 days.
The even greater thermal resistance of some bacterial cysts is well
known. Among some plants unusually high temperatures not only
are tolerated but also actually accelerate development. Seeds of the
wattlebark tree are commonly boiled 1 hour to hasten germination
before planting. The seed cones of certain species of pine trees (e.g.,
Pinus banksiana] open promptly only after fire has scorched them.
This fact has obvious implications for the ecological effects of forest
fires as will be discussed later.
Another special adaptation for dealing with extremes of temperature

is
dormancy. The term hibernation is often used loosely to describe
all instances in which metabolism is reduced during winter when the
environment becomes too cold. A great many poikilothermous ani-

mals go into such a hibernating condition in crevices, under rocks, or
in the mud. Plants similarly are sometimes spoken of as hibernating
146 Temperature
when in the dormant winter condition. Plant dormancy is induced
by lowered autumn temperatures often accompanied by desiccation.

Dormancy during the summer, when high temperatures, excessive
dryness, and/or shortage of food may occur, is called aestivation, and,
although less common
than hibernation, it is found widely among
insects and some other invertebrates, as well as among plants and
certain mammals. In many insects dormancy takes the form of a
is, a stage in the development of the animal during
diapause, that
which morphological growth and development are suspended or
greatly retarded. Species having the capacity for diapause usually
display a rhythm in the life cycle that is related to the seasons.
During periods of unfavorable climate the greater part of the popu-
lation is in the resistant, diapause stage. Anclrewartha (1952) re-
ports that in the absence of diapause the grasshopper, Austroicetes

cruciata, which maintains high numbers over a wide area in southern
Australia,would almost certainly die out in most of this area, or at
best become a very rare species. Since in general dormancy is cor-
related with severe environmental conditions, insects of the same
Alter Bodenheirner, 1938

FIG. 5.7. Variation in the seasons of dormancy and in the number of complete
and partial generations in the ladybird beetle, Coccinella septempunctata, in
different parts of its
range. development; aestivation;
hibernation.
species in different parts of the world may go into hibernation or

aestivation at different
quite periods of the year (Fig. 5.7).
Certain mammals, and at least one bird become
(Jaeger, 1949), also
dormant with reduced metabolism under seasonal extremes of climate.
Since this type of
dormancy involves a physiological change peculiar
to warm-blooded animals, a separate term should be
employed to
describe it, but in the absence of a
special word the terms hibernation
and aestivation are used for mammals
(and birds) as well as for
lower animals (Lyman and Chatfield, 1950). In mammalian hiber-
nation the internal temperature
drops to about 1 degree above the
temperature of the animal's surroundings, provided that the latter does
not fall lower than a few degrees above 0C. This is not to likely
happen in the burrows or caves where the animals usually hibernate
(Fig. 5.8). In laboratory experiments a reduction of the external
Photo by D. R. Griffin
FIG, 5.8. Hibernating bats, Myotis I. lucifugus, hanging in clusters head down-
ward by hind claws hooked to the rough limestone in a Vermont cave.
temperature below 0C
causes the hibernating mammal either to be
aroused from its dormant condition or to be killed by freezing.
Temperatures well below 0C
for long periods in the winter would
prevent any mammal from hibernating in the Arctic unless it could

find a sufficiently tempered retreat. Whether for this or other reasons
there are few hibernating mammals in the far north. The Arctic
148 Temperature
ground squirrel hibernates in burrows made in local areas of unfrozen

ground under deep snow. Other small mammals remain active in
lined nests made in the snow itself. Measurements at Fairbanks,
Alaska, showed that at less than 1 m beneath the snow surface a micro-
climate temperature of 5C existed at a time when a value of
50C was recorded in the air just above the snow.
Less is known about the physiological adjustments of aestivation
in mammals, but this type of dormancy similarly involves a shift of
metabolism into low gear and a complete change in the control of
internal temperature (Hamilton, 1939). An example of mammalian
aestivation is found in the summer dormancy of the ground squirrels
of southern California. By remaining torpid in their burrows for
several months these animals not only avoid the high summer tem-
peratures but also tide over periods of water and food shortage.
A special device for dealing with extremes of temperature while in
the active condition is homoiothermy, or warm-bloodedness. Birds
and mammals are able to maintain a remarkably constant internal
temperature despite great variations in the outside world. Limits
exist, of course, beyond which these animals cannot maintain their
temperature control, but by allowing a regulated amount of evapora-

tion to take
place from their bodies they can keep their own tempera-
tures down to normal values under the highest environmental tem-
peratures ordinarily encountered. Under extremes of cold weather

birds and mammals are able to maintain their relatively high internal
temperature through the insulating action of fur, feathers, and fat, as

well as by suitable physiological adjustments (Scholander et al., 1953).
The remarkable ability of mammals to maintain their normal thermal
levelunder severe winter conditions is exemplified by a varying hare
whose internal temperature was found to be at its normal value of
38C on a day when the thermometer stood at 46C.
Thermal Migrations. Another method for dealing with excessively
high or low temperature conditions is to move out of them. This
method is obviously available only for locomotory forms, and it cannot
be used by the majority of plants. Journeys taken by animals that
enable them to escape from extremely hot or cold situations are referred
to as thermal migrations. Some of these migrations are
relatively
short trips involving movements of only a few meters or even a few
centimeters. Thermal migrations are made on a small scale from ex-
posed positions to the shade to avoid the scorching heat of the desert
and from shade to sun in cold regions. We commonly think of desert
animals as being able to withstand extremely high temperatures. As
a matter of fact most desert animals have become nocturnal in their
habits and thus avoid the heat of the
day. This is true even of char-
acteristic desert reptiles such as the rattlesnake.
Ecologists working
in the desert areas of southern California have found that rattlesnakes
will succumb if forced to remain for more than 15 minutes on the hot
desert soil exposed to the midday sun. The red racer is one of the
few desert reptiles that venture forth regularly in the daytime. Since
this species is one of the fleetest of the snakes, it is able to cross hot
areas and get back into the shade before becoming
harmfully heated.
In less severe environments we are familiar with the short journeys
of terrestrial amphibians into shaded places in hot weather. How-
ever, the chief benefit from a movement out of the sun for salamanders
and other forms with wet, permeable skins may be conserving water
rather than keeping cool. The
excessive evaporation experienced by
an amphibian in an exposed, windy location would cause a chilling of
the body, and, in addition to avoiding water loss, the animal might go
into a sheltered place to keep warm.
The short trips into or out of the water made by frogs, turtles, and
other amphibious forms are familiar to everyone and serve to provide
cooling or wanning for the animals concerned. An unusual tempera-

ture relationis
displayed by the white pelicans that nest on islands of
the Salton Sea, California. In this extremely hot location the eggs
and young in the nest would be killed if exposed for more than about
20 minutes to the intense radiation from the sun. The brooding of
the mother birds acts to keep the eggs and young cool rather than to
keep them warm! At intervals the adult pelicans wet their plumage
by a trip into the water and take advantage of the cooling produced
by evaporation.
Burrowing animals escape excessive heat or cold by short journeys
deeper into their substrata. Soil organisms avoid summer heat by
moving deeper into the earth. Ground squirrels are included in the
desert fauna of southwestern United States but they are rarely exposed
to the extremes of heat in that area. In the hot season these rodents
retire into their burrows where a microclimate of much more mod-
erate temperatures prevails (Table 9). At a later season the same
TABLE 9
REDUCTION IN TEMPERATURE AT INCREASING DEPTHS HENEATH THE SURFACE

OF THE DESERT AT TUCSON, ARIZONA
Air (maximum) 42.5C

Surface (maximum) 71.5
10 cm below surface 41.1
30 cm below surface 29 8
.
45 cm below surface 27.9

150 Temperature
type of migration serves to avoid the severity of winter. Some ani-
mals work their way into rotten logs, others dig deep in the soil, and
species squirm into the mud of
many swamps or pools. This move-
ment of animals into winter quarters often loosely referred to as
is
hibernation, and should not be confused with the structural and
physiological changes often occurring in winter dormancy and de-

scribed by the same term.
Somewhat longer journeys that result either primarily or secondarily
in theavoidance of extreme temperatures are made by larger land ani-
mals. Bear, deer, and other game animals descend from the moun-
tains into the sheltered valleys when the weather gets cold. In the
spring they return to higher elevations and to more exposed situations.

The same type moderate length is seen in the
of thermal migration of
aquatic environment. Many and

fishes other active aquatic animals
leave the shore in the summer when the water has become too warm.
Conversely, other species migrate into deeper water during the
winter to avoid what are for them excessively low temperatures.
Such migrations at contrasting seasons are illustrated very neatly by
two species of flounder that inhabit the coastal areas of New England.
The so-called summer flounder
(Paralichthys dentatus) enters the
shallow water of the bays in June and departs in October. The winter
flounder (Pseudopleuronectes arnericanus), on the other hand, is not
able to tolerate the inshore temperatures characteristic of July and
August. Fish of the latter species are abundant in the upper reaches
of Great South Bay, Long Island, during the winter. As spring comes
on the population moves toward the mouth of the bay, and, when the
summer sun warms the water beyond about 20C, the majority of this
species migrates to the cooler, deeper water outside the inlet (Fig.
5.9). During the autumn months the winter flounder returns once
more into the bay.
Migrations of still greater length are carried out by certain mam-

mals, insects, and birds. Although change of temperature is not the
primary cause of some of these mass movements, escape from climatic
extremes is certainly an important consequence in many instances.
The caribou migrate long distances, and in former times gigantic herds
of bison traveled hundreds of miles across the American
plains from
their winter grounds to their summer feeding areas. The extensive
north-south migrations exhibited by some insect species should also
be considered as thermal migrations in part at least. Most spectacular
of all are the trips made by migrating birds. Considerable evidence
has now accumulated to indicate that change in day length is the
factor that initiates the southward migration of birds at the end of the
breeding season. However, many species of birds inhabiting high

latitudes during the summer would succumb from temperature alone
if
they remained in the far north through the winter. Other species
could perhaps withstand the winter cold but would succumb
through
an indirect action of the heat factor
resulting from the disappearance
of their food supply or from the of the birds to obtain the
inability
larger amount of food necessary to maintain their internal tempera-
ture in winter. For these reasons it seems probable that the avoid-
Mar. Apr,
12'
May June
July Aug. Sept.
Modified from Neville and Perlmulter, 1941

FIG. 5.9. Thermal migration of the winter flounder as revealed by the recapture,
at the points indicated, of fish tagged and released in Great South Bay, Long
Island.
152 Temperature
ance of temperature extremes played an important role in the evolu-
tionary development of the migration habit of birds and of
other ani-
mals, even if this factor does not provide the trigger that sets off the
seasonal journeys each year.
Action within Effective Range
Although the action of temperature extremes may be drastic at

times, the plants and animals of any habitat spend most of their lives
at intermediate temperatures. shall We now consider the action of
the heat factor in the range between the minimum effective tempera-
ture and the maximum effective temperature. Just as with extremes
of heat and cold, the influence of a given temperature within the effec-
tive range
depends upon the thermal conditions to which the species
of plant or animal is adapted. Many tropical, temperate, and polar
species have become attuned to the temperatures characterizing their
respective regions so that many of their life processes go forward at
approximately the same rate in spite of considerable temperature dif-
ferences. For example, the rate of oxygen consumption in certain
lamellibranchs is of the same order of
magnitude for species living at
0C in the Arctic, at 8C in boreal seas, at 12C in the Mediter-
ranean, and at 27C (Thorson, 1950). Seasonal
in tropical waters
adjustment to temperature is exhibited by the sand crab (Ernerita

talpoida) in the Woods Hole area to the extent that its metabolism
at 3C in winter is four times
greater than in summer. As a result
the animal can be active and can grow during the winter when many
other species in the tidal zone become inactive (Edwards and Irving,
1943). The action of specific temperatures on life processes must
be considered for each species separately, and also for geographical
subspecies, but we can delineate certain generalities in

regard to the
role of the heat factor within the effective range.
Effect of Temperature on Biological Rates. The influence of tem-
perature on the rates of biological processes most clearly seen in

is
poikilothermous animals and plants. Although mammals and birds

are definitely affected by the heat factor, the action is
largely indirect
because these warm-blooded forms carry their own temperature
around with them. When acting directly, a rise in temperature of
10C usually causes a doubling or a tripling of the rate of a biological
process, and the process is said to have a Q 10 of 2 or 3 in accordance
with Van't Hoff's law.
If you will take the trouble to listen to the cadence of the crickets on
several evenings, you will observe that the frequency of their chirping
Effect of Temperature on Biological Rates 153
is
higher in warm weather and lower in cool weather (Fig. 5.10).
This relationship was put to use by a blind astronomer who was able
to "read' the temperature by
timing the chirps produced by the crickets
outside his house and applying a formula that he had worked out
(number of chirps in 13 sec -f- 42 rr F).
Another example of the increase in reaction
speed caused by rise in
temperature is
given in Table 10. The data show that the cod egg will
develop at a temperature as low as 1C. Its

speed of development
increases regularly to a temperature of 14C, above which the egg
will no longer survive. The relationship with temperature in this
30
f
300
.20
200.2
a.
g
10
100 140 180 220
Stimulations per minute
Fie. 5.10. Relation of stridulution rate of tree cricket Oecanthus to temperature

and also to pitch. The note C is "middle C." (Modified from Matthews, 1942,
Science. )
TABLE 10
RELATION BETWEEN TEMPEKATIWE AND RATE OF DEVELOPMENT OF

OF A C()LI)-WATEIi AM) A \VAKM-WATKK FlSH
Cod Mackerel
2C No development (ecological zero) 3C No development
I Hatch in 42 davs 10 Hatch in 207 hours
150
o 18 15 10,5
8 13 18 70
10 10.5 20
12 f) 7 50
14 8.. >
r
No development
No development
154 Temperature
and in most other biological processes is not linear, and sometimes
varies in a complicated way. For the cod egg a rise of 4 from

1Cto -f* 3C
produces a doubling in the rate of development, but
a similar increase of 4 to from 10C
results in only a slight14C
acceleration. A
temperature of 10C, which is the minimum at which
the mackerel egg will develop, is near the maximum temperature for
the cod egg. As the temperature is raised from 10C to 21C the rate
of development of the mackerel egg is increased until at the upper
value it will hatch in just over 2 days.
An increase in temperature beyond a critical
point may cause a
reduction in the rate of some processes in certain species. The
life
growth of the hypocotyl of the pea seedling shown in Table 11 illus-
TABLK 11
RELATIOX BETWEEN TEMPERATURE AND THE GROWTH OF THE PEA SEEDLING

As INDICATED in THE ELONGATION OF THE HYPOCOTYL
Temperature Growth
C Mm per Day
14.1 5
18.0 8
23 5 . 30
"2(> 6 54
28 5 . 40
33 ,5 23
3(i . 5 1)
trates this point. As temperature increases from 14.1C to 26.6C

the amount of growtli recorded each day steadily increases, but at
higher values the daily elongation of the hypocotyl becomes less. In
contrast to the growth in fish eggs, the pea seedling continues to
elongate after the temperature has been reached at which growth is

the fastest. This fact does not moan that increase in temperature has
a negative effect on a reaction. Growth, like most other biological
processes, is the result of a number of biochemical reactions going on
simultaneously. In the present instance the rate of some inhibiting

reaction may have been increased more by a rise in temperature than
the reaction for elongation with the result that after a certain point
the rate of growth becomes less.
In other instances temperature may have little or no effect on the
rate of biological processes. Between certain limits the incubation
period for many reptiles is
only slightly influenced by thermal changes
in the surroundings. The diamond-backed terrapin, for example,
undergoes normal hatching between 18C and 33C, but within that
Optimum Temperature 155
range no regular correlation exists between temperature and time of

development. On the other hand, some life
processes go forward
only within a very narrow thermal range. Everyone who has raised
chickens knows that it is
impossible to persuade the eggs to hatch
more quickly by increasing the temperature. Although the hen's egg
has no temperature control of its own, it will hatch
only if maintained
between 40C and 41C and only after 21 days of incubation.
Optimum Temperature. The foregoing discussion of the influence
of temperature on the rates of
biological processes leads to a con-
sideration of theoptimum temperature. However, the concept of
the optimum in ecology is a slippery customer unless the life
process
concerned is
specified. The optimum temperature may be considered
that value at which a certain process goes on the fastest. But the tem-
perature for the maximum rate often varies considerably for different
processes within the same organism and also for the same process at
different stages in the life cycle.
Among plants in the temperate zone the optimum temperature for

germination usually differs markedly from the optimum value for the
fruiting process, and the optimum heat condition for photosynthesis
may occur at a value different from either of the other two. Similar
variations in thermal relations are found among animals. The optima
for the various developmental stages are frequently found at values
widely different from the optimum for the adult. The eggs and larvae
of terrestrial species usually require a higher temperature than the
adult stage, but notable exceptions are provided by many animals as,
for example, by the corn borer, and the eastern brook trout. The trout
spawns in October or November, and the eggs can develop at or 4C
less. The optimum temperature for the development of trout eggs is
8C. The adult trout, on the other hand, do not feed much and there-
fore do not grow until the water is warmer than 10C. The optimum
for the growth of the adult falls between 13C and 16C. These ex-
amples are sufficient to indicate that no one point on the thermal scale
can be designated as the optimum for the entire growth of an organism,
but that optimal values differ according to the life
stage of each
species.
The most favorable temperature for survival often differs consider-
for growth, reproduction, and other
ably from the optimum value
life
processes. Mackerel eggs develop at the fastest rate at 21C, but
in the same investigation the survival of the eggs was found to be best
at 15C. Other experiments showed that the life span of the common
water flea, Daphnia magna, is
greatest at 8C (Table 12). In con-
trast, the shortest time for the production of the first batch of
young
156 Temperature
Daphnia occurs at 23C, and the optimum from the point of view of
the largest number of young produced is found to be at 18C (Mac-
Arthur and Baillie, 1929). The ecological effect of these divergent
optimal values will depend upon whether they are applied to the
individual or to the population. The individual Daphnia lives longest
at one temperature, but the population as a whole grows the fastest
at another temperature. The optimum temperature for a species can
therefore be only a general concept. The optimum is best thought of
as a range of temperatures, and it is the range within which the or-
ganism as a whole functions best.
TABLE 12
TEMPERATURE RELATIONS OF THE CLADOCERAN, Daphnia magna
Average Length Time for Production Average Number of

of Life (days) of First Young (days) Young Produced
8C 108
10 88
1* 16 6
18 40 8 24
23 6 9
28 26
Other Effects of Temperature. In our discussion of temperature

thus far we have dealt chiefly with growth and survival. Many other
influences of this factor are of importance in the ecology of animals
and plants. A drop in temperature ordinarily means a decrease in
activity, especially among the cold-blooded animals and plants. As
winter comes on, insects and other invertebrates ordinarily become
dormant. In the spring on land the thermometer usually must climb
above 8C before cold-blooded animals become abundantly active.
The snow flea, Collembola, forms an interesting exception to the fore-
going generality. These small black insects are often found swarming
on the surface of the snow among the trees in February or March.
Evidently the animals can absorb enough heat from the strengthening
spring sun in their protected microclimates at the base of trees to be
active. In the aquatic environment many animals are adapted to
carrying on a very active life at temperatures only a few degrees

above 0C. No one who has hooked a salmon in a northern stream
will claim that ahigh degree of activity is impossible for cold-blooded
animals at these lower temperatures.
Certain behavior patterns are influenced by the heat factor. Some
of the more primitive animals exhibit a thermptaxis, that is, an orienta-
tion toward a source of heat Ticks are aided in locating their warm-
Other Effects of Temperature 157
blooded hosts by a turning reaction to the heat of their bodies. Rat-
tlesnakes, copperheads, and other "pit vipers" can detect the presence
of mammals or birds that
may be
only a few degrees warmer than
the surroundings. Oriented by the heat radiation, these snakes can
strike accurately at their
prey even in the dark. Other behavior
reactions are influenced by heat conditions. When the temperature
drops below 5C,
the leopard frog is stimulated to burrow into the
mud bottom pond. At the same temperature sunfish crowd
of its
together in aggregations but, if the water is warmed to 8C, the fish

tend to swim about separately. Snakes similarly gather in aggrega-
tionsand form balls in their retreats
among the rocks when cold
weather arrives in the autumn.
Temperature, as well as moisture, light, and other factors, has long
been known to affect the coloration of some animals. In warm, humid
climates many mammals, birds, and insects, tend to be more mclanic,
that darker in color than races of the same species living in cool,
is,
dry climates. This generality, already referred to as Gloger's rule, is

well recognized but many exceptions are encountered, and the relative
influences of heat and moisture are not known. In some instances
coloration seems more affected by genetic selection than by the direct
action of climate. The possibility of hereditary transmission of heat-
color relationships is demonstrated by experiments showing that the
temperature to which pupae are exposed will affect the wing color of
butterflies even to the second generation. This whole subject is con-
troversial since observations on various species in different areas do
not agree. Coloration is evidently controlled by interactions between
environmental and hereditary factors, but a more exact understanding
must await further study.
The temperature factor is also known to affect the absolute size
of many animals as well as the relative proportions of certain parts.
The general fact that among birds and mammals the same species
attains a greater body size in cold regions than in warm regions and
that among closely related species the larger ones inhabit the colder
climates is known as the Bergtnann principle. Poikilothermous ani-
mals, as exemplified particularly by reptiles and amphibians, exhibit
the reverse relationship since they tend to be smaller in colder cli-
mates.
The related observation that extremities, such as the tail, ears, and
legs, of
mammals are shorter in colder climates has been delineated
as Aliens rule. Since both these generalities apply to warm-blooded
forms, they are probably related to the difficulty of retaining heat at
low and the desirability of losing it at excessively high
temperatures
158 Temperature
temperatures. Animals with large bodies have surfaces that are rela-
tively smaller in relation to their masses than small animals. Smaller
extremities expose smaller surfaces (Fig. 5.11). Since heat is lost
through the surface, the smaller the area of an animal's skin, the more
easily may the animal maintain its

temperature in cold weather.
Conversely, the development of extremities with large areas aids heat
loss and evaporation in hot climates. These simple ecological rela-
tionships, in addition to underlying Bergmann's principle and Allen's

rule, undoubtedly account in part at least for the fact that no extremely
small mammals or birds exist, that is, as small as the majority of insects.
Fie. 5.11. Head of (left) Alopex lagopus, (center) red fox, Vulpes
arctic fox,
vulpes, and showing gradation in size of ears.

(right) desert fox, Megalotis zerda,
(Reprinted with permission from Hesse, Allee, and Schmidt, Ecological Animal
Geography, 1951, John Wiley & Sons, New York.)
A very specialized influence of temperature on morphology is the

apparent control it exerts on the number of vertebrae in certain species
of fish a
relationship known as Jordan's rule. Cod hatched off New-
foundland where temperature ranges between 4 and have 56 8C
vertebrae, whereas cod hatched east of Nantucket in temperatures
averaging 10 to 11C possess only 54 vertebrae. The relationship
is
brought about through the control by temperature of metameric
segmentation at an early stage of development. Incidentally, the
fact is of use to fishery biologists in ascertaining the origin of popula-
tions among those species of fish that exhibit this geographical dif-
ference in structure.
The temperature of the environment controls egg type and sex
ratio in certain animals. Under moderate heat conditions Cladocera
produce parthenogenetic eggs, and these hatch usually into females.
Under ordinary circumstances few if any males exist in the popula-
tion, but their production and the appearance of "winter" eggs is
favored by extremes of temperature. The winter egg of the Clado-
Mode of Temperature Limitation 159
cera, as described earlier, is able to withstand desiccation and freez-

ing but, since it is a sexual
egg, it must befertilized by a male before
it can develop (cf. Fig. 2,11). In Moina macrocopa, when the tem-
perature drops below about 14C, or rises above 30C, males appear in
the population and sexual eggs are
produced. This control of egg type
and sex is attuned to the fact that in
ratio
temporary pools the oc-
currence of very high temperatures often
precedes the drying up of
the pond in summer. At the other end of the
year a drop in tempera-
ture will presage the
possibility of freezing and the necessity that the
animal go into a life structure in which the severity of winter can be
endured.
ACTION OF TEMPERATURE ON DISTRIBUTION
Mode of Temperature Limitation
In early attempts to describe the ecological action of tem-

many
an effort was made to correlate
perature geographical distribution
with mean temperatures. Although we see many average tempera-
tures in print, we never encounter one in nature. A distinction must
be made between the use of a temperature value as an index and the
action of temperature as an agent controlling distribution. The limits
to the spread of a population are always set by ecological factors act-
ing on the individuals making up the population. In most situations

the individual is not subject to a temperature equal to the average
temperature of the region for more than a few hours of the day or
a few days of the year.
In eastern Massachusetts the mean annual temperature is about
7C but on many days in summer the thermometer stands near 24C
and in the winter near 10C with extreme records far above and
below these values. As a consequence the fauna and flora of the
region are very different from what they would be if the temperature
remained in the neighborhood of 7C all the year round. In the
latter event the local biota would be more like that of the Aleutian
Islands or of South Iceland where the mean monthly temperature

varies only from about 10 to 1C with an average of about 4C.
We may conclude that the mean annual temperature is not the chief
aspect of this factor which controls distribution, but a more pene-
trating analysis of the
action of temperature must be made.
Careful consideration of the various thermal influences shows that
for the permanent life of a species in an area the following tempera-
ture requirements must be met:
160 Temperature
a. The temperature must never be so high or so low at any time as
to kill the organism.
b. The temperature must be high enough, or in some instances low
enough, for a sufficient period to permit the reproduction and
growth of the species.
Limiting minimal temperatures for survival ordinarily occur in

winter, and minimal heat required for reproduction and growth will
be found in summer. Either or both of these influences determine
boundaries of distribution toward the poles or higher altitudes.
Limiting maximal temperatures for survival commonly occur in sum-

mer and minimal chilling necessary for reproduction and growth will
be found in winter. These influences determine boundaries of dis-
tribution toward the equator or low altitudes. On the basis of the
foregoing generalities Hutchins (1947) has delineated four basic types

of zonation in geographical distribution and has discussed them par-
ticularly in relation to the ranges of marine animals. As we shall see,

in these common types of temperature zonation as well as in more
irregular situations, failure to meet the thermal requirements listed
above may occur at any stage of the life cycle and at various seasons
of the year.
Control by Extremes. The
simplest type of control of geographical
distribution by temperature extremes is that in which the polar limit
of the range of a species is determined
by the minimum temperature
in winter and the equatorial limit is fixed by the maximum summer
value. Many illustrations of this type of control will occur to the
reader and can be found in treatments of plant and animal geography.
Sometimes the lowest temperatures tolerated by tropical species are
encountered far to the south (in the northern hemisphere), and the
highest temperatures tolerated by arctic species are found well to the
north. There is evidence, for example, that the southward ( and east-
ward) distribution of three flowering alpine perennials in Scandinavia
is correlated with the maximum temperatures in summer as indicated
in Fig. 5.12. Although the exact causal relationship in such instances
is not known, possible that the protoplasm of the northern species
it is
is
adversely affected, that respiration is increased faster than assimila-
tion, or that temperatures are too high for the proper transport of as-
similated products (cf. Went, 1950).
Intensive study of certain species has revealed the fact that control
by extremes may involve unexpected complications and that other

aspects of temperature influence may be
in operation
simultaneously.
A multiphased action of temperature will probably be found in many
Control by Extremes
Dakl, 1951
FIG. 5.12. Locations of the most southern and eastern records in Scandinavia
of three flowering alpine perennials and the maximum summer temperatures,
x Lactura alpina, correlated with 29 C isotherm.

o Ranunculus platanifolius, correlated with 27 C isotherm.
Saxifraga foliolosa, correlated with 23C isotherm.
162 Temperature
more species when they have been more closely investigated. Ac-
cording to Iversen (1944), for example, the northern and eastern

limit in Europe for thegrowth and reproduction of the ivy, Hedera
helix arborea, is determined by the minimum winter temperature in
situations where the average temperature for the coldest month is
1.5C or below, but in areas with less severe winters the northern
limit is
by fixed the occurrence of sufficient summer heat as indicated
by the average temperature for the warmest month (Fig. 5.13).
-1-10
11
o?
12
Pert
14
i
w i5
ro
X 17
19
|
21
22
6 5 4 3 2 1 0-1-2-3-4-5-6-7-8-9 -10
Average temperature for January, C
Iversen, 1944
FIG. 5.13.
Temperature conditions at localities in Europe in which the occurrence
and condition of the ivy, Hedera helix f. arborea, have been investigated.
Ivy sets fruit regularly.

O Ivy sets fruit occasionally,
o Ivy occurs but does not set fruit.
x Ivy does not occur.
The harmful action of high temperature is not necessarily limited

to midday hours in summer nor that of low temperature to the coldest
night in winter. We
have seen in the previous chapter that winter
is sometimes due to the
killing of vascular plants excessively warm,
dry wind during periods when the ground is frozen. For many plants
the danger of injurious low temperatures occurs in the spring after
Control by Extremes 163
germination, and at that time

killing will result from temperatures
that are not nearly as low as those which the
plant successfully with-
stood during the winter while it was in a dormant condition. An un-
expected temperature relationship is exhibited by certain plants in
northwest Europe that can extend their
range farther poleward in the
interior of the continent than
they can along the sea coast. The ex-
is that the milder winter and earlier rise of
planation temperature
near the ocean stimulates the
plants here to earlier germination, but
since the spring temperatures rise more
slowly the seedlings may be
killed by late frosts.
Among those species whose geographical range is limited by tem-

perature extremes we find a great variation as to the
part of the life
cycle that is chiefly concerned, The limitation is
brought about by
the most susceptible stage; the boundary of distribution is determined
by the weakest link in the life cycle in respect to temperature tol-

erance. Other life
stages may extend temporarily beyond the area
of permanent existence.
We often think of the youngest organisms as the most sensitive to
harmful influences in the environment, and this is indeed sometimes
true in respect to temperature. The early stages of development

often exhibit a very narrow range of temperature tolerance and may
thus represent the "heel of Achilles" for the species as far as damage
from excessive heating or chilling is concerned. For example, fish
eggs characteristically a greater susceptibility to harm from

show
temperature than adult fish of the same species. The eggs of birds,
although a special case because birds are homoiotherms, will tolerate
only an extremely narrow fluctuation in temperature and frequently

die as the result of unusually cold or hot weather. In some animals,
on the other hand, the egg is far more resistant to temperature ex-
tremes than any other stage. In other species the most resistant stage
is found later in the life cycle and may be represented by a cyst or
pupa. The seed of the higher plants similarly represents a tough,
resting stage and one

that is
characteristically able to withstand a
range of temperature far greater than that tolerated by

the adult plant.
can often survive a frost that would critically injure the re-
Sprouts
productive tissues of the mature plant.
Very frequently the greatest susceptibility to heat or cold damage
lies in the intermediate stages of growth. The period after the egg
has hatched or after the seed has germinated is a critical one in some
species; represents the stage at
it which pioneering individuals at the
margins of their range will be
killed off by adverse temperatures.
The fate of copepod populations carried by currents each year from

164 Temperature
the Gulf of Maine into the Bay of Fundy serve as an illustration of
this situation. Fish and Johnson (1937) report that adult copepods
brought into the Bay of Fundy find the temperature tolerable, breed
successfully, and produce eggs that are viable under the existing
thermal conditions. The temperature remains too low for the ensuing
larval stages, however, with the result that the new
generation is
eventually killed off. The is
consequently re-
copepod population
duced until the next year when it is
again renewed by currents from
the Gulf of Maine. Sometimes the harmful action of temperature is
quite indirect. Although the free-swimming life of the oyster, Cras-
sostrea virginica, is extended from 7 days at 25C to 21 days below
20C, the animal develops perfectly well at the latter temperature.
The principal significance of low temperature for the larvae is that it
prolongs pelagic life and hence increases the period during which
the larvae fall easy prey to enemies (Thorson, 1950).
In some species the adult stage is more sensitive to extremes of
temperature than any of the younger stages. Excessive heat or cold

may reduce the vitality of the plant or animal as a whole, or it may
cause damage to specific tissues or functions. The organs of repro-
duction are frequently the most susceptible. For example, the north-
ward peach trees in the United States is limited by
distribution of
damage buds by late frosts in the spring, although the other

to fruit
parts of the plant remain quite uninjured. The foregoing discussion

will suffice to illustrate the fact that extremes of heat or cold may kill
off animals or plants at whatever season the most vulnerable stage of

development occurs.
Control by Need for Minimum Amount of Heat. In order that a
locality, the temperature at some

species maintain itself in a given
season must rise above the threshold, that is, above the minimum
at which the vital processes of the organism can go on, and it must
remain above that value for a period sufficient to allow growth and
reproduction to be completed. The great variation in the length of
the frostless season within the United States is indicated in Fig. 5.14.
Many organisms require temperatures considerably above freezing

at someseason, but occasionally we find instances of plants or animals
that can get along with remarkably little heat. Lichens were found
growing on Antarctic nunatacs (rocky peaks protruding through the

ice) where extreme cold prevails for much of the year and the tem-
perature rises above 0C
on very few days. Similarly, a rich and
varied marine invertebrate fauna and associated flora are known to
grow and to reproduce along the coast of Greenland where the water
temperature never exceeds 0C.
Control by Need for Minimum Heat 165
Above the thermal threshold
generally true that, the higher the
it is
temperature, within limits, the shorter the period needed for the same
amount of development. This fact led to the belief that the heat re-
quirements for the growth of an organism from its youngest stage to
maturity, or for the completion of any other life process, might be cal-
culated quite
mechanically as a "heat sum" (or "thermal constant")
expressed as "degree days" or "degree hours." The heat sum was
obtained by finding the threshold temperature for the process con-
cerned and summing the differences in degrees between this value
and the average temperature on each day until the completion of the
process. Thus the heat sum required for the flowering of corn plants
in Ohio was found to lie between 660 and 1050 degree days, basing
the calculation on the number of centigrade degrees above 6C
recorded for the temperature each day. The lengths of the life his-
tories of many insects have been shown to depend upon a relatively
constant temperature sum over a considerable range of temperatures,
and hence of velocities of development (Allee et al., 1949, Ch. 6).
< 90 days
U| 90 - 120 J~\ J^^^^^
E3 -150
120 EZ3210-240
~~~
\
jF
33 150 -180 V V
~~\ 180-210
Data from Atlas of American Agriculture, U. S. Department of Agriculture, 103
FIG. 5,14. Average number of days between last killing frost in spring and first
killing frost in fall.
with
Although the results of temperature summation agree generally
to mature
the fact that a given type of plant or animal requires longer
in a cold than in a warm climate, closer scrutiny shows that, in many
are far more involved than
species at least, temperature dependencies
166 Temperature
are indicated
by heat sums. In the first place the heat sum for a
process is constant
only for that range within which there is direct
proportionality between growth rate and temperature. Lack of such
proportionality is usually found near both the upper and the lower
limits of tolerance. Indeed, as mentioned earlier, above certain
"optimal" temperatures the rates of many metabolic processes ac-
tually decrease. Furthermore, since heat sums are based on average
daily temperatures, they do not take into account the possible special
effects of the maximum or minimum
daily values, night temperatures,
or differences between diurnal and nocturnal
temperatures.
The critical importance of these more detailed aspects of tempera-
ture in controlling life
processes is discussed in greater detail by
Daubenmire (1947) particularly in relation to land plants. Some
plants exhibit normal responses only when grown in temperatures
that fluctuate with a regular diurnal a phenomenon known as
rhythm
thermgjyyiodism. And to make matters more complicated, the pre-
cise action of
temperature may be further modified by the intensity
of light and the length of day. Went ( 1950) points out that no heat
sum can account for the ripening of the tomato. In addition to
proper light intensity, length of day, and day temperature, the tomato
plant sets seed only if exposed to a series of five or more nights dur-
ing which the temperature remains above 15C. These findings are
compatible with the general principle of minimum heat requirements,
but they show that in certain land plants at least the matter is far
more complicated than first
supposed.
Control of the geographical distribution of many animals can sim-
ilarly be related to the need for a minimum amount of heat. The
American oyster as an adult can survive at temperatures from 32C
down to the freezing point of salt water. A temperature above 15C,
or higher in some races, is necessary for spawning to take place in this
species, and good development of the larvae requires water above
18 or 20C. The adult lobster can live at temperatures ranging from
about 17C down to 0C, but breeding will take place only in water
warmer than 11C. The lower maximum temperature tolerated
means that the lobster cannot range to the south as far as the oyster
and the lower heat requirement for lobster reproduction produces a
marked difference in the breeding range of the two animals. For a
complete study of the effects of temperature on distribution the ther-
mal conditions in each bay and inlet should be investigated, but the
range of temperature along the coast is sufficient to illustrate the gen-
eral influence of this factor.

A glance at Fig. 5.15 will show that north of Cape Cod coastal
Control by Need for Minimum Heat 167
temperatures are generally lower than 18C until the Gulf of St. Law-
rence is reached where
surprisingly warm conditions occur in summer
in the shallow areas around Prince Edward Island.
Temperatures in
the central part of the Bay of
Fundy are prevented from rising above
11C by the intense tidal stirring, but the stratified water along the
outer coast ofNova Scotia becomes considerably warmer than this.
Correlated with these temperature conditions we find that in New
England oysters grow naturally in commercial quantities only as far
north as Narragansett Bay but "seed"
oysters may be transported to
Cotuit and elsewhere on Cape Cod, where they are fattened for mar-
Modified from Ackerman, 1941

and oysters
FIG. 5.15. Regions of abundant occurrence of lobsters (small dots)
(large dots) in relation to August
surface temperatures as indicated by the loca-
tion of the commercial catch in 1934.
ket. Oysters again become sufficiently

numerous to market in the
littoral zone around Prince Edward Island and in certain warm bays
of Cape Breton and northeast New
Brunswick. Lobsters, on the
other hand, breed abundance
successfully in all
along the coast of
Maine, on the outer coast of Nova Scotia, and into the Gulf of St.
Lawrence, but these shellfish are not taken in commercial quantities

within the Bay of Fundy. Although lobsters wander into the Bay
of Fundy and some of them grow to a large size, the lack of water
168 Temperature
of sufficient warmth prevents the establishment of a native breeding
population despite the fact that the animal's range extends much
farther north in situations where the heat requirements are met.
The northward distribution of turtles in inland waters is similarly
determined not by the extremes of temperature but by the need for
the minimum amount of heat for the completion of the life cycle.
Since turtles burrow in the mud during the winter, the extreme mini-
mum air
temperature at that season is of no consequence. But these
reptiles require the heating action of the summer sun to incubate their
eggs. The snapping turtle, for example, lays its eggs in the sancl dur-
ing the month of May, and sufficient heat must reach them during the
summer in order for the young turtles to hatch out successfully in
September. The northerly distribution of turtles is correspondingly
limitedby this long heat requirement to the upper or central part of
the United States, but snakes can extend their range well over the
Canadian border because heat for the incubation of their eggs is
needed for only a short period.
Differential control of geographical range is also illustrated by the
distribution of frogs, whose temperature requirements are widely di-
vergent. Some species of frogs seem to need little more heat than
that necessary to thaw out their ponds just long enough for reproduc-
tion. The eggs of the wood frog, Rana sylvatica, develop at a tem-
perature as low as 2.5C, and the larval stage requires only 60 days.
This species extends northward in Canada to the mouth of the Mc-
Kenzie River. The pickerel frog, Rana palustris, on the other hand,
must have a temperature of 7.5C and a period of at least 90 clays for
its
development. The range
of this species is correspondingly limited
to the latitude ofJames Bay. In contrast, a third species, Rana clam-
itans, will not develop until the temperature exceeds 11C, and its
range extends only slightly above the southern boundary of Canada.

Control by Need for Chilling. Some organisms require a certain
period of cold weather for their life
cycles to be completed. No one
would plant tulips or crocuses in June and expect them to bloom in
August. These plants come into flower only after the bulbs have
passed through a winter period of low temperature. The dormant
buds of certain fruit trees and berry plants similarly require a period
of chilling before they will flower successfully. Experiments have
shown, for example, that some types of blueberries require an ex-
posure of 800 hours to temperature below 7C
before the dormant
buds will develop. In other species the initial formation of flower
buds will take place only at low temperatures.
The seeds of some plants must be chilled under moist conditions
before they will germinate In other species, such as winter
properly,
wheat, satisfactory development takes place only if a period of chilling
occurs during or after germination. It has been discovered that the
seeds of certain plants must
undergo two successive cold intervals
before the seedling will
grow (Barton, 1944). This requirement
means that under natural conditions
development of the plant can go
forward only after the second winter.
In nature all species
requiring low temperatures are limited in their
altitudinal distribution down the sides of mountains and in their
latitudinal extensiontoward the equator. This cold requirement also
imposes serious restriction on the successful cultivation of temperate
varieties of fruit trees in warm climates. Although little is known of
the physiological mechanisms upon which the need for chilling de-
pends, the empirical knowledge is put to practical use through

vernalization, that is, the artificial
exposure of seeds to cold. Such
procedure allows certain crop plants to be grown farther north and
also farther south than would otherwise be possible.
Another need for low temperatures results from the varying effect
of this factor on anabolic and catabolic processes. We have seen that
sufficient heat must be available to plants and animals so that the
growth and reproductive processes may make up for the destruction
and the death of individuals. As temperatures rise,
of tissue materials
most vital reactions are accelerated, but sometimes at very different
rates. catabolic processes are speeded up disproportionately, the
If
organism will suffer rather than gain from high temperatures. Above
certain temperatures the respiration rate of most plants becomes
higher than the rate of photosynthesis. Food manufacture is conse-

quently curtailed, and growth, reproduction, and the accumulation
of food reserves may be inhibited. The range of plants toward low
latitudes and low altitudes and their seasonal activity are limited to
areas and periods when temperatures remain low enough to permit
a favorable photosynthesis-respiration relationship.
Although the need for low temperatures during part of the life cycle
has been extensively studied in relatively few animals, there is little
question that failure to meet this thermal requirement limits the dis-
tribution of some species. Desert mammals and insects that are
forced to aestivate by high temperatures would not be able to con-
tinue active life if some portion of the year did not have a cooler cli-
mate. The gemmules of sponges hatch at a much lower temperature
and in a much shorter time after a period of freezing. The eggs of
170 Temperature
the white mayfly, Ephoron album, must be chilled to within a few de-
for several weeks (or subjected to a few days of
grees of freezing
actual freezing) for good hatching to be assured. As a result, this
insect cannot extend its in the eastern part of the United States
range
farther south than 40 N latitude.
The eggs of the Australian grasshopper will not continue growth
beyond a certain stage unless they are exposed to low temperatures

within the range that will permit diapause development to proceed
rapidly (5-13C), and later, to

high temperatures (17-35C) which
to go forward (Fig. 5.16). This dependency
permit morphogenesis
20
80h / V^ A D / 1 16 S
-148
12 J
0)
^0
|
40h / \ \ / -H8
I
6 f
20h" >T\ -J4 1
5 10 15 20 25 30 35
Temperature, *C
From Andreu'attha, 1952, copyright Cambridge Univ. Press
FIG. 5.16. The influence of temperature on (A) diapause development and (B)
Austroicetes
post-diapause morphogenesis in the embryo of the grasshopper,
cnwiata. Solid line: eggs from South Australia;
broken line: eggs from West
Australia.
nicely attuned to the normal life cycle

on chilling is of this grasshop-
per. Since the insect reaches the diapause stage at the beginning of
winter, it is endure best the severest cold weather
in the condition to
of the year and also to take advantage of low temperatures for the
completion of diapause development.

Other species in which dia-
occurs later of the life history exhibit a different
pause during stages
conditions of their habi-
timing in their adjustments to the ecological
tat. The eggs of the moth, Euproctis, hatch during July, and dia-
until the larvae have reached the second
pause does not take place
or third instar. By the time the animals enter diapause autumn has
arrived and the species overwinters in the larval stage.
In New England the gypsy moth lays eggs in summer, but develop-
ment will
normally start again only after chilling, or not until the fol-
lowing spring. This timing not only fits the life cycle of the moth
but also is most fortunate for the survival of the
vegetation in areas
where this pest is abundant. If continued
reproduction throughout
the late summer and autumn months,the trees upon which the cater-
pillars depend would
for food have no opportunity to recover from
the spring and early summer inroads of Even in species
feeding.
without a recognized period of dormancy distribution toward warmer
regions may be limited by the fact that warm conditions inhibit cer-
tain necessary
processes long before temperatures are reached that
would be immediately lethal. The southern ( equatorward ) bound-
ary for the barnacle, "Bdlamis balanoides, appears to be determined
by the isotherm for a minimum surface temperature of 7.2C (Hutch-
ins, 1947). Evidently this barnacle requires a certain period of low
in order to
temperature complete reproduction successfully; this
its
conclusion is supported by the fact that the species breeds during the
summer in the northern part of its range, but breeds only during the
winter in the southern areas of its distribution.
The foregoing consideration of the limitation of distribution by
thermal requirements has called attention to the complexity of the re-
lationships involved. Temperatures that are harmful, or that are

necessary, differ widely with species, age, and physiological condi-
tion. Resistant stages of a species may be found far beyond the
boundaries of the area within which all parts of the life cycle may
be carried on. For marine bottom invertebrates, for example, Thor-
son (1950) has delineated three areas of distribution: (1) an exten-
sive area throughout which the adult animal may live at least vegeta-
tively; (2) a smaller area within the first in which the animal will
ripen its sexual products and spawn; and (3) a still smaller area
within the second in which the embryos and larvae will develop
successfully. Other types of animals and plants have different pat-
terns of temperature sensitivity at different life stages. As we have
seen, the effect of temperature may vary for different life processes
in the same or even for the same process in different parts
organism
of the organism. Each species of animal or plant can live only in
those regions in which the temperature pattern is tolerable and ade-
quate for the needs of the species.

The organism is restricted to
habitats in which the annual, seasonal, and diurnal fluctuations in
times and in such magnitudes as to allow
temperature occur at such
all life processes to be completed but do not occur so as to cause
serious harm. The intriguing interdependencies outlined here cer-

tainly need
and invite further investigation by ecologists.
172 Temperature
Results of Temperature Limitation
One far-reaching consequence of the action of temperature in the

various ways that have been reviewed is to impose a north-south
gradation on the distribution of many animals and plants both in
water and on land. The ranges
of species that are controlled pri-
marily by temperature, rather than by moisture or some other ecologi-

cal factor, have boundaries tending to run generally east and west.
Latitudinal zonation is considerably modified, however, by tempera-
ture anomalies due to currents in the ocean, and to differences in alti-
tude, proximity to oceanic areas, and other influences on land. Across
the United States, for example, less severe cold is experienced along
the Atlantic and Pacific coasts than in the center of the continent, and
cold weather occurs farther south in the mountain areas. The ranges
of species correspondingly extend farther north along the coasts
many
but are restricted to more southerly regions in the Rockies and in
the Appalachians. On the other hand, in northwestern Europe
species for which the northern boundary is determined by summer
warmth range farther north toward the interior of the continent be-
cause the temperature in summer is higher there than on the coast.
There are no temperature boundaries to which the ranges of all
plants and animals conform. The complex influence of thermal con-
ditions must be investigated for each species separately before gen-
eralities applying to groups of animals and plants can be established.
As we go north or south species drop out irregularly and their ranges

overlap. The pines of the north, for example, extend into the realm
of the palms of the south. The live oaks, typical of southern United
States, overlap in their distribution the white oaks, whose center of
distribution lies farther north. The quail extends its range northward
into New Hampshire, but the grouse, a similar type of bird common
in Canada, spreads southward as far as Virginia.
The same sort of overlapping occurs with aquatic organisms. Of
40 species of marine invertebrates common along the eastern coast of
the United States, 16 have their northern boundary at Cape Cod, 14
have their southern boundary at the same point, but 10 species occur
on both sides of the Cape. In the study of geographical distribution
as influenced by temperature, we see again the necessity for the ana-
lytical approachfor dealing primarily with the mechanism of control
of individual species.
Special Cases of Common Boundaries. Only in special instances

do a large number of plant and animal species share a common ther-
Special Cases of Common Boundaries 173
mal boundary for their distribution. Under unusual circumstances
the horizontal
temperature gradient may change abruptly. At the
northern edge of the Gulf Stream, for
example, the temperature may
drop as much as 5C
within a few miles-an extreme thermal change
for the oceanic environment. We are not surprised that
species many
inhabiting Sargasso water have northern boundaries and species in-
habiting continental slope water have southern boundaries at the Gulf
Stream. Similar sudden
changes of climatic conditions occur on
land in relation to unusual
topographical changes, such as an abrupt
in altitude at the
change edge of a plateau. In these situations a
considerable number of plants and animals share common boundaries
based primarily upon the temperature factor.
Frost Line. The frost line on land constitutes another special
thermal situation but one in which there is no sudden change in tem-
perature geographically, The fact that the tissues of animals an<}
plants in active condition freeze not far below results in the0C
occurrence of a boundry of general importance at the frost line.
All plants and all animals that are not frost hardy and that do not
have modes of escape are limited in their poleward and altitudinal
distributionby the harmful results of free/ing. At the same time
any plants or animals dependent upon these frost-sensitive species
will drop out. The palm trees are an example of plants that are
unable to stand a continued frost. In North America the natural
range of the true palms does not extend north of the central part
of Florida, the Gulf coast, and the southern part of California.
Smaller plants and animals associated with the palms are necessarily
limited in their distribution to essentially the same boundaries.
Tree Line. Other abrupt changes in the fauna and flora commonly
occur where one major type of vegetation gives place to another, since
the dominant vegetation influences the presence of many subordinate
species of plants and animals.

An outstanding instance of common
boundaries due to the dependence of many species on the type of
vegetation is seen at tree line. Here again there is no sudden
change in the distribution of temperature. When
thermal and other
conditions have been modified to such a point that trees no longer can
grow, an change in the whole ecological situation takes

important
place. In any forested region trees have a profound controlling in-
fluence on the subordinate plants and on the associated animal life.
Thus the environmental situation that limits the trees indirectly sets
boundaries to the distribution of many other species as well. Tree
lines dependent in part at least on temperature occur across the con-
tinents at the poleward margins of the forests and also on mountain
174 Temperature
slopes at the upper limits of forest vegetation. We
may consider first
the northward boundary of timber in Canada as an example of a
continental tree line.
A traveler going north in the eastern part of North America passes
from the southern region of palm trees into the belt of southern
pines and then into the "central hardwoods," dominated by white oaks
and hickories. The central hardwoods extend into southern New
York, Connecticut, Rhode Island, and maritime Massachusetts. In
east central Massachusetts and up the valleys in New York, Vermont,
and New Hampshire the vegetation changes to the "transition forest,"
in which the maple, red oak, ash, and black birch are the most
abundant species. In northern New York, central Vermont, and
New Hampshire the forest becomes the "northern hardwood" type,
with maple, beech, and white and yellow birch predominating.
When the traveler reaches northern Maine and the corresponding
parts of Ontario, he finds himself in the spruce forest. Here red
spruce, firs, and white and yellow birch are the common trees. Still
farther north, about at the southern edge of Hudson Bay, the trees
have become so small that the Indians referred to the area as the "land
of little sticks." Then
the forest stops.
Although temperature is believed to play a major role in determin-

ing the poleward boundary of forest vegetation, it is difficult to de-
termine just how this factor operates. We
know that tree line is not
related primarily to the coldness of winter since even in Siberia,
where extremely low temperatures are recorded, heavily forested
regions are found. Nor is the tree growth determined directly by
maximum temperatures in summer, since in
the occurrence of certain
Labrador and in Alaska, far beyond the tree line, temperatures of
32 to 38C have been recorded. High daily maxima are beneficial
since they hasten the melting of snow. In order to survive trees must
have sufficient heat for growth in the interval between the arrival
of effective temperatures in the spring and their disappearance in the
fall. In Alaska, and also on the San Francisco mountains, the conifers
reach but do not pass the July isotherm of 10C (Griggs, 1946). On
the other hand, the position of the tree line in Scandinavia appears
to be chiefly influenced by the occurrence of a season with sufficient
heat for seed germination, especially of the birch.
In some regions the curtailment of tree growth is brought about by
excessive soltfluctionthe heaving of the soil by frost action. Trees
are tilted and their roots broken by severe and in such
solifluction,
situations this indirect action of climate may be more significant
than its direct effect. The occurrence of permafrost (a stratum of
Life Zones 175
permanently frozen soil) prevents the penetration of tree roots, and,

although trees will grow in an unfrozen stratum over permafrost, its
presence usually increases the severity of solifluction. Raup (1951)
regards timber line in Alaska as occurring at thepoint where frost
action prevents the white
spruce from surviving in numbers on the
uplands. Isolated stands of spruce trees are found growing hundreds
of miles north of the where local drain-
recognized tree line in areas
age conditions result in little or no solifluction.
the student of ecology
Fortunately, may observe the profound
effects of the tree line without the necessity of visiting Alaska or the
vicinity of Hudson Bay. In the United States many of the accessible
mountains both in the east and in the west rise to sufficient
heights
to
display altitudinal tree lines. At the upper edges of the coniferous
forest, the spruce, fir, and birch become progressively reduced in size
until they culminate at tree line in a dense scrub
growth or dwarfed
individual specimens. The actual altitude at which tree line occurs
differs
widely according to local circumstances. The harmful effects
of strong wind, including especially excessive evaporation, are be-
lieved to restrict tree line to lower altitudes on exposed mountain
peaks than would be the case if

temperature were acting alone
(Griggs, 1946). Physical influence of ice formations, avalanches,
landslides, and solifluction may exert a further control locally (Fig.
5.17). Thus on different slopes of Mt. Washington, New Hampshire,

described as having "the worst weather in the world," trees give way to
alpine tundra at altitudes ranging from about 1200 m

to 1740 in,
whereas in the Rocky Mountains forests may extend to 3300 m (Fig.
5.18). Tree line occurs at sea level in parts of Belle Isle Strait but
at an altitude of about 300 m
at Bay of Islands, Newfoundland.
Life Zones. The progressive change in temperature and other
climatic factors from the equator toward the poles and from the low-
lands to the mountain peaks controls the distribution of certain major
vegetation types and these in turn are accompanied by

characteristic
sets of subordinate plants and associated animal species. As a result,
we may recognize a series of life zones extending across the land from
the to the polar regions, and, on a much smaller scale, extend-
tropics
slopes from the warm
lowlands to the alpine condi-
ing up mountain
tions of high elevations. Although many irregularities exist and
the distribution of
although the special circumstances influencing
each individual species must be kept in mind, the general sequence
of the latitudinal and altitudinal life zones is clear, and the two series
have recognizable counterparts (Fig. 5.19). The continental life
zones are formed by the roughly latitudinal arrangement of certain of
176 Temperature
FlG. 5.17. Winter view ot tret- line on Mt. Mansfield, Vermont, with spruce trees
in foreground encased in iee. Severe frost action and other adverse climatic
influences reduce the trees to stunted dwarfs.
Photo by D. R. Clarke
FIG. 5.18. General view in midsummer of tree line on Mt. Rainier, Washington,
and of the alpine meadows above it. Tree line here is believed to be relatively
stationary because of the occurrence of both old and young trees among the out-
posts of the forest.
178 Temperature
the principal biotic formations, or bionics, that will be discussed more
fully in Chapter 12.
A synoptic view of the life zones of the world could be obtained

by chartering a plane at the equator and flying toward the north
pole, or toward the south pole. On such a trip, you would observe
that the life zone characteristic of the equatorial region is the tropical
rain forest. In this* zone a wide variety of trees provides continuous
shade and humid conditions for the many species of plants and ani-
mals that live in and under the forest. North of the tropical rain
forest the flying ecologist comes to a life zone in which dry seasons
alternate with rainy seasons. Farther to the north the plane passes
over a belt of varying width of desert or grassland and thence over
deciduous forests of oak, hickory, beech, and maple. The colder
region to the north is heavily forested with spruce and other conifers
that form a circumpolar life zone. At the limit of the spruce forest
the plane crosses the tree line and reaches the tundra where dwarf
birches or willows, scattered grasses or other flowering plants, mosses,
and lichens struggle to keep alive on ground that is frozen for much
of the year. Eventually all land vegetation is lost to view in a land-
scape of universal snow and ice. If the plane trip had been made
from the equator southward, the same general sequence of life /ones
would have been traversed except for the gap caused by the great
expanse of the Antarctic Ocean.
4,000m
3,500
JIMBER LINE
>
Spruce-Fir
3,000
, ....2,500
Submontane shruoV
7,000 L -^
5,500
4,500
UINTA MOUNTAIN S TAVAPUTS PLATEAU
North
FIG. 5.20. Diagrammatic representation of the altitudinal life zones in the Uinta
Basin. (Graham, 1937.)
Without an airplane you may observe the same general series of

lifezones on a much smaller scale by climbing a mountain. If the
mountain is a high one and located in the tropics, as is Mt Popo-
catepetl in Mexico with an altitude of 5448 m, the basal slopes may
Life Zones 179
be covered by rain forest and the summit by snow. As
you climb
upward you emerge from the dense vegetation of the tropical forest
and pass progressively through life zones of savannah, deciduous
forest, and coniferous forest. Higher up you reach a belt of dwarf
fir and alder. Above this a life zone of grasses, lichens, and
pine,
mosses exists, corresponding to the tundra and giving way finally to
TABLE 13
OUTLINE OF PLANT COMMUNITIES i\ THE UINTA BASIN AND THEIR RELATION

TO ALTITUDIXAL LIFE /ONES (Graham, 1937)
Alpine zone
Upper
altitude
Spruce-fir zone
Lodgepole pine zone
Mid
altitude
Aspen zone
Juniper-pinyon zone
Low
altitude
bare rocks, snow, and ice. Altitudinal zonation on the slopes of

mountains in the temperate region is often equally striking but may
not be as extensive. The report by Graham (1937) of the Uinta
Basin of Utah and Colorado contains a detailed study of the life zones
at different altitudes with many excellent photographs of representa-
tive plant communities (Fig. 5.20 and Table 13).
180 Temperature
Although temperature relations are sometimes chiefly responsible
for the demarcation of certain life zones, it is obvious that moisture
extremely influential in many instances, and other ecological

is factors
such as soil condition and length of day play contributory roles. The
positions of life zones cannot be universally correlated with any
one
influence of the environment but are the result of a complex of in-
teracting factors.
Nevertheless, an attempt was made by Merriam in 1894 to set up
zones based exclusively on temperature into which the entire fauna
and flora of North America could be divided. These were called
biothermal zones and, unfortunately, were accepted uncritically by
many There are certain specific objections to the wide
ecologists.
application of Merriam's scheme besides the general warning that the
presence of an organism in a certain locality cannot be accounted for
by saying that it is a "member" of a certain biothermal zone (

Dauben-
mire, 1938). In the place the heat requirement calculations for
first
Merriam's zones were based on 6G

as a threshold temperature for the
germination of wheat, and therefore obviously should not be directly

applied to other species of plants and particularly not to animals.
The assignment of birds and mammals to zones based on environ-
mental heat conditions is especially inappropriate since these warm-
blooded animals carry their own temperature around with them.
Other objections to the use of Merriam's zones appear from a critical
understanding of the varied For example, the
effects of temperature.
number of the degree days required fordevelopment is riot constant
over a wide temperature range. Fifty days at 2C
above the thresh-
old have a very different influence from that of 5 days at 20C above
the threshold, although the "thermal constant" would be the same in
both cases. Furthermore, as explained above, temperatures at seasons
of the year other than those used by Merriam may be critical in con-
trolling the distribution of many species. The fact that distribution

of a species may agree with certain isotherms of the region does not
necessarily prove a direct causal relation. The existence of many
minor species of plants is controlled by the presence of dominant or
accessory species. The geographical range of the dominant species
may in fact be controlled by temperature, or it may be dependent
primarily on the moisture factor, which in turn is influenced by tem-
perature.
Animal populations also may exist in characteristic zones for reasons
that are only indirectly related, or unrelated, to temperature. Horned
larks, for example, are found all the way from Colombia to the Arctic
and occur in very divergent climates and hence in various biothermal
Temperature and Moisture Together 181
zones. The primary relationship is with the vegetation: these larks
are found wherever broad
expanses of shortgrass occur regardless of
thermal conditions. Cone-feeding birds are
similarly found wherever
suitable conifers are growing. The range of this type of bird may
include areas in one lifezone where a northern conifer is growing and
also in another life zone where a southern conifer is growing. It is
obviously absurd to think of the range of this bird as primarily de-
termined by biothermal zonation.
Temperature and Moisture Acting Together

A discussion of temperature as an ecological factor should not be
concluded without calling attention again to the close interrelationship
between temperature and moisture in the terrestrial environment.
Emphasis has been placed on the fact that temperature affects relative
humidity and the rate of evaporation. The converse is equally true
that when evaporation and condensation occur, they tend to modify
the temperature. We
are accordingly not surprised to find that tem-
perature and moisture often interact in such a way as to make it dif-

ficult or
impossible to disentangle the individual effects of these two
factors. Does the salamander go into a sheltered place to keep cool,
to get warm, or to avoid excessive evaporation? Added to the com-
plication is the fact that for many organisms the indirect effects of
these factors may be more significant than the direct effects because
of their control of the vegetation.
We have evidence that the influence of humidity on insects is often
greatly modified by the existing temperature. The dual effect of
these factors on the rate of development of the cotton boll weevil pro-
vides an admirable illustration (Fig. 5.21). This insect pest cannot
develop if the relative humidity is less than 40 per cent or more than
88 per cent, no matter how favorable the temperature may be. On
the other hand, the animal remains dormant regardless of humidity
if the
temperature is lower than 17C or higher than 39C. Within
these ranges the speed of development depends upon the values of
both factors.At a temperature of 28C, for example, the boll weevil
requires 21 days to develop under a relative humidity of 40 per cent,
but it develops in only 11 days if the humidity is between 60 and 65
per cent.
In the foregoing example of the combined influence of temperature
and humidity on rate of development it is of interest to consider what
the limiting factor is.
Picking the point on the diagram representing
a temperature of 24C and a relative humidity of 55 per cent, we see
182 Temperature
that the speed of development will be increased if either the tempera-
ture or the humidity is raised. In this instance more than one limit-
ing influence is
present. In certain situations, two (or more) in-
dependent factors may be unfavorable and both of these must be im-
proved before the organism can continue its

growth or extend its
range. In other situations, the limiting effect of one factor depends

upon the value of another, as in the development of the boll weevil.
The two factors acting together produce the limitation, and may be
referred to as a "limiting combination." The ecologist should always
be on the lookout for possible multiple adverse or modifying in-
fluences in the environment. Limiting combinations are probably in
operation under natural conditions more frequently than is now real-
ized (
Shelf ord, 1951).
-Absolutely
fatal
^Dormancy
or death
-Dormancy
-No
development
-21 days
18 days
16 days
12 days
11 days
Absolutely
-10- fatal
10 20 30 40 50 60 70 80 90 100
Mean relative humidity, %
FIG. 5.21.Generalized scheme indicating the interaction of temperature and
humidity in controlling dormancy and the number of days required for develop-
ment in the cotton boll weevil. (By permission from Animal Chap- Ecology by
man, 1931, McGraw-Hill Book Co.)
An combined action of temperature and moisture

illustration of the
on vegetation is the effect of these factors on the form of growth of
plants and on the life form of the whole plant. The climate exerts a
major control over the type of plant that can exist in each region, and
consequently the life form of the vegetation is to a certain extent an
Temperature and Moisture Together 183
expression or an indicator of climate. This idea has been extensively

developed by the Danish botanist Raunkiaer (1934), whose work
should be consulted for a thorough treatment of the subject. Raun-
kiaer stresses the significance of the adaptations of buds and shoot
tips for withstanding adverse temperatures and surviving drought.
On Raunkiaer delineates different groups of plants. Trees
this basis
and shrubs are placed in one group in which the surviving buds pro-
ject into the air.The perennial grasses are included in another group
in which the buds are situated on or near the soil surface. Plants
with bulblike buds that are protected during adverse seasons by being
buried in the ground are placed in a third group.
Dry cold Wet cold Dry cold Wet cold
Dry hot Wet hot Dry hot Wet hot

Precipitation-evaporation ratio Precipitation-evaporation ratio
VEGETATION SOIL
of temperature and
FIG. 5.22. representation of the interaction
Schematic
moisture effectiveness in controlling the general distribution of vegetational life
forms and soil groups in North America. (Modified from Thornthwaite, 1931.)
Deciduous trees, conifers, shrubs, succulent xerophytes, mosses,

life forms.
grasses, annual plants, and epiphytes represent contrasting
As we have seen the major life zones are distinguished by the life form
of the vegetation. life form characterize the
Plants of distinctive
and these communities are the dominant
major plant communities,
feature of the biotic formations, or biomes, of the world, as described
in Chapter 12.
Thornthwaite ( 1931 ) has worked out indices of the effectiveness of
thermal and moisture elements of climate as related to the range of
in influencing the distribution
temperature and the rate of evaporation
184 Temperature
over the continent of certain vegetational life forms. The scheme
presented in Fig. 5.22 shows in diagrammatic fashion that above a
certain value of thermal efficiency, moisture tends to be more re-
sponsible than temperature for the control of life form. In colder

climates temperature is generally more important. Soil groups are
influenced in a similar way by the climate as well as by the vegeta-
tion. With low precipitation-evaporation ratios and high tempera-
is more In
tures the moisture factor significant than the heat factor.
the far north the temperature is by far the more critical, but in inter-
mediate regions moisture and temperature play more nearly equal
roles in determining soil groups. Here then we have another illustra-
tion of the extremely wide direct and indirect influences of the tem-
perature factor in the ecological adjustments of both plants and ani-

mals.
6
Light
The chief natural sources of light are

sunlight, moonlight, starlight,
and the light from luminescent organisms. In this chapter we shall
deal primarily with visible light, but since the sun's emission also in-
cludes ultraviolet and infrared radiation these components of sunlight
will alsobe considered. Practically all the energy of importance for
organisms under natural conditions is derived directly or indirectly
from the sun. It is true that man has learned to obtain energy from
other sources such as tides, and more recently from nuclear fission,
but man like other organisms depends primarily on the sun for his
main supply of energy.
The radiation from the sun produces a direct heating effect, and
also produces photochemical transformations. After these transfor-
mations have been completed, the energy appears in the form of heat.
Accordingly, all the sun's energy eventually ends up as heat, but cer-
tain portions of the radiation from the sun take part in vital photo-
chemical processes before becoming heat.
Light as an ecological factor is generally highly directional. It
differs in this respect from the temperature factor since heat often
reaches the living organism from many different directions at the
same time. Light is extremely variable. It changes over a tre-
mendous range, often very rapidly. Many organisms can respond to a
value of light that is only one ten-billionth of full sunlight. Although
we often think of temperature as varying widely on the earth's sur-
face, the magnitude of its variation is extremely small compared to
that of light. Light can change from a value near zero to its maxi-
mum within a few tours.
Light is most plants and animals, though some can do
essential for
without For the continued existence of organisms two require-
it.
ments must be met. First, light must not be so strong as to cause

serious harm at any stage in the life history. We shall see that a con-
185
186 Light
siderable variation exists in the upper limits of tolerance to the light
factor. Second, for those animals and plants that require light it must
be sufficient in intensity and in duration. The intensity of the light
must be above the threshold for the organism concerned, and the total
amount of light received during the period when it is needed must
be adequate.
DISTRIBUTION OF LIGHT
In discussing the distribution of light it is convenient to start with
sunlight as the chief source and trace the radiation as it passes through
the air and water environments. The magnitude of the solar radia-
tion as reaches the outer atmosphere referred to as the solar con-
it
stant has the value of about 1.9 g-cal/cm 2 /min. At sea level the in-
2
tensity of solar radiation averages about 1.5 g-cal/cm /inin, If the
radiation received from the sun were evenly distributed over the sur-
face of the globe, it would be sufficient to melt a layer of ice 35 m
thick during the course of a year. At a latitude of 44 N the energy
received on the earth's surface from the sun is
equivalent to the light
that would be produced by hanging a 250-watt lamp over each square
meter of the ground.
We know very well, however, that the radiation from the sun is not
evenly distributed cither in time or in space. We
wish to inquire
what its variations both in quality and in quantity are. Light changes
in spectral distribution and in angular distribution. The light factor
also varies in intensity and in duration, resulting in differences in the
total amount of light falling on each unit of surface for each unit of
time, such as a day or a month. The changes in the light factor in

these respects will be examined as they affect the terrestrial and the
aquatic environments.
Light on Land
Spectral Composition. The spectral distribution of light as it
reaches the earth's surface isshown in Fig. 5,3. Authorities differ as

to the exactwavelength limits to be assigned to the different portions
of the spectrum. Roughly speaking, radiation of wavelength longer
than 7600 Angstrom units is considered to be infrared, and that of
wavelength shorter than about 3600 A is designated as pltra violet.
Almost one-half of the total emission of the sun is infrared radiation,
and almost one-half is visible light. These proportions remain ap-
proximately the same at the earth's surface, regardless of the total
intensity of sunlight. The ultraviolet component, however, is

always
Intensity of Light 187
only a small percentage of the total radiation, and it may be reduced

to immeasurably small
quantities under certain circumstances.
Intensity of Light. The intensity of light reaching the earth's
surface varies with the of incidence and with the amount of ab-
angle
sorption by the atmosphere and by obscuring features. The lower the
altitude of the sun, the smaller the angle of incidence and the
is
longer
is the of the
path light through the atmosphere with corresponding
reduction in intensity. Changes in the sun's altitude result from dif-
ferences in latitude as well as from
changes in the season and in the
time of day. The greatest intensity of sunlight occurs at
positions
on the earth's surface and at times at which the sun is most nearly
overhead. At higher latitudes, the intensity of light is correspond-
ingly reduced. At 50 N latitude, for example, during the period of
the equinox in March and in September when the day is
everywhere
12 hours long, the intensity of sunlight is
only about one-half of what
at the equator ( Fig. 6.1 ).
it is
2
FIG. 6,1. Total solar radiation g-eal/day/cm of horizontal surface) on March
(
21 with average cloudiness. (Modified from Kiiuball, 1928.)
Latitude thus has a definite effect; but other factors may have much
greater influence upon the light factor. Moisture, clouds, and dust
in the atmosphere have a profound and irregular effect in reducing
illumination. Living organisms may also act to diminish the in-
forest vegetation, and
tensity of daylight, as is clearly shown by the
this represents a reciprocal action in which the inhabitants modify
the light factor in their own environment.
Different forest communities vary widely in the degree to which
they diminish the sun's radiation. Cottonwood (poplar) trees tend

188 Light
to grow rather widely spaced, and the relatively sparse foliage
allows
many patches of sunlight to reach the ground. The canopies of pines

and oaks usually have fewer gaps. Measurements made in Illinois
revealed that the portion of the forest floor exposed to direct sunlight
was 84 per cent for poplar, 77 per cent for pine, and 35 per cent for
oak. In elm-maple forests and in tropical rain forests the canopy
typicallyis
complete, with the result that none of the direct sunlight
reaches the forest floor. Other measurements showed that the light
on the which had passed through the leaves of the canopy
forest floor
was reduced its value above the tree
in intensity to 3.5 per cent of
tops in the oak forest; corresponding figures for elm-maple and rain
forests were 0.4 per cent and 0.2 per cent, respectively. Thus less
than 1 per cent of the light outside the forest reaches the ground if
the canopy is complete.

Different types of forests exhibit different seasonal influences on the
light factor. Underneath a stand of pine trees the light is reduced

by about the same amount throughout the year since the trees are
evergreen. The illumination on the floor of a pine forest reaches a
maximum in the early summer and a minimum in winter correspond-
ing to the seasonal variation in the intensity of the incident sunlight.
In a maple forest, however, a very different seasonal picture is pre-
sented. During January, February, and March the plants and ani-
mals living on the forestfloor receive
increasing amounts of light, but
in April the leaves begin to appear and the intensity of light drops
rapidly reaching a minimum in the middle of the summer. Then, as

the leaves of the maple trees begin to curl and drop off a larger
amount of light is allowed to filter
through the forest canopy. The
striking seasonal variation in the amount of ultraviolet radiation
pene-
trating different forest types is shown in Fig. 6.2. The changes in the
total visible illumination arc of a

corresponding nature (Park, 1931).
Accordingly, if
you were a beetle, or, if yon prefer, a lily growing in
a maple forest, the brightest month of the year for you would be April
and the darkest would be July. In this situation living organisms have
modified the environment in respect to the light factor so that the
seasons have been practically reversed.
Duration and Amount of Light. The total amount of light received

by an organism is determined both by the intensity of the light and
the duration of the period of irradiation. The situation is similar to
the exposure of a photographic plate. The amount of blackening of
the plate is determined by the intensity of the light times the period
of exposure. In natural habitats, the variation in the length of day
Duration and Amount of Light 189
.080
.070
'"Jan. June Dec,
FIG. 6.2. Mean monthly maximum intensities of solar ultraviolet radiation
(2900-4000A) in forest communities of northern Indiana. ( Stroliecker, 1938.)
often has a greater effect on the total amount of light received than do
differences in the intensity of the sun at noon.
On the equator the day is always 12 hours long, but in the temper-
ate regions the day grows longer as spring progresses. This effect is
accelerated at higher latitudes, and the day becomes 24 hours long
during the summer in the polar regions.

The day becomes cor-
the summer solstice (Fig. 6.3). Up to
respondingly shorter after
latitudes the increase in length of day during summer
moderately high
has more effect on the total amount of light received per day than the
190 Light
i
.
n .m .iv. v 4
vr vii viii ix. x xi xii i n m, iv v vi
60 N
40*
60 S
80* S
FIG. 6.3. Variation in length of day at the indicated latitudes during the indicated
months. Arrowheads indicate periods of decreasing daylength. The heavy line
shows the course of zenith sun ( Baker, 1938. )
7
FIG. 6.4. Total solar radiation g-cal/day/cm of horizontal surface) on June 21
( "
with cloudless sky. ( Modified from Kimball, 1928. )

reductions in solar
intensity due to the greater angle of incidence. The
greatest northward development of this influence is illustrated in Fig.
6.4 in which the total solar radiation, determined
by the combined ac-
tion of intensity and duration, is
plotted for June 21 under conditions
of cloudless sky. At that period the amount of solar radiation received
each day at 50 N latitude is than that received
considerably greater
at the equator, and the daily amount of light received in Scandinavia
is
roughly the same as that falling on tropical Africa. If
you are
amazed to find potatoes
growing in
Norway, or wheat being harvested
in Alaska, don't
forget that once the plants have begun to develop in
the late spring, sufficient light exists in these latitudes for the
plants
to continue all
growing and a
day of the
good part "night."
The foregoing review of the circumstances of light on the earth's
surface has shown that light is everywhere sufficient for life of some
type at least for part of the year. Plants and animals exist from pole
to pole. Mosses and lichens grow in abundance right up to the edge
of the ice in the polar regions. On the other hand, light is nowhere
too strong for animals and plants of some sort. Thus the whole range
of light on the earth's surface generally compatible with life. Of
is
course, light may sometimes be too strong or too weak for individual
species, and in these instances it controls growth and distribution.
Light in Water
The sunlight available for plants and animals in the aquatic en-
vironment has entered the water from the air and hence has first been
subjected to all the changes imposed upon it by the conditions above
the surface. Ten per cent or more of the light is lost by reflection at
the surface or in the special conditions just beneath the surface
(Clarke, 1939). In addition, in passing downward, the light is fur-

ther modified by the water medium in respect to intensity, spectral
composition, angular distribution, and time distribution.
Extinction and Modification of Light. The light factor in the water
environment subject to a number of variable influences that modify
is
it
profoundly. In order to consider the situation first in its simplest
terms, imagine a lake from which we have pumped out all the natural
water and refilled with distilled water. Our lake then contains water
of uniform and maximum transparency from top to bottom. L Light is
reduced in intensity both by absorption and by scattering, and the
rate of reduction is measured as the extinction rate, although the term
absorption rate is sometimes

used loosely for the combined effect.
Pure water causes the extinction of light at different rates in different
192 Light
parts of the spectrum. Measurements of illumination made with a
photometer placed watertight case and lowered into our imag-

in a
inary lake would reveal the changes presented in Fig. 6.5 At a depth
of 70 m the blue component of sunlight has been reduced to about
70 per cent of its
intensity at the surface. At the same depth the
yellow component of sunlight has been reduced to 6 per cent of its
incident value. The orange and red components have been ex-
tinguished very much more rapidly. At a depth of 4 meters, red
light has already been diminished to about 1 per cent of its surface
intensity.
Percentage of incident light
5 10 50
TTT
FIG. 6,5. Reduction in intensity (logarithmic scale) of the color components of

sunlight (indicated by initial letters) at increasing depths (linear scale) in a lake
of optically pure water. (Clarke, 1939, AAAS Pub. No. 10.)
Since we know that the daylight incident upon the surface of our
lake differs markedly in intensity in different parts of the spectrum,
we are dealing with the unequal extinction of an unequal spectrum.
As shown in Fig. 6.6, the extinction by pure water of light at the two
ends of the spectrum is much more rapid than in the middle of the
spectrum. vThe blue component of light is also the most penetrating
in the clearest ocean and lake waters. v At depths of 100 m or more
the blue light becomes

completely predominating;) this was observed
directly by Beebe in his
bathysphere dive off Bermuda. An observer
looking straight down into the water from the deck of a ship in the
resulting from this
clear tropical parts of the ocean sees the blue color
selective absorption. Since sunlight has been shorn progressively of
its
longer and shorter wavelengths, the only component remaining to
be scattered upward again to the eye is the blue. Anyone who has
seen the intense indigo of the tropical oceans will never forget it.
- uv-
3000 A 4000 5000 6000 7000 8000
Wavelength
The of solar energy at the earth's surface and
FIG. 6.6. spectral distribution
of pure water. Similar
after modification by passage through the indicated meters
light conditions would be found in the clearest ocean and lake waters. (Clarke
1939, AAAS Publ No. 70.)
The foregoing has shown that even pure water absorbs light at a
to air and causes a profound change in
very rapid rate compared
In the clearest parts of the ocean and in ex-
spectral distribution.
clear lakes the optical properties of the water are closely
ceptionally
similar to those of pure water, Other natural waters contain sus-
and dissolved material in sufficient quantities to
pended particles
cause a further reduction in and a further alteration in
transparency
spectral composition.
includes living organisms that in-
Suspended material in the water
crease the extinction of the light and thus modify their own environ-
ment in this respect. Illumination is reduced by beds of kelp along
194 Light
the seacoast and by submerged or floating vascular plants along the
shores of inland waters in much the same way that light is cut down
by the larger vegetation on land. In the free water of ponds, lakes,

and the oceans the phytoplankton is sometimes sufficiently abundant
to produce a noticeable reduction of light. Plankton populations may
cause an additional extinction of light indirectly by adding detritus or
stains to the water after the organisms have died and disintegrated.
A thick "bloom" of algae in a pond may thus reduce the light supply
to such an extent as to curtail its own growth and that of other plants
in the water layers beneath.
B G Y
Wavelength
FIG. 6.7. Reduction in intensity and shift in spectral composition of light in
heavily stained Rudolph Lake, Wisconsin. (Clarke, 1939, AAAS Publ. No. 10.)
In temperate and coastal seas and in the majority of clear inland

lakes fine particles or stains are present that tend to absorb or scatter
the blue component of light more strongly than occurs in pure water.
As a result the green component of sunlight is usually the most pene-
trating in these situations and gives the

water its characteristic emerald
color. The organic stains occurring in some ponds and rivers absorb
the shorter wavelengths so strongly that the red or orange components
of sunlight become the most penetrating. In Rudolph Lake, Wiscon-
sin, for example, the combined absorption of the water and of stains
in the water causes not only a very rapid reduction in the light with
Extinction and Modification of 195
Light
depth but also a shift in the
position of the maximum light to the red
region of the spectrum (Fig. 6.7).
Since natural waters differ
very greatly in transparency and in se-
lective a of the
absorption, study penetration of each part of the spec-
trum in each body of water would have to be made for a
complete
description of the light conditions. An approximate comparison of
the illumination in various natural waters can be made on the basis
of the relative
transparency to the central part of the spectrum.
Sample values are given in Table 14 for representative bodies of
TABLE 14
TRANSPARENCY OF WATER TO CENTRAL PART OF VISIBLE SPECTRUM

(5000-0000 A)
Extinction Depth for \% of Secchi Disc

per Meter (<*;.) A:* Surface Light Depth f
Distilled water 3.8 . 039 118 m 44 m
where /o = initial intensity.

7 = intensity at depth, L, in meters.
f
c/
= 27
/c. i .
t Depth at which a 20-cm white disc disappears when lowered from the
surface.
water. Since the great differences in the extinction of light per meter
have a cumulative effect, an even greater contrast is presented in the
depths at which a given fraction of the light is found. The depth
at
which the light intensity is reduced to 1 per cent of its surface value
is of particular importance because it represents the approximate
lower limit for plants, as will be discussed later. In general the table
and the representative curves in Fig. 6.8 show the profound variation
in the intensity of light available for organisms at increasing depths
in different natural waters.
196 Light
Another way in which light is
changed by water is in its angular
distribution. The direct beam
of light from the sun is bent by re-
fraction toward the perpendicular at the water surface. Since waves
or ripples almost always exist in natural situations, the light from the
sun tends to be broken up to a considerable extent in passing through
the surface. Within the water itself further diffusion is produced by
scattering. When
larger amounts of suspended materials are present,
scattering proceeds at a very rapid rate, with the result that measur-
Percentage of surface light

0.1 0.5 1.0 5 10
INN I
FIG. 6.8. Comparison of the transparencies of representative natural waters.

Curves based on the average extinction rate for the yellow-green (5000-6000A)
component of daylight. (1) Midge Lake, Wis., (2) Trout Lake, Wis., (3) Gun-
flint Lake, Minn., (4) Woods Hole Harbor, (5) Thatcher Pass, San Juan Islands,
and Buzzards Bay, Mass., (6) Vineyard Sound, (7) Baltic Sea, (8) Crystal Lake,
Wis., (9) English Channel, (10) Gulf of Maine, (11) off Vancouver Island, and
off Nantucket Shoals, (12) Crater Lake, Ore., (13) Gulf Stream, (14) Caribbean
Sea, ( 15) distilled water. (Clarke, 1939, AAAS Publ No. 10.)
Changes in
Transparency 197
able light is
passing in every direction. In the upper strata the light
in the direction of the retracted
passing rays from the sun is the
strongest. With increasing depth the direction of strongest light in-
tends to move toward the vertical, and a
tensity limiting pattern of
diffusion is established Thus, although a variable de-
(Jerlov, 1951).
gree of diffusion occurs, the directional character of the light in the
water is never entirely lost.
Changes in
Transparency. Parts of the ocean and of lakes that
are well stirred exhibit a uniform
transparency from the surface down-
war6\ but, when strong density gradients exist, considerable changes
in transparency may occur with Furthermore, the transpar-
depth.
ency of coastal marine areas and of inland water bodies often changes
profoundly from season to season, owing to differences in amount of
stirring, in the discharge of muddy rivers, and in the growth of plank-
ton. Such variationsin transparency are added to the seasonal
changes the
in intensity of the light received at the surface and in the
length of the day. The* combined effect of these influences results
in surprisingly great fluctuations in the light factor at subsurface
levels.
Observations made off the coast of Massachusetts illustrate the mag-

nitude of the foregoing seasonal influences (Clarke, 1938). In this
region the maximum hourly intensity of light received at the surface

during the summer averaged about twice that received during the
winter period. The average total solar radiation received per day
in summer was about four times as great as that received during the
winter. Significant departures from the average for periods up to
several weeks occurred, and such variations are undoubtedly im-
portant in causing changes in the response to the light factor from
year to year. Values for individual days may also depart markedly
from the average. During the investigation referred to above the
maximum light received on the brightest day of summer was forty
times greater than that received during the dullest day in winter.
In the course of the same year the extinction coefficient of the water
varied fourfold. Since the effect of extinction is cumulative, the il-
lumination in the subsurface layers varies more because of changes
in transparency than because of seasonal changes in radiation reach-
ing the surface. By combining the two effects it was found that for
an organism living at a depth of only 30 in the minimum hourly
light intensitywas 7000 times greater and total daily radiation was
10,000 times greater in May, when high incident light was combined
with high transparency, than in December, when both incident il-
lumination and transparency were low. The magnitude of seasonal
198 Light
changes in illumination in the aquatic environment are therefore fre-
quently very much greater than in the terrestrial environment. Any
water body that displays a much higher transparency in winter than
in summer have less light available during the summer in its
will
deeper layers and thus suffer a reversal of the usual seasonal change
in the
light factor. The foregoing instances are sufficient to show
that in the aquatic environment light becomes profoundly altered
quantitatively and qualitatively and that its changes may go far be-
yond those experienced by plants and animals on land.
BIOLOGICAL EFFECTS OF LIGHT

Before concerning ourselves with the influence of the light factor
on growth, reproduction, locomotion, and other activities of the or-
ganism as a whole, we shall consider certain general effects of radia-
tion as strikes the surface of the plant or animal.
it In addition to
heating the tissues, the absorbed radiation affects biological processes

in the
exposed tissues, including particularly their pigmentation.
General Effects
Among green plants light is required for the production of chloro-

phyll in the chloroplasts. Plants germinated under insufficient il-
lumination will not develop their normal green color. Normal plants
become etiolated in the absence of light; that is, they lose their pig-
ment and develop abnormal form. On the other hand, excessive il-
lumination causes the destruction of chlorophyll. In some plants ex-
cessive absorption of light by the deeper tissues is prevented by the
screening action of thickened chloroplasts or of increased numbers of

chloroplasts near the surface. In some species when the light be-
comes too bright, the chloroplasts line up one behind another so that
a larger proportion of the radiation passes through the leaf between
the chloroplasts. When the light becomes weak, the chloroplasts
spread out and absorb a maximum percentage of the incident illu-

mination.
Light may influence th(^ pigmentation of animals in several ways.

Skin color may be indirectly affected by light through the mediation
of the eyes or other receptors. In other instances differences in color
may have arisen as a result of selective survival. The abundance of

pigment exposed in the chromatophores is sometimes directly con-
trolledby the intensity of the light received. The characteristic lack
of pigment in cave animals is associated with darkness, and certain
aquatic forms have been shown to lose their color when removed from
light. Blind cave amphibians and fishes with little or no color have
been found to develop abundant pigment in the skin after exposure
to normal daylight
(Rasquin, 1947). Intense radiation may be harm-
ful because of undue
heating or evaporation, because of the lethal
action of the ultraviolet component, or in other
ways.
The excessive absorption of light by animal tissues must also be
avoided. Most animals simply move into the shade, burrow into
the ground, or descend to deeper levels in the water. The
develop-
ment of a relatively transparent body would appear also to help deal
with this Since only the radiation that is actually absorbed
problem.
can be effective, the highly transparent tissues of many types of plank-
ton can retain but little light energy. This fact may enable some
planktonic animals to endure higher light intensities in the surface

waters than would otherwise be possible. As we shall see, many
species of plankton are extremely sensitive to light. In other kinds

of animals evolutionary development has gone in the opposite direc-
tion with the production in the skin of abundant pigment which pro-
tects the deeper tissues. Anyone who has lain too long on the beach
in June has been painfully aware of the harmful effect of excessive
radiation on animal tissue and is familiar with the deposition of pig-
ment in the skin of man to produce tan. In addition, transparency or
pigment patterns also serve to render animals less conspicuous.

Protective Coloration. The pigmentation of a great many animals
forms a coloration that appears to afford protection from enemies.
One very common
type of protective coloration is a simple matching
of the background in respect to color and pattern. A quail squatting
in the grass or a bittern standing motionless among the swamp reeds
is
exceedingly difficult to distinguish from its surroundings, as is a
moth on the bark of a tree or a katydid among green leaves. Many
other instances of remarkably close resemblance of birds, insects, and
other animals to their background will occur to the reader. A second
common type of protection, often combined with the first, is oblitera-
tiveshading which the
in or fish displays darker pig-
bird, mammal,
mentation on its back and lighter color underneath. This difference
tends to counteract the stronger illumination received from above
with the result that the animal blends with its
background.
Theprotective aspect of other color patterns may result chiefly from
the confusion of the enemy by the disruption of the animal's usual
outline. Birds with necks or heads of contrasting colors or tropical
fishwith strong transverse stripes are not easily recognized by man
under certain conditions of illumination because of the unexpected
200 Light
pattern and presumably are not easily recognized by their natural

enemies. Any such visual deception will, of course, not affect preda-
tors that locate their
prey exclusively by smell, sound, or other meth-
ods. Some species when disturbed produce a flash of some brightly
colored or contrasting part that is concealed when the animals are
quiet, and the startling effect may serve to distract or to frighten off
pursuers.
Another type of protective resemblance is
mimicry a phenomenon
that particularly well developed among the insects.
is Here one
species closely resembles in color and form another totally unrelated
species sometimes in an entirely different order. The mimic is be-
lieved to derive protection from the fact that it is mistaken by preda-
tors for the species it resembles. If the model
species is distasteful or
harmful as a food organism and hence is avoided by the predators,
the mimic may also escape unmolested. Extraordinarily close
exists between insect species, but the the
mimicry certainly reality of
benefit of the mimicry, its mode of operation, and its evolutionary
origin are controversial subjects which still await conclusive investi-

gation. For a further discussion of the far-reaching problems of pro-
tective coloration the reader may consult more extensive treatments
of the subject such as that by Cott (1940).
Some animals
are able to change their color or pattern sometimes
within a matter of minutes or even seconds, Such changes in ap-
pearance occur characteristically as adaptations to the background,

and are found among reptiles, amphibians, fishes, crustaceans, insects,
cephalopods, and other invertebrates. The mechanisms by which
color changes are brought about have been summarized by Prosser
(1950, Ch. 21). In many instances they involve the nature of the
light received from the background through the eyes, but in other in-
stances, they are activated by direct radiation. A flounder changes
its
general color tone and also the pattern of the black and white
its skin as it moves from one
patches of type of bottom to another.
Such changes in coloration tending to match the background furnish
an obvious advantage in concealment (Summer, 1935). In other in-
stances color changes serve as protection from high illumination, take
part in thermoregulation, or are associated with breeding, as in cer-

tain lizards, fishes, and squids. The seasonal color changes of the
varying hare, weasel, and ptarmigan from brown in summer to white
in winter are obviously related to the conspicuousness of such animals
against bare ground or snow-covered landscape.

The different, and usually more brilliant, coloration of the males
of many birds and of some other animals is familiar to everyone. In
attempting to explain the evolutionary origin of this sex difference,

Darwin drew attention to the antics of peacocks, pheasants,
courtship
and other birds, in which the males
appear to vie with one another
in showing off
plumage. Darwin proposed that the special
their
plumage male
of the evolved after many generations in which the
females selected and mated with the "most beautiful" birds. Since
we have no reason to suppose that birds judge the attractiveness of
male suitors on the same basis as we would, this explanation seems
unsatisfactory. As yet no convincing demonstration has been made
of any basis for selection or of
any other method by which the elab-
orate decoration of the male
may have arisen.
The duller coloration characteristic of the female is
undoubtedly
related to her greater need for concealment while brooding the eggs.
The striking coloration of the males of many species can rarely have
any protective value for the male himself, although his conspicuous-
ness might draw attention away from the female on the nest. In some
birds, such as Wilson's phalarope, the tables are turned, for the fe-
males are brightly colored and the drab males do the housework of
incubating the eggs. Difference in appearance may play a useful
part in aiding sex recognition. The
breeding plumage of
brilliant
the male is often replaced by a duller garb during the winter season.
The length of day has been shown to influence breeding, migration,
and color change in many birds and mammals. The light factor may
thus be involved in coloration through its effect on reproductive ac-
tivities as well as through its role in protective resemblance, and the
two may be interrelated.

Mention was made in earlier chapters that desert animals charac-
teristically display
a pale coloration in contrast to the darker hue of
the inhabitants of humid regions. Although temperature, moisture,
and light may directly affect the general color of terrestrial animals
under some circumstances, these factors often act indirectly through
their influenceon the color of the ground in relation to the conceal-
ment of the animals. Evidence indicates that pale or dark coloration
has evolved in many species and races as a result of selective survival
as influenced their conspicuousness against the background. Se-
by
lection would thus account for the occurrence of a white mouse
(Perognathus apache) inhabiting an area of white gypsum sands in

New Mexico and of a black mouse (P. intermedius) living on an ad-
lava (Benson, 1933). The possibility that cli-
joining area of black
mate sometimes exerts a direct effect, however, is suggested by the
fact that nocturnal animals in the desert have the same pale coloration
202 Light
as those which are abroad during the daylight hours when light would
appear be much more important.
to
It is a mistake to assume that the
pigment developed by an organism
'
Necessarily plays a critical role in its

present relationships. In some
instances the pigment; may be primarily a by-product of metabolism
without any ecological significance. The red color of the deep-sea
shrimp is a case in point. Since there is no red light deep

in the
water, the shrimp must appear as black as the deep-sea fish which
share the same habitat. Similarly, it is unnecessary to assume that
the coloration of the scarlet tanager, the oriole, and other birds with
brilliant plumage provides any protective resemblance. Certain
writers have practically turned themselves inside out in attempts to
find a protective function for all bright colors. No one in his right
mind would try to claim that Chromodoris, a bright blue nudibranch
with orange and gold spots, was protectively colored as it creeps over
the gray rocks and brown seaweeds of the tidal zone of southern
California. In such instances either the bright colors do not attract
enemies, or the organism survives in spite of being conspicuous be-
cause of the possession of other, sufficiently advantageous attributes.
Activity and Vision
Photokinesis. Light controls the locomotory activity of many of the

lower organisms by a direct action upon their speed of locomotion
a phenomenon known as photokinesis. The magnitude of this reac-
tion among animals without eyes was well illustrated by a laboratory
test on the larvae of the mussel crab, Pinnotheres maculatus (Welsh,
1932). These animals swam the length of a 29-cm trough toward a
light of
0.5 meter-candle in 34 sec. As the light intensity was in-
creased, their swimming rate was accelerated regularly until at 46
meter-candles they made the trip in 17 sec. Many other animals
show a similar increased activity under increased illumination whether
or not they are oriented to the light. This simple direct relationship
is of
profound importance in the lives of many of the aquatic inverte-
brates and of the smaller terrestrial forms including insects.
Vision. Whenactivity is controlled by light among higher forms
it
usually through vision.
is We
should inquire as to the circum-
stances of illumination under which vision is possible. On
land day-
light is everywhere sufficiently strong at some period for the vision of
those animals that possess eyes. In the water because of the rapid
which light is reduced in intensity we may suspect that depths
rate at
would soon be reached at which vision is no longer possible. Let us
Vision 203
inquire what is the minimum illumination for aquatic animals to see

sufficiently to feed, to find their mqtes, or to avoid dangers. Interest
also centers on the maximum
depths in the aquatic environment at
which responses to day and night still exist.
A laboratory test can be set up in which the intensity of light is
varied over a considerable range with relative ease. But it is not so

easy to ask a fish whether or not he can see small objects at least
it isnot so easy to get him to
reply. An answer was obtained, how-
ever, from the fresh-water sunfish, Lepomis, by using the response of
the fish to background motion. If this type of fish is placed in a
glass cylinder with a surrounding screen made of bars and spaces, the
fish
responds by a turning motion if it sees the screen rotate. It is
through this kind of reaction that many fish maintain their position
in a stream. Reactions of this sort are involved in rheotaxis, dis-
cussed in an earlier chapter. By reducing the light intensity in an
experiment of this type until the sunfish no longer responded, the

minimum illumination under which the fish could see small objects
similar in size to the bars of the screen, was found to be one ten-
ln
billionth (or 10 )
of the value of full sunlight (Grundfest, 1932).
The threshold sensitivity of the human eye is similar, and the value
for other vertebrates is
probably generally of the same order of
magnitude.
Assumingthat the threshold illumination for other species of fish
issimilar to Lepomis, determinations were made from transparency
measurements of the approximate depths at which this minimum
intensity of would occur under different circumstances, and
light
hence the maximum depths at which vision would probably be pos-
sible, although other optical conditions such as color and contrast with
background would have to be taken into account (Clarke, 1936).
In most lakes except the most turbid and in typical coastal areas
vision would appear to be possible for fish similar to Lepomis at all
levels right down to the bottom. In the open ocean beyond the mar-
gins of the continental shelf the water is far deeper than this maxi-
mum depth for vision. If deep-sea fishes can see as well in blue
light as the sunfish tested could see in green light, then vision would
be possible for them at depths of more than 700 m in the clear
tropical ocean.
Manyaquatic animals may show some activity response to the
increase or decrease of light at still lower intensities. Studies of the
diurnal vertical migration of zooplankton indicate that certain species
may react to light from the surface at 800 m

and possibly at 1000 m,
or more than half a mile down. This depth probably represents the
204 Light
biological zero for the response to daylight penetrating from the sur-
face. Below this level day and night no longer exist, and no seasonal
change in illumination can be detected.
Since the average depth
of the ocean about 4000 m, it is clear that more than three-fourths
is
of the volume of this environment is devoid of any influence of day-
light.
The whole of the deep sea is
by no means completely dark,
however. Light of biological origin occurs irregularly through the
marine environment, and bioluminescent organisms are sometimes
very abundant in the sea.
Bioluminescence. Our discussion up to this point has dealt pri-
marily with light emanating directly or indirectly from the sun, but
light of biological origin, known as bioluminescence or popularly as
y
"phosphorescence," has ecological significance under certain circum-

stances. As mentioned above, bioluminescence is the only source
of light in the deep sea. Near the surface of the ocean and on land
it is
frequently prominent during the night, but it
rarely occurs in
fresh water. Luminescence produced by members of various taxo-
is
nomic groups scattered through the animal kingdom including in the

sea certain fishes, crustaceans, coelcnterates, and many other in-
vertebrates, and on land particularly the insects. Fungi and such
microorganisms as the dinoflagellates and many groups of bacteria

also are luminescent. For a complete summary of the occurrence
and physiology of bioluminescence the reader should refer to the
book on the subject by Harvey (1952).
The very considerable amount of illumination that can be provided
by luminescent organisms may be appreciated by anyone who catches
some fireflies and brings them into a darkened room. A few fireflies
in a bottle will provide sufficient light for reading newsprint. The
"phosphorescence" of the sea sometimes is almost dazzingly brilliant
on a dark night, at which time waves and the wake of a boat appear
like"burning water." The bodies of fish and other organisms swim-
ming through the water are outlined by millions of tiny lights. This
illumination is caused by the luminescent discharge of the plankton
organisms. Measurements have shown that the intensity of light

emitted by the surface waters of the sea may be one thousand times
greater than the threshold intensity for the vision of man.
Bioluminescence is thus sufficiently strong to evoke reactions under
a variety of circumstances. Although this living light may serve no
useful purpose in some situations, in other instances it probably
one or more of the following functions: (1) illumination, (2)
fulfills
recognition, (3) lure, and (4) warning.

The luminescence of deep sea animals undoubtedly provides a use-
Biolwn inescence 205
ful amount of illumination for the individuals it as well
producing
as for other inhabitants in the immediate neighborhood. In shallow
water and on land
during the night animals may similarly use their
photophores as lanterns. The employment of luminescence in recog-
nition of one sex another
by is
clearly exemplified by the firefly.
the American
Among Lampyridac each species has a characteristic
code of flashing by which the females can
recognize males of the
same species and distinguish them from males of other species. After
dark the female climbs up on a blade of and when a flying male
grass
signals in her vicinity she attracts him to herself by recognizable
flashes in
response (Harvey, 1952, Ch. 13).
In the abyssal
depths of the sea, fish can perhaps locate each other
by recognizing the pattern of lights presented by the luminescent
It has also been
organs. suggested that squid are able to keep to-
gether in a school during the dark hours of the night by means of
their characteristic
flashing.
Animals may lure prey by means of their luminescent organs.
Since many small fish and planktonic invertebrates are attracted to
light, these animals would be expected to move toward the lumi-
nescent organs of predatory fish, many of which are located near the
jaws or even on filaments dangled in front of the mouth. On the

other hand, the sudden flash of luminescent organs may act as a
warning to scare off predators. Certain deep-sea shrimps produce

a luminescent secretion that may be discharged into the water. This
is an interesting counterpart to the sepia produced by squid and
cuttlefish in shallow water. When the latter are attacked, they can
discharge the black secretion into the water and escape from their
enemies in the "smoke screen'* thus produced. In the inky blackness
of the deep sea the shrimps that produce a luminous discharge when
attacked may be
able to escape in a "cloud of light" (Fig. 6.9)-~or
these sudden emissions may act simply by distracting or frightening
the enemy.
The possession of luminescence may be a definite disadvantage for
some organisms if it
gives away For other
their presence to enemies.
species the emission of light may be an accidental by-product of

metabolism and hold no ecological significance whatsoever. can- We
not imagine, for example, any possible benefit for bacteria that could
be derived from their production of luminescence.
A correlation has been thought to exist in the marine environment
between the occurrence of eyes, the type of coloration, and the pres-
ence of daylight or luminescence. Eyes are well developed among
animals inhabiting the upper layer of the ocean, and luminescent
206 Light
organs occur in widely different groups. At somewhat deeper levels
"telescopic" and in some species
the eyes of fish tend to be enlarged or
directed upwards. At greater depths
eyes are often degenerate or
entirely absent. Eyes would perhaps be of little use here since
both daylight and animal light produce very meager illumination.
N.C.S. Copyright, 1934

FIG. 6.9, Drawing to illustrate the
appearance of the photophores of the deep
sea fish Photostomias guernei and of the luminous discharge of the
shrimp
Acanthephyra purpurca in the inky blackness of the ocean abyss.
Orientation 207
In the ocean abyss by contrast,
many fish possess well-developed
eyes and the bottom-living organisms produce abundant luminescence
(Sverdrup et al., 1942).
In caves no luminescence is
produced by the aquatic inhabitants
and only one form-the New Zealand glowworm (Arach-
terrestrial
nocampa lwninosa)-is known to be luminescent. The fauna of caves

that are completely cut off from
daylight consists typically of species
that are blind or have degenerate Cave animals usually have
eyes.
little
pigment and contrast strongly with the jet black or dark red
deep-sea animals. The heavy pigmentation of the lattermay perhaps
be explained as a protective
adaptation serving to reduce the reflec-
tion of luminescent light from their surfaces and hence the chance
of detection. Since cave waters are completely dark, the whitish
coloration of the inhabitants would be of no
disadvantage on this
score.
Orientation
Light often plays a significant role in orienting the growth or

locomotion of plants and animals. Since orientation to light is often
associated with reactions to other factors in the environment, such as
gravity, we shall first consider the subject in general terms. Orienta-

tion is brought about either by the differential growth or movement
of parts of the organism or by a change in the direction of locomotion
of the whole organism. The question of why plants and animals grow
or move in the directions they do has occupied the attention of in-
vestigators for a long time. Many of the theories and terms used have
been in conflict, and for a more extended discussion reference should
be made to Fraenkel and Gunn (1940) and to Griffin (1953).
The term tropism is best used for orientation by growth or turgor
movements as exhibited by sessile forms. These forms are usually
plants, but essentially
the same type of orientation is exhibited by
many animals such as the hydroids. If the orientation is to

sessile
gravity, the term geotropism is used. If it is to light, the growth

movement is referred to as phototropism, and other prefixes are used
for other orienting factors. OfTthe other hand, the orientation of
the locomotion of motile organisms is best referred to as a taxis,
although it is sometimes also

called a tropism. Here the forms in-
volved are usually animals, although motile plants such as the green
or zoospores are included. Ap-
flagellates and
motile plant gametes
plying suitable prefixes,

we obtain the terms geotaxis, phototaxis, etc.,
for orientation by the corresponding factors. Orientation in the direc-
tion of an orienting force is referred to as a positive tropism or
208 Light
taxis; orientation in the opposition direction is a negative tropism or
taxis,
Occasionally a transverse tropism or taxis is displayed.
Control of speed of locomotion by the
intensity of the factor is
termed a kinesis. Gravity never changes significantly in intensity,
but the speed of swimming, flying, or creeping is influenced by altera-
tions in the strength of other factors; the common occurrence of
photokinesis has already been mentioned.
The position of the main axis of the body is the primary orientation
of the plant or animal. This usually determined gravity, as
is is
by
seen in the upward growth (negative geotropism) of the shoot and
the downward growth (positive geotropism) of the root of a
plant
seedling (Fig. 6.10). In the aquatic environment the buoyant action
FIG. 6.10. Negative geotropism of shoots and positive geotropism of roots of four
kernels of maize that have germinated in different positions. ( By permission from
Botany by Sinnott, 1929, McGraw-Hill Book Co. )
of water often reduces the effect of gravity so that primary orienta-

tion is to light or to current. When the source of illumination is
from the the primary orientation of a land plant will be a
side,
compromise between the influence of gravity and that of light.

The primary orientation of motile organisms is also usually to
or a flying animal continually adjusts its posi-
gravity, and a walking
tion to it. However, a fly on the ceiling or on a wall is quite uncon-
cerned as to which side up it is, since it is oriented primarily in rela-
tion to the surface upon which its feet rest. Sometimes orientation
is
brought about passively by the resistance of the appendages as
Orientation 209
gravity draws the organism through the medium, as is seen in a

"floating" butterfly or in slowly sinking plankton.
Superimposed upon the orientation,
primary many secondary re-
sponses to orienting influences take place. The tips of green plants

grow toward the light and hence exhibit positive phototropism. The
leaves are oriented at to the incident radiation,
generally right angles
thus receiving maximum illumination, and sometimes form a
they
symmetrical pattern (Fig. 6.11) and sometimes a mosaic, as is dis-
played by ivy leaves on a wall.
leaves on shoot of
FIG. 6,11. Alternating position and horizontal orientation of
Norway maple, resulting in maximum exposure to sunlight. (Shipley, 1925, after
Kerner, Copyright, Cambridge Univ. Press.)
In such species as the sunflower the top portion of the plant or its
leaves are turned by turgor changes during the course of the day,
in the direction of the sun. In regions where light
keeping always
and heat from the sun are excessive, the leaves of some plants, such
as the compass plant, Silphium laciniatum, are oriented so as to
toward the sun. These examples will suffice to
present their edges
210 Light
illustrate the
turning reaction of phototropism by actual growth or by
turgor movements.
Phototaxis, geotaxis, and other tactic reactions involve the orienta-
tion of locomotion toward or away from
light, gravity,
or some other
source of stimulation in the environment. In many instances the turn-
ing of the moving organism appears to be due to the unequal stimu-

lation of symmetrical sense receptors that control the tonus of the
body, leg, or wing musculature. When the animal moves, it is caused

to turn by the unequal posture of its locomotory apparatus. For
example, in a positively phototactic animal such as the swimming
insect Ranatra, if the left eye receives more light than the right eye,
the legs on the left side of the body will be more strongly flexed,
whereas the legs on the right side of the body will be more greatly
FIG. 6.12. (Upper) Symmetrical swimming position of positively phototactic

Ranatra moving toward a light source in front of it. (Lower) Position of normal
animal turning toward a light source at its left. This position will be produced
permanently if the right eye is removed; if both eyes are removed, the animal's
position becomes symmetrical again. (Crozier, 1929, Copyright, Clark Univ.
Press.)
Orientation 211
extended and will make longer
sweeps. As the animal swims in this
it will tend to turn toward the source
posture, of light. Turning will
continue until the amount of received in the two is
light eyes equal,
whereupon the legs will function symmetrically and the animal will
swim straight ahead (Fig. 6.12). In such instances the turn-
many
ing of the animal appears to be produced quite mechanically, and
such reactions were referred to as "forced movements"
by Jacques
Loeb (1918) who originated the theory of orientation just described.
In other instances orientation
appears to come about bya trial-and-
error procedure in which the animal changes its direction when it
encounters unfavorable conditions and tries other directions until it
finds a course in which conditions no
longer stimulate it adversely.
Fie. Arrangement of experiment to measure the geotactie orientation at

6.13.
of eaterpillar Malacosoma placed on
angle plane inclined at angle a. Lateral
movement of head indicated by h and ti. (Crozier, 1929, Copyright, Clark Univ.
Press.)
Whether one adopts or rejects a mechanistic viewpoint in inter-
preting the reactions of organisms, the fact is that tropistic and

tactic responses of plants and lower animals
play a major role in
their livesunder natural conditions, and their orientation often ap-
pears to be rather rigidly controlled. An example of the mechanical
way in which an orienting force may act is provided by an experi-
ment involving the negative geotaxis of the tent caterpillar, Mala-
cosoma. If this
caterpillar is placed horizontally on a steeply sloping
surface, an unequal stimulus of the proprioceptors located within
the two sides of the body will result, owing to the animal's weight.
A differential response in the tonus of the body muscles will then be
produced, with the result that, as the animal creeps, it will turn to
move up the slope (Fig. 6.13). As the caterpillar turns from its
initialhorizontal position toward the vertical, the difference in stimu-
lation of the two sides of the body becomes progressively less, and
eventually reaches a threshold beyond which no further turning is

elicited. If the plane upon which the caterpillar is creeping is only
212 Light
slightly inclined (angle a), threshold discrimination is reached after

a smaller amount of turning, or at a smaller final angle of orientation
(6). When
the plane is steeper, the animal continues turning to a
greater angle of orientation. The result of a series of trials made

with varying angles indicates the mechanical nature of the taxis and
the quantitatively exact way in which the response of the animal is
controlled by the magnitude of the orienting stimulus (Fig. 6.14).
90
80
90 70
s?
80 60
S 70 50
60 40
50 30
7
40
30
10
7 20 30 40 50 60 70 80 90
Inclination of surface, a, degrees
FIG. 6.14. Relation between the angle of inclination of plane (<*) and the mean
angle of orientation (0) of caterpillar creeping upon it. Open circles are means
for one individual; black circles are means for all individuals tested. The curve
is that for A0/A log sin
<* constant. ( Crozier, 1929, Copyright, Clark Univ.
Press. )
Orientation of growth and locomotion in relation to light and other

factors plays a profoundly important role in the lives of plants and
lower animals. The responses of all species that survive must ob-
viously generally bring the organism into favorable surroundings, but
lack of flexibility in these reactions often produces harmful results
under extreme or unusual conditions. The rigidity of the tactic
responses of the tent caterpillar is often seen to lead to its destruction.

When the animals emerge from their nests in the crotches of trees,
they crawl upward in response to their negative geotaxis to the tips

Orientation 213
of the branches where no doubt the juiciest leaves are found. The
caterpillars later may return to the nest by following the silk threads
they have left behind them on their upward journey, but they will
not ordinarily move further downward over fresh surfaces. The two
wild cherry trees shown in Fig. 6.15 have been completely defoliated
FIG. 6.15. Wild cherry trees in southern Rhode Island defoliated by tent cater-
pillars (Malacosoma), whose nests are seen in the forks of the branches. Neigh-
boring oak trees were not attacked.
tent caterpillars. Following the consumption of all the leaves in

by
their tree,many caterpillars starve, or die of disease (Craighead,
1950) but others crawl or fall to the ground and eventually find their
way to another tree. For some reason, as yet unknown, this species
does not ordinarily eat oak leaves, and in the area photographed the
oak trees only a meter or so away were not attacked. As the time for
pupation approaches, negative geotaxis becomes strong and the ani-
mals tend to climb up any vertical object encountered. In a neighbor-
ing area a series of fence posts was found,
on the top of each of which
was a seething mass of tent caterpillars that had gathered there as a
result of the reactions just described. Since the animals were slaves to
their geotaxis, they remained on the fence posts, and, finding nothing
else to eat, they proceeded to devour each other.
When an animal moves vertically under normal illumination from
above, it is often difficult to ascertain whether the animal is
reacting
or to light; that is, if it moves upward, may be
primarily to gravity
it
214 Light
displaying a negative geotaxis, a positive phototaxis, or both. Ex-
periments can be devised that distinguish between the responses
of which the animal is
capable. An example of such an experiment is
given in Fig. 6.16 in which the response of a negatively geotactic
and a negatively phototactic Agriolimax is shown. When light is
allowed to act at right angles to gravity, this lowly garden
slug can
resolve the forces with more alacrity than is displayed
by some col-
lege students! The first trial indicated in the diagram was made
when the animal was dark-adapted and its reaction to light was
stronger than that to gravity. A decrease in the strength of photo-
taxis as the animal became
light-adapted is indicated by the change
in the angle of orientation in successive trials.
\
V I
FIG. 6.16. (Upper) Orientation of negatively geotactic and negatively photo-

tactic Agriolimax on a vertical plane with light from the right. L = phototactic
vector, g geotactic vector, ~
angle of orientation. (Lower] Successive trials
at 1 -minute intervals made by on a vertical
initially dark-adapted Agriolimax
plane. (Crozier, 1929, Copyright, Clark Univ. Press.)
Other complications occur in the orientation to light. Diffuse
light has been shown to exert an effect upon planktonic copepods that
is different from that
produced by light from a single source. Many
plankters are found to be phototactic to an electric light,
positively
Orientation 215
but these animals do not
generally swim upwards to the surface of
the water toward the sun since the diffuse side illumination in the
ocean inhibits the reaction to a single source (Schallek, 1943).
The familiar "bee-line" course which the honeybee takes from its
hive to a source of food is determined as a definite
angle to the direc-
tion of the sun. The bee does not fly directly toward or away from
the sun, but orients at an angle to the
changing azimuth of the sun.
Upon returning to the hive, the bee communicates to its fellow work-
ers by a special "wagging dance" both the
approximate distance and
the direction relative to the sun of the food that it has discovered.
Still more remarkable is the fact that the worker bee is not required
to see the sun directly but can be oriented
by a small patch of blue
sky. The bee's compound eye has been shown to be sensitive to the
angle of polarization of sky light, and the beeapparently uses this

information in combination with other orienting forces to determine
the proper line of flight when the sun is obscured. The bee usually
restricts its visits to one species of plant at a time a fact that is of
obvious advantage both to the bee's efficiency and to the plant's
successful pollination. Flower species are recognized by shape, scent,
and color; the bee's eye can distinguish yellow, blue-green, blue, and
ultraviolet, but it is blind to red (
von Frisch, 1950 ) .
The ability of homing and migrating birds and fish to find their
way over long distances perhaps even more amazing. The precise
is
method of navigation used by such far-ranging vertebrates has mysti-

fied observers for generations. Although birds may use landmarks,
persisting cloud formations, prevailing winds, and other ecological
cues to some extent, these possibilities do not appear to explain all
instances (Griffin, 1952). Evidence has been obtained that certain
species of birds may use the direction of sunlight as a means of
orientation and that the birds are able to allow for the change in the
sun's position during the day (Kramer, 1952).
Under natural conditions the direction of growth or of locomotion
of plants or of animals is usually determined by several different in-
fluences acting simultaneously. In addition to such common orient-
ing factors as light, gravity, temperature, and moisture, certain species

may be guided by sound or other types of vibration, by scent or other
chemical sense, or by other special reactions to environmental
stimuli. We know that tropistic and tactic responses play a very large
role in the lives of most lower organisms in nature. Reactions to the

direction and
intensity of the orienting influences of the habitat are
getting motile organisms where we find
responsible for them. In
order to understand the mechanism of distribution of these forms,
216 Light
it is
necessary to analyze the reactions involved. Many practical
applications of such knowledge suggest themselves, such as the control
of insect pests.
In attempting to interpret the
locomotory reactions of organisms
and to apply them to natural situations, the
following considerations
should be kept in mind: (1) Responses to individual stimuli are
usually limited by the influence of other conditions. Thus the posi-
tively phototactic moth does not attempt to fly to the sun. (2) The
observed distribution may be the result of movement without direct
orientation. The reduced locomotory activity of wood lice in situa-
tions where moisture
is
high results in a tendency for these animals
to congregate moistinplaces although they are not specifically
oriented to them, just as the traffic on a main highway becomes
denser in a bad stretch of road because the speed of the cars is
reduced. (3) For an oriented response the orienting factor must
provide a stimulus above the threshold of sensitivity, and either the
direction of the flux of the orienting force or the direction of a gradient
produced by it must be
perceptible to the organism. The tempera-
ture gradient in the ocean, for
example, may be shown to be below
the threshold for the response of zooplankton (Clarke, 1934). (4)
Factors whose gradients are below the threshold often exert an in-
direct effect by controlling the sign or the speed of the response to
another factor. (5) Responses may similarly be altered under
changed internal physiological conditions, as before or after feeding,
or breeding. Within the same species the males, females, and various
immature stages sometimes orient quite diversely. Thus, in gen-
eral, we see that the distribution of plankton in the water, of microbes
in the soil, or of insects in the vegetation is the result of a complex
interplay of the orienting factors of the environment acting directly

and indirectly on the changing physiological state of members of the
population.
Periodicity
Diurnal Periodicity. A
good number of the fundamental rhythms
in nature are related to the light factor. Many animals and plants
exhibit a 24-hour cycle in their activities; this has long been known
as diurnal periodicity. However, the term diel periodicity may be
substituted if confusion arises from the fact that diurnal is also used
for daytime activity as opposed to nocturnal for nightime activity.

The most fundamental diurnal rhythm that of photosynthesis itself,
is
which necessarily fluctuates because of the daily change of light.
Many plants exhibit other more specialized reactions to the alterna-

Diurnal Periodicity 217
tion of day and night, such as the opening and closing of flowers and
the folding of leaves.
Although often not conspicuous, a very large number of the lower

animals are controlled in their activities by the in light from
change
simple photokinesis. The
day to night. This often a matter of
is
animals are more active in the light, less active in darkness or vice
versa, but behavior may be modified by concomitant diurnal

also
changes such as temperature and humidity. Among

in other factors
the higher animals the reactions are often more elaborate, and some-
times the daily rhythm of activity is only indirectly related to light.
Many of the relatively defenseless forms, such as mice, come out to

forage chiefly at night when they are less likely to be detected by their
enemies. As a further complication many of the predaceous animals
are nocturnal as an adaptation to the night activity of their prey or
to the fact that they can stalk their prey more successfully under
cover of darkness. A patch of woods may be inhabited by two sets
of animals which practically never meet because one set is active
only during the day and the other active only during the night (Park,
1940).
The fact that a diurnal rhythm in the activity of an organism is
sometimes deeply ingrained can often be demonstrated by bringing
the organism into the laboratory and observing it under constant or
under changed conditions of illumination. In one such experiment
the deer mouse, Pcromtjscus, which is normally active at night and
quiescent during the day, continued to display a diurnal rhythm in

its behavior after seven months in continuous darkness. This and
other experiments summarized by Welsh (1938) and by Park (1941)
indicate that certain internal physiological processes have become
attuned to a 24-hour cycle. In many organisms this rhythmicity per-
sists for
long periods and tides over periods when the usual controlling
environmental stimulus fails to occur. Under normal conditions the
timing of the periodicity is re-enforced by changes in light or other
factors each day.
The diurnal shift in the activity of animals frequently results in
significant changes in their position

in the community. Perhaps no
better illustration of this phenomenon could be found than the vertical
migration of zooplankton in the sea and in lakes. In general, vast

numbers of copepods and other planktonic animals tend to swim to-
ward the surface at night and to move downward to
deeper levels
during the day. The diurnal changes in vertical distribution of fe-
males of the copepod, Calanw, in the Clyde Sea area are indicated
in Fig. 6.17. The time of ascent in the evening and of descent in the
218 Light
morning, as well as the depth to which the animals migrate, is corre-

lated with the differences in time of sunset and sunrise and in the in-
tensity of the noon sun during January and July. The complexity of
the response to light is revealed by the fact that in this same area, the
males and the various young stages of the same species exhibit quite
different patterns of behavior.
25-26th January
4ipm 7 pm 10 pm 1 am 4 am 7 am i 10 am 1 pm 4 pm
!M2thJuly
50 20 20 50
Fie. 6.17. Diurnal vertical migration of female Calanus finmarchicus in the Clyde
Sea area during 24-hour periods in January and in July. The abundance of the
population is indicated by the width of the figures. The times of sunset and
sunrise are indicated by arrows. (Nicholls, 1933, Copyright, Council of Marine
Biol. Assoc. of United Kingdom.)
Reactions to gravity and to temperature change often modify the

course or the extent of the vertical migration (Clarke, 1933). The
various theories dealing with the reactions of the zooplankton to the
factors believed to control vertical migration have been summarized
by Gushing 1951 )
(
The magnitude of the migration is indicated by
,
the fact that the copepod population may often move more than 100
m in a vertical direction. This represents a journey of more than
15,000 times the animal's own length twice each day. Even more ex-
tensive diurnal vertical migrations are carried out by euphausiids,
fish, and perhaps other large active forms. The widespread occur-
rence of changes in level of huge populations in the open ocean, dur-
ing the course of the day, has been revealed by midwater

echoes re-
ceived by the underwater acoustical equipment of ships. A discus-
sion of the kinds of animals probably responsible for this "deep scat-
tering layer" and its vertical oscillation is presented by Moore ( 1950).

Similar vertical movements occur in terrestrial environments al-
though these are generally less extensive. Many animals move from
the surface to deeper levels in the soil at regular
periods each day.
Others emerge from under the ground litter at definite times during
the diurnal cycle and ascend the vegetation. Harvestmen, or
"daddy-
long-legs" (Leiobunlum rotundum), in an English oak wood were
observed to descend from the tree trunks in the evening to the forest
floor where they hunt their
prey and to move up onto the trees again
in the early
morning (Todd, 1949). Many other insects move reg-
ularly from lower levels in the herbs and shrubs to higher positions
in the trees atdawn and return at dusk; others migrate in the reverse
manner. These changes in levels of whole populations, both in the
water and on land, have profound repercussions on prey-predator
relations and other interdependencies among the inhabitants. For
a further discussion of these aspects of stratification and periodicity
in the
community, the reader should refer to Allee et al. (1949, Ch.
28).
Lunar Periodicity. Since the days of classical Greece interest has
been attracted to the correlation of certain animal activities with pe-
riods of the moon. Oppian in the time of Aristotle wrote:
The shellfish which creep in the sea are reported, all of them when the
moon waxes, to fill up their flesh proportionately to her disk, occupying then
a bigger space, On the other hand when she wanes they shrivel and their
members grow thinner.
It is Red Sea the gonads of the sea urchins ( the edible

true that in the
portion) do enlarge during the period of the full moon, but the be-
lief was
spread fallaciously around the Mediterranean and elsewhere
that the lunar cycle controlled many more animal activities than is
actually the case. One even meets the statement occasionally that
the 28-day menstrual period in man harks back to our marine an-
cestors. Since we know
that the menstrual cycle in other mammals
has very different periods, the agreement of the human period with
the 28-day lunar cycle is merely a coincidence.
Modern investigation has shown that the activity of certain widely
different types of organisms, usually in relation to the reproductive
cycle, shows a correlation

with the moon. The striking nature of
these lunar periodicities is well illustrated by the fluctuation in the
abundance of conjugants produced by a ciliate living as an ectopara-
site on the
gills
of a fresh-water mussel (Fig. 6.18). The distinct
peaks occurred regularly on the days following the new moon and
were not correlated with temperature or other known environmental
changes.
220 Light
19 19 17 17 15 14
100
20
90 20
80
03
18
70
22
50
40
30 -
20 -
15
10
-
Sept. Oct. Nov. Dec. Jan.

i
Feb.
FIG. 6.18. Average daily number of conjugants of the ciliate Conchoplrthirius

lamellidens on the gills of the fresh-water mussel Lamellidens marginalis. Dates
of new moon (top) and of peak numbers are shown. (Ray and Chakraverty,
1934, Nature.)
Most of the organisms exhibiting lunar periodicities are marine and

hence may be affected by the amplitude of the tide which is greatest
at times of new moon and full moon (spring tides) and smallest at
the times of the quarter moon (neap tides). For organisms living
within the influence of the tides it is
obviously difficult to determine
how much may be
of the effect due to moonlight itself and how much
to the action of the tidal cycle. The marine alga, Dictyota, for ex-
ample, produces its
gametes at the time of the full-moon spring tide.
The spawning of a number of marine polychaete worms shows various
time correlations. The palolo worm inhabiting the waters of the
South Pacific islands comes to the surface in great numbers on the
last quarter of the moon during October and November, producing a
luminescence and discharging eggs and sperm into the water. The
natives know of the occurrence of this swarming of the palolo worm
and take advantage of the opportunity to scoop up large quantities
of these animals for food. The worms swim about in small circles in
dense masses, giving the sea an appearance of spaghetti soup. The
natives gather the worms in crude baskets and celebrate the occasion
with religious rites and feasts.
The Bermuda fireworm puts on a similar display of fireworks in the

shallow water early in the evening at the time of full moon. The
reaction appears to be set off by the drop in light intensity
following
sunset (Huntsman, 1948). Professor E. L. Mark, who was for
years
director of the Bermuda
Biological Station for Research, took a special
interest instudying the swarming of this marine polychaete. He
found that swarming and luminescence began at about 55 minutes
after sunset and lasted for half an hour. On one occasion when Pro-
fessor Mark was traveling to Bermuda he realized that he would ar-
rive on a day when the fireworms were due to appear. He interested
his fellow passengers in the
phenomenon, telling them that the timing
was so exact that one could set one's watch by it and invited a large
group of people to come to the shore that evening. When the moment
arrived for the beginning of the display, not a worm was in sight,
nor was there after 5 minutes, 10 minutes, and more. Soon people,
feeling that they had been hoodwinked, began moving away, but 20
minutes after the appointed time the worms appeared in great num-
bers and a spectacular demonstration was given for the visitors who
still remained. Not until the next day was Dr. Mark's embarrassment
relieved when it was discovered that during the winter Bermuda
time had been changed from local sun time to zonal time, resulting
in setting the Bermuda clocks ahead 19 minutes.
Perhaps the most fascinating of all the responses of animals to the

lunar or tidal cycle is that exhibited by the grunion or California
smelt. On the three or four days after the spring tides from April to
June the grunion swims in on the beaches of southern California to
lay eggs in the sand. During this period the higher of the daily
its
high tides comes at night. About an hour after the water reaches its
highest level on each of these nights the fish allow themselves to be
carried up on the beach by the waves. As each wave recedes, an
observer can discern hundreds or thousands of fish left behind, wrig-
gling in thewet sand (Fig. 6.19). Within a few seconds the females
have burrowed tail first into the sand where they deposit the eggs
while the males curl around them discharging the sperm. When the
next wave comes in, the fish are washed out to sea once more. Since
the eggs have been discharged during the period of descending tides
after the highest spring tides and during the hours following high
tide each night, the waves do not reach them again during the next
two weeks. The eggs are thus left undisturbed to develop in the
warm, moist sand. When the next spring tide occurs, the eroding
surf of the rising tide uncovers the eggs, now ready to hatch out, and
the young larvae swim away. In order for this complicated relation-
of the fish and the tides to be carried out,
ship between the life cycle
the fish, which ordinarily remain in the waters offshore, must be
stimulated in some way to move in to the beach on the proper
day
and to allow themselves to be stranded on the shore by the waves
222 Light
FIG. 6.19. Flashlight photograph of grunion (Leuresthes tenuis) spawning on

Malibu Beach near Los Angeles, California. The females bury themselves tail
first in the sand. The males then circle around them and fertilize the eggs.
(Photo by Moody Institute of Science.)
only during the proper hour, Just how this timing is controlled has
not yet been discovered despite a considerable amount of study. The
complexity of the interplay of the tidal and lunar influences and the
remarkable precision of the responses of the grunion are indicated in
Fig. 6.20.
The foregoing are representative of the varied occurrences of lunar
periodicities in nature of which many more instances could be cited

(Korringa, 1947). A
definite control of the timing of the
reproduc-
tive cycle is usually indicated, and
frequently other special activities
such as swarming or luminescent are involved. The mech-
discharge
anism by which the timing is controlled is far from clear, however.
For some organisms the changing amount of moonlight may be of
chief importance either directly or as a degree of contrast with sun-
light or starlight. Since the intensity of full moonlight is more than

1(H of noon sunlight, it is well above the threshold for the response
of many organisms. As an alternative the cyclic change in the rela-
tion of the moon's gravity has been suggested as
possibly controlling
the activity of fish and other organisms. Many sportsmen
strongly
believe that such lunar gravitational fluctuations influence the success
of their fishing. At the present time, however, we have no evidence
224 Light
that the sense
organs of any type of animal could respond to the dif-
ferences in the moon's gravitational effect.
For marine animals of the littoral zone the moon's influence often
appears to be mediated at least in part through the change in the
tides, but certain lunar periodicities occur in regions where tides are
slight or lacking. One great difficulty in studying this problem is

that often the behavior
pattern persists for a while under changed
conditions, as when the moon is obscured, apparently because of a
correlation with an internal rhythm. Much more investigation will
be required before these intriguing periodicities are completely un-
derstood.
Seasonal Periodicity. The seasonal activities of plants and animals
are sometimes due to changes in
temperature and are sometimes con-
trolled by the cycle of dry and rainy seasons; seasonal periodicity may
also be influenced by the light factor. The seasonal effect of light
is much due to differences in intensity as it is to differences in
not so
total amount of and in the relative lengths of day and night.
light
The response of organisms to daylength is known as photoperiodism.
The most striking manifestation of photoperiodism is the control of
the reproductive cycle in certain plants and animals, although other
life
processes are involved in some instances. In these organisms the
reproductive phase of the life cycle is initiated by days that are shorter
or longer than certain critical lengths. Under natural conditions short
days are accompanied by long nights and vice versa. Experimental
manipulation of the photoperiod has revealed facts indicating that the
length of the night may be more influential than the length of the day
in
controlling the photoperiodic responses of at least some of the
plants and animals, but the precise mode of action of the light factor
has not yet been determined (Farner et al., 1953).
Photoperiodism was discovered in relation to plants. Species

first
that flower only when

the days are longer than a certain number of
hours and the nights are correspondingly short are known as long-
day plants. Short-day plants, on the other hand, flower naturally only
under conditions of short days and long nights. However, the re-
productive cycle of certain other species is not affected by daylength;
these are referred to as indeterminate or indifferent plants. This
aspect of periodicity in the light factor controls the season at which

long-day and short-day plants flower in any locality, and it also in-
fluences the geographical distribution of the species.
Long-day plants flower naturally only in the middle or high lati-

tudes during the late spring or early summer. Familiar examples
are the radish, iris, red clover, evening primrose, spinach, the smaller
and timothy. If such plants are
cereals, grown under daylengths
shorter than the critical
photoperiod, stems tend to be shorter and
flowering suppressed (Fig. 6.21). Other effects may also be pro-
is
duced in the
plant, as illustrated by long-day potatoes which produce
the best tubers when daylength is below the for shoot
optimum
growth.
FIG. 6.21.Control of flowering and of vegetative growth by daylength in
timothy, a typical long-day plant. The daily exposures (in hours) to light are
indicated on each container; C natural daylength. Photo taken in July.
(Evans and Allard, 1934.)
In the growing season outside the tropical zone short days occur
both in the early spring and in the late summer. Certain short-day
plants require a long growing period before they
are sufficiently
mature to react to the flowering stimulus, and such plants can flower
only in the shorter days following the summer solstice. Familiar
plants that can bloom naturally only
late in the year are tobacco,
goldenrod, aster, dahlia, ragweed, cosmos, and chrysanthemum. In

daylengths above the critical abnormally great vegetative growth
takes place and flowering is much delayed or entirely inhibited (Fig.
6.22). Short-day onions and beets develop the largest storage organs
under photoperiods that are longer than those best for the growth of
the upper part of the plant.
The plants that bloom in the short days of early spring are mostly
perennials in which the flower buds
were set the previous autumn,
The few annuals blooming at this season either germinated during
the winter or are so small that they need little time for vegetative
226 Light
FIG. 6,22. Control of flowering and of type of vegetative growth by daylength

in garden chrysanthemum, a typical short-day plant. The daily exposures (in
hours ) to light are indicated on each container; C = natural daylength. Photo
taken in July. Flowering occurred as follows: 10 hr, July 6; 12 hr, Oct. 14; 13
hr, Oct. 19; 14 hr, Oct. 31; natural day, Oct. 31. (Allard and Garner, 1940.)
growth before flowering. Early spring flowers growing on the floor

of the deciduous forest are able to take advantage of the great amount
of light available before the tree leaves are fully developed.
Since short days occur later in the summer at higher latitudes, the
northward distribution of short-day plants may be stopped by the
lack of sufficient time for the ripening and hardening of the seed be-
tween the occurrene of the critical daylength and the occurrence of
a killing frost that terminates the growing season. Ragweed, for
example, which flowers when the days have shortened to 14% hours,
will begin reproductive phase about the first of July in Virginia,
its
and hay-fever sufferers notice its pollen in the air by the middle of
August. In northern Vermont the summer days are not reduced to

14% hours until after the first of
August, with the result that seed
formation is not completed before cold weather, and consequently
ragweed is unable to establish itself in abundance at this latitude.

The distribution of long-day plants toward the equator is limited by
the reverse action of this aspect of the light factor. Species of Sedum
that require a daylength of 16 hours or more will flower abundantly
in Vermont but bloom in Virginia (Naylor, 1952).
will not
Plant species in which flowering is not controlled by daylength
also occur at all latitudes, but all plants growing at high latitudes must
be able to tolerate long days and short nights since these light condi-
tions exist during the growing season, and tropical plants must be
able to complete their
development under photoperiods of about 12
hours* duration. Since the vegetation of intermediate latitudes is
composed of a mixture of long-day, short-day, and indifferent plants,
the seasons of vegetative growth, and seed dissemination
flowering,
the species of the
vary among community, and competition is reduced
accordingly. In some instances genetic strains of the same species
growing under different climatic conditions respond to
differently
given photoperiods, apparently, as an adaptation to differences in the
time of onset of adverse conditions. The of
possibility intraspecific
variation of this sort plants and animals in their response
among both
to all types of environmental factors must
constantly be kept in mind.
The control of the reproductive cycle by the length of day is a re-
sult of acomplicated balance between life processes that go on in the
light and in the dark
(Leopold, 1951). Since the intensity of light
which influences the photoperiodic response is far below that needed
for photosynthesis, we know that the 'reactions responsible are quite
different. The delicate nature of the processes involved are indicated
by the fact that only a few minutes, or even a few seconds, of light
during the middle of a long night may reverse the flowering reaction
of the plant. The tasseling of sugar cane can be inhibited by a short
light period during the night, and the possibility of retarding the
ripening of the crop until the most favorable time for harvesting by
sweeping the cane fields with search lights has been investigated by
Hawaiian planters. In greenhouse plants and other commercial
species the timing of the production of flowers, seeds, fruits, or storage
organs is controlled artificially by means of the photoperiodic re-
sponse.
Photoperiodism similarly plays an important role in the life cycles
of many kinds of animals and, interestingly enough, also most fre-
quently controls the reproductive cycle. However, wing production
in aphids,metamorphosis in mosquitos, pelage changes in fur bearers
(Lyman, 1942), and other life processes are known to be influenced
by daylength. The most extensive studies have been carried out on
mammals and birds, and these have been reviewed by Bissonette
(1936) and Burger (1949). Trout that ordinarily spawn in Decem-
ber can be induced to lay their eggs in August by artificially chang-
ing the daylength. The fresh-waterpulmonate snail, Lymnaea
palustris, will not lay eggs
on an 11-hour day but lays abundantly
when the days are 13% hours long or longer. The control of repro-
duction by daylength in this species rather than by temperature was
neatly demonstrated under
natural conditions by the fact that snails
living in a spring with practically

no seasonal change in temperature
228 Light
began laying eggs during the same week as did snails inhabiting a
shallow ditch that underwent
rapid vernal warming (Jenner, 1951).
Very additional investigation of photoperiodism among the in-
little
vertebrates has been made, but this reaction to

light may eventually
be found to play an important role in controlling the life cycle of
many of the less

conspicuous animals. Among mammals certain
forms, such as ferrets, are brought into breeding condition by the
long days of spring, but others, such as deer, come into heat during
the short days of autumn.
Photoperiodism in birds was first discovered as a result of studies

It had
of migration. long been realized that the arrival of birds in
the spring was not closely related to temperature or other common
aspects of the weather since birds are often caught by late winter con-
ditions. A similar difficulty arose in explaining the start of the south-
ward migration. Many birds leave the northern regions during July
or August when the temperature is
high,and when an ample food
supply is still available (Fig, 6.23). The length of clay was finally
Photo by Allan D. Crmckshank, National Audubon Society

FIG. 6.23. Tree swallows flocking previous to migration.
recognized as one environmental factor that changed regularly year

in and year out and that could account for the exact
timing of the
migration regardless of the weather conditions. Present evidence
indicates that the effect of the
daylength is
brought about through
controlling the amount of light
reaching a sensitive tissue in the
bird's body (the pituitary gland). Under natural conditions the
larger amount of light received during the long days of spring causes
an increase in the size of the gonads and also, in some way as yet un-
known, sets off the northward migration. Following the breeding
season the diminishing daylengths of late summer and autumn cause
a decrease in gonad size and provide the stimulus for the southward
migration. Evidence on the mode of action of light and other factors

on bird migration has been summarized by Farner (1950).
Although photoperiodism thus appears to be basically involved in
the annual stimulus for the migration of birds, it does not explain the
evolutionary origin of the migratory habit nor, most mysterious of

all, how the birds find their way. Once established, however, migra-
tion would tend to advantages. At higher lati-
be perpetuated by its
tudes the longer days make more time available for nest building
and for feeding the young. The young grow more rapidly, and hence
the whole process of rearing the family, with its attendant dangers,
is
completed more quickly than would be the case nearer the equator.
By leaving the north during the winter, on the other hand, the birds
avoid the hazards of low temperatures and shortage of food.
The migration of birds is evidently a secondary effect of the in-
fluence of light. The primary effect is in the control of the breeding
cycle since this can be demonstrated in species of birds that do

not
migrate. The occurrence of breeding in the English sparrow at dif-

ferent latitudes over the face of the earth is indicated in Fig. 6.24.
Near the equator some breeding takes place in every month of the
year. In areas farther north and farther south reproduction tends to
be confined to the months of the year with longer days. North of
50 N latitude breeding is strongly centered in the month of May with
no breeding at all during the months of short daylength, and south
of 50 S latitude most breeding takes place in December. Outside of
the tropics, therefore, the light factor plays an important role in con-
to the months of the year when reproduc-
fining the breeding period
tion most successfully carried forward. Since short daylength pre-
is
vents birds from entering the breeding condition too early and the
long days of midspring act to speed up the laggards, the whole popu-
lation tends to breed within the same favorable period ( Bartholomew,
1949).
230 Light
1 " ! IV v v' ix x XT xii i ii in iv v vi
. ." , , , v'lyniT 1 1 T 1 \ 1 1
10 20 N
0- 10 Ni
0-10S.
10 -20Si
20 -30 Si
30 -
40 $
40 -
50 S
50 60 S
J L
Vm
1 ' ' 1 ' ' 1 '
v
'
i ii in iv vi vii ix x xi 'xn i n m' iv v vi

FIG. 6,24. Relative intensity of breeding of the English sparrow in each month of
the year at the indicated latitudes. (Baker, 1938.)
Ultraviolet Light
The small fraction of the sun's radiation that reaches the earth as
ultraviolet light has certain
very special biological effects, and some
of these may be of
ecological significance. The sensitivity of the bee's
eye to this part of the spectrum and its use in orientation has already
been mentioned. The population size of certain land animals in north
central United States was found
by Shelford (1951 and 1951a) to be
correlated with the intensity of ultraviolet
light, although the reactions
that underlie the correlation are not known. The bactericidal
action of ultraviolet is familar and causes the destruction of micro-
organisms that are excessively exposed to the direct rays of the sun.
Ultraviolet radiation produces sunburn, or erythema, in man and
per-
haps causes harmful effects in other animals under natural condi-
tions, althoughquestion has not been investigated from the
this
ecological viewpoint. Ultraviolet also brings about the production of

vitamin D
with its antirachitic effect. Vitamin is formed
by the D
irradiation by ultraviolet light of certain sterols, or
fatty substances,
Ultraviolet Light 231
that occur in both plants and animals. The relative effect of different
wavelengths in producing the foregoing influences is shown in Fig.

6.25. Although the curves differ very much from one another
throughout most of the range, it is clear that in no instance is ultra-
violet radiation effective at
wavelengths longer than about 3100 A.
250 260 270 280 290 300 310 320

Wavelength
FIG. 6.25. Relation between wavelength in the ultraviolet region and the bac-
tericidal, erythemic, and antirachitic effects of ultraviolet radiation, together with
the absorption of ergosterol. ( Coblentz, 1939, Copyright, J.
Am. Medical Assoc. )
Under ordinary conditions no measurable radiation from the sun

reaches the earth's surface at wavelengths shorter than 2950 A. It is
clear then that under natural conditions the whole action of ultraviolet
must occur between 3100 A and 2950 A. Under the best circum-
stances solar radiation is
extremely weak and when dust
in this region,
or smoke is abundant the shorter wavelengths are still further re-

232 Light
duced. During the winter in the
neighborhood of cities no measur-
able radiation shorter than 3100 A can be found, and hence no
ap-
preciable action of ultraviolet can take place at that season. When
birds that ordinarily migrate south were retained in Alberta, they
died during the winter apparently from lack of vitamin D. Neverthe-
less, we have little evidence that the lack of ultraviolet in winter is
generally of crucial ecological importance, but the matter should be

investigated further.
In the aquatic environment ultraviolet is still further reduced by
the very rapid rate at which it is absorbed even in the clearest water.
Although in some quarters popular imagination has fixed upon the
idea that ultraviolet is the most penetrating region of the spectrum,
actual measurements have shown that radiation at 3100 A is reduced
to 10 per cent of its surface value at a depth of 15 m
in the extremely
clear water of the Mediterranean and at 1 m
in Gullman Fjord on the
coast of Sweden. The and biologically more active wave-
shorter
lengths are reduced even more rapidly by absorption and scattering

in the water (Jerlov, 1951). In most natural waters the effectiveness
of ultraviolet diminishes very rapidly with depth. Carefully con-
trolled experiments have revealed no bactericidal action below 1 m
and probably none at depths greater than 20 cm even in clear ocean
water. The
ecological effect of the lethal action of ultraviolet in very
shallow water awaits further study. It is possible that the bacterial
population on the bottom mud in the littoral zone is seriously affected
when exposed at low water.
How can we account for the formation of vitamin D in fish liver
oils? Cod and whose oils are particularly rich in vitamin D
halibut
live far below the depth at which any measurable ultraviolet could
penetrate. It has been

suggested that the vitamin D might reach
the fish livers through a food chain starting with the phytoplankton
and the zooplankton that live near the surface of the sea. Since cod
and halibut are bottom feeders, a long food chain must be postulated
to extend to the forms on which these fish live. Another suggestion
is that the vitamin D formed in the Sargassum weed as it floats near
the surface of the tropical oceans finds it way via the food chain into
such species as the cod. For the most part we know that the animals
associated with the Sargassum weed do not eat the weed directly but
live on plankton or materials accumulating on the surfaces of the
fronds. In addition we know that a sharp temperature barrier exists
between the regions where the Sargassum weed is abundant and
areas inhabited by such fish as the cod and halibut. The amount of
vitamin D in the liver oils of these species is very great. It seems in-
Land Plants 233
conceivable that the supply could have resulted from such tenuous
food chains as are described above, even if the fish were able to retain
mammals are not. Nor has any method
the vitamins in their bodies as
been shown by which an animal could manufacture vitamin D with-
out the presence of ultraviolet light. The actual source of the large
quantities of vitamin in theD fish liver oils thus remains a mystery.

It
emphasizes again how little we know about ecological influences
of this region of the spectrum, both in the water and on land.
Ecological Aspects of Photosynthesis
Light is
fundamentally important as the essential direct source of
energy for the growth of all
green plants and the purple bacteria.
The photosynthetic plants form the
first
step in the ecological cycle
in every natural situation. They are the first link in the food chain
and hence are the base of the production pyramid that will be dis-
cussed in greater detail later on.
Land Plants. On the surface of the land enough light is received
everywhere for the growth of plants of some sort. Within the soil
and in the interior of caves light is insufficient to allow photosynthesis.

A striking illustration of the limiting action of
the light factor is found
in caves to the walls of which electric lamps have been secured for
the benefit of tourists. Within the cone of light immediately around
each lamp mosses have grown from spores brought in by air currents.
These "islands" of plant growth present a sharp contrast to the com-
plete lack of vegetation in the
remainder of the cave.
Although illumination is generally sufficient for photosynthesis over

the entire surface of the globe, the local distribution of individual
species of plants is definitely influenced by differences in the avail-

ability of light. Species that can grow in shady places are termed
tolerant, and, although the degree of tolerance is affected by soil
moisture, temperature, and other factors, the illumination is the prin-
cipal controlling influence. Plants that require strong illumination

and will not survive or develop in reduced light are referred to as
intolerant species.Trees such as the spruce, hemlock, beech, and
sugar maple, shrubs, such as spicebush, and herbs, such as blood-

root, can grow in deep shade and are examples of tolerant species.
The sugar maple can photosynthesize adequately in situations where
the illumination is reduced to less than 2 per cent of full
sunlight.
Birch trees, poplars, willows, and several species of pine, as well as
many shrubs and herbs such as sumac, bluestem grasses, and sun-
234 Light
flowers are intolerant of shade. The ponderosa
pine, for example,
requires a light intensity equal to at least 25 per cent of full sunlight
Intolerant species cannot develop in the shade of a dense stand
of their own or other species. Ecological consequences of this fact
for the
community
plant will be discussed further
in later chapters.
Even among somewhat tolerant species the first
plants to become es-
tablished in an open area will grow to better
advantage than indi-
viduals subsequently developing around them because of the com-
petition for light and other needs. Differential growth often results
in "pyramiding" in the development of such a
group of plants (Fig.
6.26). As we have seen in an earlier section, the light on the floor
FIG. 6.26. Pyramiding due to competition exhibited by a group of spruce trees
growing in a Michigan bog.
of the temperate deciduous forest becomes seriously reduced when
Many of the smaller plants are adapted to a period

the trees leaf out.
of rapid growth in the early spring while light on the forest floor is
sufficiently intense. In the marginal rainforest of Panama a dense
undergrowth is
possible because during the dry season the leaves fall
from the trees and allow light to penetrate. In the equatorial rain-
forest of Columbia, on the other hand, trees shed their leaves in-
Aquatic Plants 235
dividually and a continuous canopy is maintained, with the result

that little vegetation can grow on the forest floor.
In such rainforests many smaller plants have gained access to suf-
ficient illuminationby the evolution of the epiphytic habit, that is,
by growing in the crotches or on the horizontal branches of the trees.

The vertical distribution of the many species of bromeliads in the
forest trees of Trinidad reveals their
varying degrees of tolerance
(Fig. 6.27). One group of species grows only near the tree tops in
situations fully exposed to the sun, a second
group is found at inter-
mediate levels in partial sunlight, and a shade-tolerant third group
inhabits the lower branches of the forest where the illumination is
quite inadequate for the other groups.

Aquatic Plants. Reduction of light presents even more serious
problems in the aquatic environment. As we have seen illumination
diminishes rapidly with depth even in clear water and becomes
changed in spectral composition and in other respects. Plants at-

tached to the bottom in the marine environment consist principally of
algae with a few species of vascular plants such as the eel grass. In
fresh water vascular plants as well as algae are well represented in
the submerged vegetation.
Early investigators reported an apparent color zonation in the depth

of occurrence of attached plants. In Puget Sound, for example, the
green algae are generally the most abundant in shallow water, brown
algae dominate the zone from 5 to 20 m, and red algae are usually
most numerous in These color types of algae
depths of 10 to 30 m.
often occur at depthswhere the complementary color of the penetrat-
ing daylight predominates red algae, for example, tend to be abun-
dant in deep water where the blue or green component of daylight is
the strongest. It was formerly believed that the predominating color
of the light controlled the depth distribution of the color types of
algae because the plants would absorb light of complementary color

more This generality was referred to as chromatic adap-
efficiently.
tation. Many exceptions occur, however, and we now know that the
wavelength of light does not control in any precise way the depth of
distribution of algae on the basis of their color ( Dutton and Juday,
1944). Furthermore, the pigment of algae living in weak light deep
in thewater is sufficiently thick to absorb all the light incident upon it,
regardless of the part of the spectrum in which it occurs. Evidence
exists that some
of the light energy adsorbed by pigments other than
chlorophyll can be transferred to the chlorophyll present and thus

enhance the ability of certain algae to live in weak blue-green light
(Smith, 1951, Ch. 13).
236 Light
Group I
(exposure)
Group II (sun)
Group III (shade tolerant)

FIG. 6.27. Vertical distribution of various species of bromeliads in the forest trees
of Trinidad. ( Pittendrigh, 1948.)
Aquatic Plants 237
When algae are grown in the laboratory in light of different colors,
often tend to change color, and this
they response has also been
referred to as chromatic
adaptation. Experiments have shown that
individual plants photosynthesize most and best in
efficiently grow
that color and in that
intensity in which they have been living. In
the course of time fixed algae can
adapt themselves to habitats with
illumination of widely different conditions of color and In
intensity.
view of the great range in the transparency of natural waters, it is not
surprising that the maximum depth at which attached plants can live
varies widely in different situations (Table 15). In general benthic
TABLE 15
MAXIMUM DEPTH IN METERS FOR GROWTH OF ATTACHED PLANTS
Crater Lake, Ore. 120

Crystal Lake, Wis. 20
Trout Lake, Wis. 12
Mediterranean Sea 160

Challenger Bank 100
Off Iceland 50
Puget Sound 30
Baltic Sea 20
Off Cape Cod 10
TABLE 16
COMPENSATION DEPTHS IN METERS FOR PHYTOPLANKTON

(for midday periods)
Sargasso Sea 100

English Channel 35
Gulf of Maine 30
Off British Columbia 19
Trout Lake, Wis. 16
Woods Hole harbor 7
plants will not grow at depths at which the light intensity is less than
0.3 per cent of the surface value.
A glance at a chart of the oceans or of a typical lake reveals
that
only in a very narrow fringe around the margin is the water suffi-
ciently shallow for enough light to reach the bottom for the growth of
plants. Weare forced to the realization that in any deep body of
water far more organic matter is synthesized by the phytoplankton
present everywhere in the surface layers than by the benthic plants
238 Light
limited to the littoral zone. Photosynthesis, the first step in the
ecological cycle of the sea upon which all marine life ultimately
depends, is carried forward
primarily by the minute diatoms and
other microscopic planktonic plants. The growth of the phyto-
plankton is limited not only by the rapid diminution of light with

depth but also by the fact that vertical movements of the water sub-
changing light conditions.
ject the plants to continually
The greatest depth at which phytoplankton can grow successfully
in any body of water may be ascertained by suspending bottles
containing samples of the plant population at various levels beneath

the surface. Photosynthesis going on within the bottle adds oxygen
to the water, and respiration taking place simultaneously removes
oxygen from the water/ The rate of respiration alone may be meas-
ured by placing at each level a blackened bottle from which all light
is excluded. The sum of the gain in oxygen in the clear bottles and
the loss in the dark bottles gives a measure of total photosynthesis.
The increment oxygen in the clear bottles is the amount produced
of
by photosynthesis minus the amount consumed in respiration and

represents the actual increase in energy content or growth of the plants.
With the diminution of light in any environment photosynthesis
is reduced but
respiration remains approximately the same, pro-
vided, of course, that temperature and other factors are essentially
unchanged. For each species a point is reached in the reduction
of the illumination at which the rate of photosynthesis is just equal
to the rate of respiration; this is known as the compensation point or
better as the compensation intensittj. At light intensities below this
value photosynthesis may still go on but the plant is fighting a losing
battle: it cannot survive indefinitely under these conditions because
the loss of energy duetto the catabolic processes represented by
respiration exceed the gain in energy brought about by the anabolic

process of photosynthesis. With further light reduction a minimum
intensity is reached below which no photosynthesis at all can take
place.
In the aquatic environment the level in the water at which the
compensation intensity is found is called the compensation depth.
Sample values for phytoplankton based on measurements in the
middle of the day are presented in Table 16. For phytoplankton in
general the compensation intensity has been found to be about 1 per
cent of the value of full sunlight at the surface. In Fig. 6.28 the
change in the amount of photosynthesis at various depths during the

course of the day is plotted for a type of diatom representative of the
marine plankton. It will be observed that during the noon hours the
Aquatic Plants 239
illumination close to the surface was excessive and depressed photo-
synthesis. The highest rate of oxygen production occurred at 5 m
in the middle of the day. No significant amount of photosynthesis
occurred at any depth before 0600 in the or after 2200 and
morning
none below 35 at 1400. m
o
10 noon 14
Time >
FIG. 6.28. Photosynthesis of a species of phytoplankton (Coscinodiscus) at the
indicated depths during the course of the day off Stoke Point, England. (Jenkin,
1937, Copyright, Council of Marine Biological Assoc. of United Kingdom.)
If a plant is
photosynthesis during the day must build
to grow, its
up enough organic matter to more than make up for the material

lost by respiration not only during the day but also during the night.
In other words, the crucial value for the continued existence of the
plant is the compensation depth for the 24-hour period. This will
obviously occur at a shallower depth than the values reported for
experiments limited to the middle of the day. The compensation
depth for the complete day ranges between 20 and 30 during the m
summer in clear coastal water of the temperate oceans. In the winter
and transparent water the compensation depth occurs cor-
in less
respondingly nearer the surface. No constructive growth is possible

for diatoms or other pelagic plants below these levels. When we
recall that the average of the ocean is more than 4000 m, it is
depth
240 Light
apparent that the zone in which productive growth of the crucially

important phytoplankton can take place is a relative skimming of
the surface. The photosynthesis in the upper few meters of the
ocean and of lakes accounts for the main portion of the initial pro-
duction of organic matter for the whole breadth, length, and depth
of the water.
Surface
100m- Limit of growth for planktontc plants
160 - Greatest depth for benthic plants
250 - Daylight 0.001% of surface value
500 - Approximate limit for vision of fish

-
550 Complete darkness for man
800 - Limit of response to day and night

by Crustacea
- Photographic plate blackened

1,000 after80 minutes' exposure
No perceptible light
from the surface
1,700 -P Plate not affected after 120 mfnutes'

exposure
4,000 - Average depth of ocean
10,860
- Deepest recorded sounding in ocean
Summary of limitations by light factor in aquatic environment, Values

are for clearest water.
A summary of the conditions of light in the aquatic environment

and its critical limiting effects is given in Fig. 6.29. Here are indi-
cated the maximum depths for the growth of phytoplankton and
slightly deeper,
of benthic plants. Since the mean illumination
necessary for the vision of aquatic animals is so very much smaller,

Aquatic Plants 241
is at a much
the depth limitation greater level. Animals can respond
to the difference between day and night at somewhat greater depths.
Below this level no perceptible light from the surface penetrates and
the water is
completely dark except for light provided by the lumi-
nescent organs of deep-sea animals. Terms used in deep bodies of
water for zones based on the light factor are as follows:
Euphotic Zone: Sufficient light for photosynthesis.
Disphotic Zone; Insufficient light for photosynthesis but sufficient light for
animal responses.
Aphotic Zone: No light of biological significance from the surface.
The depth limits of these zones differ widely according to trans-
parency.
The discussion in this chapter has revealed the many important
roles played by light in the world of life. Although there is gen-
erally sufficient light on land for plants and animals, the special cir-
cumstances of this factor exert a wide control over the activities of
many terrestrial organisms. In the aquatic environment light is even
more crucial since, in addition to its various periodicities, its rapid
changes with depth impose serious limitations on the lives of both

animals and plants.
7
Oxygen and
Carbon Dioxide
With this chapter we turn from the more mechanical or physical

features of the environment to a consideration of some of the chemical
factors. Two
fundamental substances taking part in the chemical
exchange between the organism and the environment are oxygen and
carbon dioxide. These substances enter into the basic processes of
photosynthesis and respiration, as may be indicated by the overall
reactions in which C H 12 O
6 6 is taken to represent the carbohydrate of
the organism.
Light Photosynthesis
C0 + H O ;=i C H
2 2 6 12 O, + 2
Respiration
Oxygen and carbon dioxide thus stand in a

reciprocal relation to each
other as regards these fundamental reactions of life, and their
abundance in the environment is of direct critical concern to the
organisms of every habitat. In addition, these materials take part in

subsidiary reactions involved in the ecological relations of plants and
animals. Of particular ecological importance in this connection are
the decomposition and transformation of organic matter, carried out
primarily by microorganisms, since in these processes oxygen is con-

sumed and carbon dioxide is released.
OXYGEN
Oxygen is needed by almost all
organisms to make available the
energy contained in organic food materials. The great majority of
plants and animals use free oxygen from the air or from the water for
the oxidation of organic substances; these are aerobic organisms.
242
Anaerobic forms, on the other hand, get their energy by partial de-
composition of organic matter without free oxygen. Anaerobic or-
ganisms nevertheless depend upon the aerobic forms to produce the
organic matter upon which they live. As we shall see, the abundance
of oxygen environment may become critically low for aerobic
in the
organisms but it never becomes harmfully high for these forms.

Natural concentrations of oxygen may, however, be seriously detri-
mental to some anaerobes.
Oxygen is present in the air as one of the gases that are physically
mixed together, and it occurs in water in simple solution. Dissolved
oxygen does not combine chemically with water itself but it does react
with iron and other inorganic materials in the water. Points of con-
trast in this respect with carbon dioxide will be discussed later in the
chapter. Since oxygen is taken up and given off by life processes,

its concentration in the environment can sometimes be
appreciably, or
even seriously, altered by the activities of the plant and animal in-
habitants. Under these circumstances oxygen is a modifiable factor
of the environment.
Availability of Oxygen
Terrestrial Environment.
Oxygen constitutes 21 per cent of the
atmosphere, and this value varies by less than 1 per cent the world
over. Although plants and animals are continuously drawing upon
the oxygen supply in the air, and plants are periodically adding to
it,the concentration in the atmosphere is not changed appreciably
by these life activities because of the great volume and mobility of

the air medium. Most of the terrestrial environment is thus pro-
vided with a uniform and adequate supply of oxygen.
In two types of situations of importance in the terrestrial environ-
ment a lack of oxygen exists at high altitudes and in the subsurface
layers of the soil. At great elevations the concentration of oxygen is
low because all
gases have become rarer. The amount of reduction
in oxygen with altitude is
proportional to the reduction in total at-
mospheric pressure, discussed in Chapter 2. Thus at an altitude of
5500 m
O 2 is only half as abundant as it is at sea level. The oxygen
in soils drops from near the atmospheric value of 21 per cent at the
surface to about 10 per cent within well-drained loams and to lower
values in poorly aerated soils and in layers below the water table.
Lack of air circulation results from very fine texture or from the
flooding of the pore spaces.
Under these circumstances the respira-
tion of roots and of soil organisms, particularly those involved in the
244 Oxygen and Carbon Dioxide
decomposition of organic matter, reduces the oxygen supply faster
than it can be replenished from the atmosphere.
Aquatic Environment. The total amount of oxygen that water will

hold at saturation varies with temperature, salinity, and pressure.
Sample oxygen saturation values for fresh water and sea water at
temperatures within the range of ecological interest are given in
Table 17. When we realize that the 21 per cent of oxygen present
TABLE 17
CONCENTRATIONS OF OXYGEN AT SATURATION

Fresh Water Sea Water
in the atmosphere is equivalent to 210 cc per liter, the contrast be-

tween the amounts available in air and in water is brought into relief.
There may be 25 times as much oxygen in a liter of air as in a liter
of water. When the oxygen in the water is in equilibrium with
that in the air, however, the pressure of the oxygen is the same in
both media.
From
the foregoing it is clear that the world's main Reservoir of
free oxygen is in the atmosphere. The chief source eft oxygen for
the water environment is the oxygen which can be absorbed from
the air, but a second source of supply in aquatic habitats is the oxygen
released beneath the surface by the photosynthesis of submerged
and planktonic plants. Water may lose oxygen by diffusion from its
surface out into the atmosphere. Oxygen is also used up by the
respiration of aquatic organisms and by the decomposition of organic

matter in the water. Living organisms are thus seen to influence
both the supply and the depletion of oxygen in natural bodies of
water; oxygen is a modifiable factor in the aquatic environment but it
is
essentially an unmodifiable factor in the air. The actual amount
of oxygen present at any one time and place in the water is the result
of a balance between the processes of supply and depletion. Here is
another situation in which an ecological factor is controlled by a
dynamic equilibrium involving both physical and biological processes.
(Three general situations may be discerned in regard to the supply
of oxygen in the water environment: the surface stratum, the sub-
surface zone, and the deep layers. The water at the surface tends
to be in equilibrium with the air above it so that the oxygen value
Aquatic Environment 245
at or near the saturation concentration for the
existing conditions
is
of temperature and salinity. The thickness of the surface stratum

in which saturation equilibrium is found varies
greatly according to
the amount of turbulent mixing of the water in contact with the air.
In the subsurface zone great variation in oxygen concentration
may
occur because the water does not have easy exchange with the at-
mosphere and factors causing both decrease and increase in oxygen
are present. Respiration and decomposition, tending to deplete
the oxygen supply, occur at all levels, and photosynthesis, tending to
increase the abundance of oxygen, takes place down to the com-
pensation depth, that is, to the lower limit of the euphotic zone, as
described in the previous chapter. /When large populations are
dead material are decomposing, the
respiring or large quantities of
oxygen concentration reduced to a low level. ^ In stagnant ponds
is
and swamps choked with organic matter, and in rivers or other

bodies of water receiving excessive amounts of sewage or other
pollutants, the available free oxygen often becomes completely ex-

hausted/ Under other circumstances supersaturation may occur in
subsurface water layers. If water which was saturated during the
winter becomes warmed as the season advances, more oxygen will be
present than can be held in true solution at the higher temperatures.
Similarly, when photosynthesis exceeds respiration
at intermediate
levels in the euphotic zone, oxygen may be released into the water
faster than it can be carried away. As a result of these physical and
biological influences, supersaturated values as high as 180 per cent
in the sea and 300 per cent in inland lakes have been reported.
Often cyclic fluctuations in oxygen content are observed in natural
bodies of water. Seasonal changes in temperature and circulation
with differences in rates of photosynthesis, respiration, and
coupled
decomposition result in changes both in the absolute amount of
oxygen present in the water and in the degree of saturation. Under
certain circumstances a pronounced diurnal fluctuation in oxygen
concentration is found; this is known as an oxygen pulse. During the
daylight hours photosynthesis tends to cause an increase in the

amount of oxygen present, but, after the sun has set, the respiration
of the aquatic organisms and the decomposition processes going on
draw on the free oxygen in the water. The amplitude of the oxygen
pulse may reach considerable proportions in quiet waters in which
green plants are growing in abundance. Measurements in an Ohio
lake, forexample, revealed a diurnal fluctuation in oxygen concentra-
tion extending from 6.7 ppm at 8:00 A.M. to 13.0 ppm at 5:00 P.M.
(Fig. 7.1).
How does oxygen get to the bottom of the ocean? have seen We
that the sources of oxygen are at or near the surface, but we know
that many kinds of aerobic animals live at the bottom of some lakes
and ocean abyss at depths of several miles beneath the surface.
in the
Although oxygen moves slowly through the water by direct diffusion,

it can reach the
deeper levels at an adequate rate only by the circula-
tion of the water itself.
6 pm 12 mid. Sam 12 noon 5pm

ppm
15
fo 10
"?
Free carbon dioxide
>o--o
x
(f
f
3-5
-10
Modified from Tressler, Tiffany, and Spencer, 1940
FIG. 7.1. Diurnal cycle of temperature, oxygen, hydrogen ion (as pH), and car-
bon dioxide in the surface waters of Buckeye Lake, Ohio, during August 12-13,
1930. Negative values for CO 2 represent amount of CO 2 required to make water
neutral to phenolphthalein.
Oxygen-rich surface water is carried downward by wind stirring,

eddy conduction, and mass sinking. Wind stirring is very effective
in aerating the upper layers of lakes and of the ocean at times when
the water circulates freely. Whenever a water body is covered by
ice, no turbulence caused by wind or waves is possible. At other
seasons stirring is greatly curtailed whenever the water is stratified in
respect to densityas
a result of salinity differences occurring at any
time of year, or as a result of a thermal gradient arising primarily
in summer. During periods of pronounced stratification the circula-
limited to the layers above the
tion produced by wind and waves is
density discontinuity (Fig. 7.2). When stratification has been de-

stroyed, wind stirring may extend sufficiently deep to reach the bottom
of shallow lakes and coastal waters, and thus to
replenish the oxygen
supply. There is a limit, however, to the depth to which turbulence
caused by wind is effective. In the open ocean, where the highest
winds and largest waves occur, little influence of wave action is felt
below 100 m.
Direction of wind
r -*" ~* -" Plane -*

,
Shearing
y ~^-_ "^vr "^currents*-*
^
f__ ^^~ "*^n Return ^-,
Thermocline
Fir,. 7.2. Wind-driven currents with a pronounced thermocline. Circu-

in a lake
lation is limited to the epilimnion. (By permission from Limnology by Welch,
Copyright 1952, McGraw-Hill Book Co.)
Oxygen may be carried to depths beyond the limit of effective wave

action by eddy conduction. Wherever differential movement of cur-
rents occurs, eddies are generated along the current margins, and
these cause units of water to be exchanged (Fig. 7.3). The exchange
will bring about a transfer of any characteristics of the water that
differ in the adjacent water masses. The rate of transfer depends upon
SURFACE WATER
Eddy transfer
O-O-O-O-O-O
CURRENT
>-O-O-O-0-O
Eddy transfer
DEEP WATER
FIG. 7.3. vertical section of a body of water, indicating vertical

Diagrammatic
transfer eddies across the current boundaries. Horizontal eddy conduction
by
lateral margins of the current.
similarly takes place at the
(a) the rate of mixing of water units and (b) the difference in con-
centration of each characteristic. The rate of mixing, or austausch
coefficient, is a property of the water under the existing conditions of

turbulence, and it affects all characteristics that are being exchanged
between water masses, At the same time that oxygen is being trans-
ferred, heat may also be moved downward, phosphate moved upward,
and other exchanges carried out by the same eddy action between
the water masses. This is an illustration of the principle of common
transport: a current that transfers one property of the medium also
transfers other properties at the same time (Redfield, 1941).
Eddy conduction is a process by which materials are transferred at

right angles to the direction of current. The transfer may take place
vertically, as
in the situation just described, horizontally, or in some
other direction. The downward
transfer of oxygen by ecldy conduc-
tion is slow but it
commonly a thousand times greater than the
is
diffusion rate. Although eddy conduction helps furnish oxygen to

water layers at middepths wherever currents of different speeds or
directions are flowing, it cannot supply an adequate amount of this
essential material to the deepest parts of lakes or of the ocean.
Fortunately for the abyssal animals oxygen can reach them by

another method, namely, the mass sinking or deep circulation of the
water. In the typical deep lake of the temperate region water
masses charged with oxygen at the surface sink to the bottom during
the spring and fall overturns, as described in relation to the tempera-
ture cycle. As a result of the extensive circulation of the lake at these
periods the water may become turbid and formerly this was often
noticeable in poorly filtered water supply systems. When muddy
water issued from the taps, the residents would nod knowingly and
say, "The pond working." The limnologist, however, refers to
is
the periods of overturn that bring a fresh supply of oxygen to the
deeper layers after winter and summer stagnation as occasions when

"the pond takes a breath."
An example of the seasonal changes in oxygen at various depths in
a temperate lake is taken from the classic work of Birge and Juday
(1914) (Fig. 7.4). During the period of this representative study
oxygen
was abundant at all depths after the spring overturn but a
rapid depletion began in May following thermal
stratification^ Oxy-
gen was almost completely absent at depths greater than 15 during m

July, August, and September. The fall overturn in mid-October
caused a sudden mixing and equalization of oxygen at all levels; in
succeeding weeks increasing amounts of oxygen were supplied

throughout the lake by deep circulation. In
certain very deep lakes
the seasonal circulation does not reach the bottom, with the result
that the deepest layers may be completely devoid of oxygen through-
out the year.
In the temperate regions of the ocean the thermal stability built up
during the summer is similarly broken down in the autumn. Turbu-

lence produced by wind and waves then carries oxygen-rich water
downward, and this deep stirring continues all winter. The down-
ward extent of the winter mixing is sufficient to reach the bottom in
coastal areas but its effect does not go below the permanent thermo-
cline of the open sea (Fig. 5.4). Seasonal changes in temperature
and wind action thus do not provide for the delivery of oxygenated
water to the great depths of the ocean.
May June July Aug. Sept. Oct. Nov.
FIG. 7.4. Changes in oxygen during the warm part of the year at the indicated
depths in Lake Mendota, Wisconsin. ( Needham and Lloyd, 1937, after Birge and
Juday. )
Oxygen supplied to the ocean abyss by deep permanent currents

is
that originate near the surface in high latitudes, sink to intermediate

levels or to the bottom, and flow for thousands of miles toward and
beyond the equator (Fig. 7.5). This deep flow of water is dynam-
ically integrated with the horizontal
current systems with the result
that the vertical movements of water follow a complicated pattern
(Sverdrup et al, 1942). As a simplified generality, however, the
cold water that sinks to great
depths may be understood eventually
to find its the
way to surface again in the central part of the ocean,
and a return flow of water from the equatorial regions toward the
poles brought about as a part of the horizontal circulation of the
is
upper water masses.
5000
70 S 60 50 40 30 20 10 S N 10 20 30 40 50 60N
FIG. 7.5. Cross section of the Atlantic Ocean, showing the distribution of oxygen
in cubic centimeters
per liter. The north-south and vertical components of the
oceanic currents are indicated by the arrows. (Modified from Svcrclrup et al.,
1942, after Wiist.)
Sea water near the surface in tropical regions contains about 4 cc

of oxygen per liter, becomes colder at higher latitudes,
but as it
oxygen is absorbed from the atmosphere and produced by photo-

synthesis until a concentration of about 8 cc per liter is attained.
When the water masses leave the surface and sink toward the bottom,
depletion of oxygen begins as a result of respiration and decomposi-

tion, but as much as 5 cc O 2 per liter remains when the water reaches
the ocean abyss. Animals living at the bottom of the sea thus
receive oxygen via a direct flow of water from the enriched polar
seas, but this is perhaps the slowest delivery service in the world.
Calculations based on the rate of consumption of oxygen and also on
the relative proportion of CM in deep water indicate that movement
of Arctic surface water to deep layers at midlatitudes may require a
thousand years or more.
The concentration of oxygen at any point in deep water depends
upon (1) the amount that the water contained when it left the
which oxygen is used up en route by respira-
surface, (2) the rate at
tion and decomposition, and (3) the time that has elapsed. Mini-
mum values occur at positions in the ocean at which the combined
effect of the foregoing factors has reduced the oxygen to the greatest
extent. The lowest oxygen values are not found at the bottom but
at certain intermediate regions, as indicated in Fig. 7.5. In no part
Oxidation-Reduction Potential 251
of the open Atlantic Ocean
is the
oxygen completely used up, and
minimum values are generally above 3 cc
per liter. In the Pacific
Ocean, however, oxygen is reduced to zero in certain oxygen minimum
layers, as for example, at depths between 300 and 1300 m off the
coast of lower California.
Other instances in the marine environment of the complete lack
of oxygen, or of very low concentrations, are found
only in special
situations in which circulation is or cut off and de-
largely entirely
composing organic matter has accumulated. The classic example
of this condition is in the freewater of the Black Sea, in which no
measurable oxygen is m
found from a depth of about 150 to the
bottom at 2200 m. Vertical stirring is prevented by a layer of fresh
water at the surface supplied by the discharge of the Danube and
other large rivers entering the Sea, and deep exchange with outside
water is stopped by the shallow entrance at the Bosporus. Organic
particles carried in with the river water use up the oxygen as they
sink to deeper layers, and anaerobic decomposition produces great
quantities of hydrogen sulphide. Similiar conditions on a smaller

scale are found within deep fjords with shallow sills and within other
estuaries in which the circulation does not supply the oxygen as fast
as it is used up (Fig. 7.6). The bottom material of the marine en-
vironment also likely to develop a shortage of oxygen unless it is
is
sufficiently coarse to allow good

movement of water through it.
Fine muds containing decomposing matter contain little or no
oxygen at depths of more than a few centimeters from their surfaces.
Oxidation-Reduction Potential. Another aspect of oxygen avail-
ability in the aquatic environment, but less
well known, is the oxida-
tion-reduction potential, or redox potential. The redox potential
(Eh), measured in an expression of

volts, is the
electropositivity or
negativity of a substance in solution as referred to a hydrogen stand-
ard. A
positive redox potential
indicates a condition tending to cause
oxidation, and a indicates the reverse condition,
negative potential
tending to bring about reduction. As used to describe an aquatic
habitat, redox is the overall summation of the redoxes of all the solutes
present in the water. Redox potentials have two general properties

that must be considered: intensity or Eh as defined above; and ca-
pacity or poising, which is analogous
to the buffering of a pH
system.
A high capacity reflects the tendency of the system to retain its in-
tensity despite minor changes
in its constituents.
In well-aerated natural waters oxygen concentration usually gov-
erns the redox potential and produces positive values. However, fer-
rous complexes or oxygen shortage may allow various inorganic and
organic ions and molecules to set more negative values. Such low
redox systems represent reducing agents that would tend to combine
with any introduced oxygen. An aerobic organism, needing oxygen
for its
respiration, would find such competition exceedingly difficult,
ifnot insuperable, if the redox system were well poised. Facultative
and obligative anaerobes, however, have no such difficulty and are
able to tolerate low redox values.
FIG. 7.6. (A) Basin with local formation of heavier water and deep outflow
across the sill, as in the Mediterranean Sea, the Red Sea,and the Inner Gulf of
California. ( B ) Basin with surface outflow of lighter water and occasional inflow
of denser water across the sill, as in the Black Sea, Baltic Sea, and many fjords.
(From The Oceans by Svedrup et al, 1942, Copyright Prentice-Hall, N. Y.)
Zonation of organisms according to redox potentials has been shown,

especially in sediments, but whether the oxygen concentration or the
redox potential is the determining factor is hard to ascertain since the
latter generally depends chiefly upon the former. In marine muds
the redox potential decreases with depth in the sediment with the
most rapid change occurring in the few uppermost centimeters. The
abundance of aerobic organisms correspondingly drops off sharply,
and at increasing depths the anaerobes become relatively more
abundant. The type

of bottom fauna in certain lakes has also been
found to be related to the redox potential. In lakes in which the
water above the deepest mud had an Eh above 0,4 volt, the mud
itself supported a pure or mixed fauna characterized
by midge larvae
of the genus Tanytarsus, whereas in lakes with an Eh below 0.3 volt
a true Chironomus fauna occurred (Hutchinson et al., 1939). For a
further discussion of the ecological effects of redox potentials the
reader is referred to ZoBell (1946); it is obvious that the influence of
this aspect of the oxygen factor is due for closer
scrutiny in the future.
Effects of Oxygen Availability
Terrestrial Environment.We may now examine the ecological

effects of the
varying amounts of oxygen present in different habitats.
Since oxygen is
amply abundant in the lower
portions of the earth's
atmosphere, this factor has no important limiting action in most ter-
restrial situations.The reduction in the partial pressure of oxygen
at high elevations imposes a restriction on the altitudinal distribution
of organisms having a high oxygen requirement. No mammals can
livepermanently at altitudes at which the partial pressure of oxygen
is than about 45 per cent of its value at sea level. Oxygen short-
less
age as well as the thinness of air imposes a similar ultimate limit on

the altitude at which birds can live. The altitudinal range for most
types of lower animals and for plants is limited by low temperatures

or by other ecological factors long before the diminished oxygen con-
centration becomes a serious handicap.
Organisms excluded from soils and

requiring free oxygen are
ground debris within which the circulation of air is inadequate.
The lack of oxygen in poorly aerated soils becomes seriously harmful
to the roots of most plants at concentrations below about 10 per cent.
Root penetration for most of the vegetation is stopped by soil layers
containing less than 3 per cent oxygen and also by the water table
since even less oxygen is available in the ground water. If areas with
established vegetation are subjected to prolonged flooding, caused for
example the construction of beaver dams, most of the plants will

by
be "drowned" or killed by the cutting off of root aeration.
The roots of some plants, however, possess special adaptations en-

abling them to grow in soil layers devoid of oxygen. In some types,
including particularly the herbaceous hydrophytes, air passages within

the stem and major roots allow oxygen to be brought from the
atmosphere, or possibly from internal sources in the upper part of the

plant, and delivered to the lower extensions of the root system. In
the black mangrove, common along the
margins of Florida lagoons,
root branches, known as pneumatophores, grow vertically
upward
through the mud and water until they project into the air, where they
have ready access to oxygen. Other special features of plant growth
found in permanent swamps are the "knees" and enlarged trunk bases
displayed by such trees as the bald cypress (Fig. 7.7). The knees
Photo U. S. Forest Service
FIG. 7.7. Enlarged trunks and "knees" of cypress trees growing in permanent
swamp conditions in the Francis Marion National Forest, South Carolina.
were formerly believed to provide aeration for the roots, but evi-
dence now indicates that these root excrescences and the enlarged
trunks represent excessive cambial growth resulting from the com-
bination of abundant water and oxygen supply for the tissue at the
water line. The roots of a few vascular plants without observable
adaptation for securing outside oxygen, such as the willow, are able
to remain active during prolonged flooding; the root tissues in these
exceptional instances are presumed to be capable of anaerobic

existence.
Most cryptogamic plants and soil animals depend upon aerobic
respiration and hence are excluded from soil layers devoid of oxygen.
A few of the higher forms, such as the earthworm, may be able to
do without oxygen for short periods, as when the soil is rain soaked,
but for the most part earthworms
migrate to the surface when oxygen
becomes depleted. Many organisms encyst, or go into a dormant
condition,when oxygen becomes too scarce for the species concerned.
However, some of the microorganisms inhabiting the soil can follow
an anaerobic existence (Waksman, 1932). Included are members of
the genera Clostridium and Rhizobhnn that are
important as nitrogen-
fixing bacteria; Clostridium tetani is of particular concern to man as
the microbe that causes lockjaw. We realize then that the highly
variable conditions of oxygen availability often exert a critical control
over the distribution and activity of both aerobic and anaerobic
organisms involved in the ecology of the soil.
Aquatic Environment. We have seen that the supply of oxygen
in the aquatic environment varies over the complete range from super-
saturation to total exhaustion, but that, even when saturated, water
contains a much lower concentration of oxygen than the atmosphere.
Some fresh-water organisms have hit upon certain dodges by which
they can live in the aquatic medium and still breathe air. Many
water bugs, such as Notonecta, and certain beetles come to the surface,
entrap bubbles of airbeneath their wing cases, and swim down with a
a period of time. Other animals,
supply of oxygen that will last for
like the water scorpion Ranatra ( Fig. 6.12), the mosquito larva, and the
rat-tailed maggot, possess breathing tubes that extend to the surface.
Some kinds of snails periodically refill their lungs with air at the
surface, whereas other water animals are able to bore into the stems
or roots of emergent hydrophytes and obtain oxygen from the internal
air spaces of the plants.
Most aquatic organisms, however, must get along with the rela-
tively meager supply of oxygen
dissolved in the water. Let us con-
sider the ecological conditions in well-aerated water before taking up
the effects of severe oxygen depletion. Although saturated water
contains only a small fraction of the amount of oxygen in an equal
volume of the atmosphere, the partial pressure of oxygen in the water
is to that in the air with which it is in equilibrium. Accord-
equal
the of the in water to cross the respiratory
ingly, tendency oxygen
membranes of organisms is just as great as it is in air. The lower
concentration of oxygen in water simply means that less is in reserve.
The relatively small amount of oxygen in the water is adequate for
aquatic organisms if the medium in contact with the absorbing mem-
branes is
rapidly replenished.
Inhabitants of the water environment display a wide variety of
a sufficient supply of oxygen from the
adaptations that serve to obtain
medium. Sessile animals cause water to flow through their respira-
tory chambers or wave their respiratory organs about in the surround-
ing water. The echiuroid worms keep a current of water flowing
through their burrows in the sand when the tide is in by rhythmic
contraction of their bodies, but reduced metabolism allows them to
withstand periods of 18 hours when their tide flats are out of water.
More active animals, such as most fish and higher crustaceans, possess
special mechanisms for pumping water continuously over their gills.
Certain however, reverse the process and produce a flow over
fish,
the gills by swimming rapidly through the water with their mouths
open. Some species, as, for example, the mackerel, have come to
rely entirely on this latter method, and in the course of evolution the
gills and opercular muscles have become reduced, with the result
that these fish cannot obtain sufficient oxygen while remaining sta-
If a mackerel is confined in a small
tionary. space, or prevented from
swimming rapidly, it will die of asphixiation even though the sur-
rounding water is saturated with
oxygen ( Hall, 1930 ) .
In view of the foregoing one might expect that in those aquatic

habitats in which the water is not saturated with oxygen, the lack of
this vitalsubstance would immediately have serious consequences.
However, many aerobic aquatic organisms are quite able to survive
with concentrations of oxygen much below normal pressures. The
metabolism of some species is so low that a small
oxygen supply is
sufficient. Certain species possess special respiratory pigments or
other physiological adaptations that aid in the absorption of oxygen
at very low partial pressures. Chironomid larvae, inhabiting the
muddy bottoms of ponds, are furnished with a type of hemoglobin
that becomes 95 per cent saturated under a partial pressure of oxygen
of only 10 mm
Hg, whereas mammalian hemoglobin is less than 1 per
cent saturated at this pressure. These insects are also capable of a
remarkably simple anaerobic metabolism in which the lactic acid

formed from glycogen breakdown is excreted, thus eliminating this
toxic compound rather than oxidizing it as most animals do.
The relation of oxygen consumption by animals to the tension of
oxygen in the environment follows two general patterns: (1) the non-
regulatory type in which consumption is highly dependent upon ten-
sion, as seen in certain annelids and arthropods (e.g., Nereis, Hornarus,
Limulus, and Callinectes), and (2) the regulatory type in which con-
sumption is
independent of oxygen pressure over a wide range, as
seen in certain crustaceans and mollusks (e.g., Astacus, Carcinus,
Aplysia, and Eledone). In the latter type the harmful effects of in-
sufficient oxygen supply appear rather suddenly when the oxygen
drops below the critical tension for the species concerned. The
criticaloxygen tensions for a large number of animals, a further dis-
cussion of adaptations to low oxygen
supply, and a consideration of
physiological relations to the oxygen factor in general are to be found
inProsser (1950, Ch. 8).
It is difficult to determine the minimum concentrations of
oxygen
required by various aquatic animals. The value is influenced by
temperature, pH and other modifying factors, as well as by the
y
degree and rate of acclimatization to low oxygen tensions. In the

hypolimnion of lakes, where oxygen is likely to be depleted, the tem-
perature is low, with consequent reduction in the animal's metabolism

and in its need for oxygen. Organisms require more oxygen when
in an active condition than when quiescent; some species enter a
dormant condition in which they can withstand varying degrees of
oxygen deprivation for varying lengths of time. The ecologist thus
recognizes that the animals caught in a water layer containing little
or no oxygen are not necessarily able to live permanently under the
observed conditions: they may be in a dormant condition for a
limited period, or they may have made a temporary excursion into the
poorly aerated zone. Nevertheless we have obtained a knowledge of

the approximate oxygen requirements of many common aquatic
animals, and even among completely aerobic forms great differences
in the minimum values tolerated have been revealed. At one extreme
are those like the toadfish that seems to be able to survive if any de-
tectible amount of oxygen exists in the water. Carp and other fish
adapted to live in muddy ponds often show no harmful effects until
oxygen has been reduced to 0.3 cc per liter or even to 0.1 cc per liter.
At the other extreme are active fish like the mackerel that requires
3.6 cc per liter (70 mm
Hg) at 24C (Hall, 1930).
In those ponds, lakes, and fjords, in which the oxygen concentration
of the lower levels is reduced to zero during the summer ( and in some
instances also during the winter) the inhabitants must either migrate
out of the oxygenless zones or enter upon some sort of anaerobic
existence,if
they are to survive. Copepods and other types of zoo-
plankton are known to leave the hypolimnion of lakes as summer stag-
nation begins. Fish and the motile bottom fauna undertake similar
seasonal vertical
migrations. Many animals and plants with no
means of escape die off in large numbers after the exhaustion of the
oxygen supply. However, certain species of worms, crustaceans,
insect larvae, mollusks, and other invertebrates are known to remain
in thebottoms of lakes and to survive for periods of many weeks each
year with no detectible oxygen present in the bottom water. A
further discussion of the
types of metabolism displayed by inverte-
given by von Brand (1946).
brate animals capable of anaerobiosis is
When the supply of oxygen in a confined aquatic habitat becomes
reduced below the level of toleration, the lethal effects sometimes
appear quite suddenly. We the fish
occasionally see reports that all
in a pond have been found dead on one whereas

morning the day be-
fore they had appeared to be in a Such a
perfectly healthy condition.
situation occurs most
frequently after a hot calm spell in summer.
Because of the high temperature the amount of oxygen that the water
is low and the rates of
can hold respiration by the pond organisms and
of the decomposition of organic materials in the water are high. In
calm weather no effective stirring takes place to replenish the sub-
surface water levels. As a result of this combination of circumstances
the oxygen minimum reached by the diurnal pulse at the end of each
night becomes progressively lower. Early one morning the oxygen

concentration drops below the toleration point, and the whole popula-
tion of fish in the pond is suffocated.
In aquatic habitats permanently and completely devoid of oxygen
no aerobic organisms can live. Included in this category are: the
deep waters of certain lakes, the Black Sea, and certain arms of the
ocean with insufficient the subsurface layers of mud
circulation;
bottoms; and many heavily polluted waters. Such places can be per-
manently inhabited only by forms that are able to function indefinitely
on an anaerobic basis. The population of these habitats consists
chiefly of anaerobic Protozoa and bacteria. Many of these obligate
anaerobes are killed by the presence of oxygen. This fact is put to
practical use in the employment of thorough aeration as one step in
the purification of water supplies. Certain higher animals, such as
slime worms (Tubificidae), pea clams (Pisidium), and insect larvae
(Corethra and Chironomus), are found in the bottoms of very deep

lakes where oxygen is
very low or absent.
The organisms that inhabit the oxygenless water layers carry out a
partial decomposition of the organic matter filtering down from above,

and frequently release large quantities of hydrogen sulphide in the
process. In the Black Sea the permanent absence of oxygen in the
deeper water layer brings the vertical distribution of aerobic benthic

animals to an end at depths varying between 115 and 165 m accord- m
ing to the locality. The plankton is similarly limited to the upper 125
m in the central part of the sea but may extend to 175 m
in marginal
areaswhere the surface water is stirred more deeply. Lack of oxygen
and abundance of poisonous hydrogen sulphide exclude all life except
anaerobic microorganisms from the lower 2000 m of the Black Sea.
Carbon Dioxide 259
oxygen we have seen that this life-

In the foregoing review of
giving material is sufficiently abundant in most major habitats, but

that it becomes critically scarce in certain situations. Most
special
land organisms find enough oxygen in their habitats, and the great
majority of marine animals are so adapted that the amount of oxygen
in sea water ordinarily meets their requirements, although its con-
centration is far less than in the
atmosphere. As green plants photo-
synthesize and grow, they produce more oxygen than they consume,
and hence run short of oxygen only in rare instances. But we have
seen that in fresh water and under certain conditions on land and in
the sea, available oxygen may become seriously reduced, with the
result that whole populations
may be killed unless they have means
of escape, or are capable of anaerobic existence. The distribution of
aerobic plants and animals is sharply curtailed at the margins of
habitats permanently devoid of oxygen, and within these areas the
relatively few groups of completely anaerobic organisms hold un-

disputed domain.
CARBON DIOXIDE
A consideration of carbon dioxide as an ecological factor is of vital
concern first of all because it is one of the essential
ingredients of
the photosynthetic reaction. This material thus provides the carbon
source necessary for the growth of all green plants and indirectly for
allother organisms. Carbon dioxide also influences other features of
the environment. Quite in contrast with oxygen, carbon dioxide
combines chemically with the water medium itself, and forms car-
bonic acid. Through this reaction it influences the hydrogen ion
concentration (measured as pH), and it forms compounds with
calcium and other elements of ecological importance. Carbon di-
oxide affects the respiration of animals and, in combination with
calcium, takes part in the formation of their bones and shells.
When animals and plants respire, organic matter is oxidized and
carbon dioxide is produced. This process is the reverse of photo-
synthesis as indicated in the equations at the beginning of the chapter.
Carbon dioxide is also released when dead organic matter is decom-
posed by the oxidizing or fermenting activities of microorganisms.
On the other hand, carbon dioxide is withdrawn from the environment
by the growth of green plants and by lime secretion. Thus, carbon
dioxide, like oxygen, enters the living complex in an absolutely vital
way. Since the abundance of carbon dioxide in many situations is

strongly affected by living organisms, we may definitely list this ma-
terial as a modifiable property of the environment. Because of the
in the funda-
reciprocal relation between carbon dioxide and oxygen
mental reactions of nature mentioned above, we generally find that
wherever the supply of one of these materials has been depleted the
concentration of the other has been increased.
Carbon Dioxide in the Terrestrial Environment
It is a striking fact that carbon dioxide a material so essential in

the ecological relations of organisms is
present in the earth's atmos-
phere in an extremely small amount; constitutes only 0.03 per cent
it
of the air medium. Carbon %

dioxide is thus only about o as
abundant in the air as oxygen. Because of the great mobility of the
atmosphere, however, carbon dioxide is well distributed, and its
low
concentration is generally sufficient for at least a moderate amount
of photosynthesis the earth over. Possibly the reduction in carbon
dioxide at great altitudes would seriously curtail the photosynthetic
rate of plants, but low temperature or poor soil conditions are usually
far more important in limiting plant growth in the mountains,
0.200
Carbon dioxide 0.010 volume percent
400 600 800 1000

Light intensity, foot-candles
Fie. 7.8. Relation between rate of photosynthesis of wheat plants and light in-
tensity at normal atmospheric, at decreased and at increased concentrations

of car-
bon dioxide. (From Meyer and Anderson, 1939, after Hoover et al., Smithsonian
Misc. Coll, 1933.)
COt in the Terrestrial Environment 261
The normal amount of carbon dioxide available in the
atmosphere,
although allowing adequate plant growth under usual circumstances
at ordinary altitudes, is insufficient for
photosynthesis to attain rates
that would be possible under favorable natural conditions if the con-
centration of carbon dioxide were
higher. During conditions of low
illumination the light factor controls the rate of photo-
ordinarily
synthesis, but under high illumination the small amount of carbon
dioxide in the air acts as the limiting factor. This situation is illus-
trated in Fig. 7.8 in which rate of
photosynthesis of wheat plants in
relation to illumination is plotted under conditions of normal, in-
creased, and decreased tensions of carbon dioxide. In this experiment
under a normal carbon dioxide tension of 0.037 per cent the rate of
photosynthesis (as measured by utilization of carbon dioxide) in-
creases with increasing illumination up to 1000 ft-c. At higher il-
luminations, beyond the
limits of the graph, little further increase in
photosynthetic rate took place, as the availability of carbon dioxide

became limiting. With carbon dioxide at 0.010 per cent the limiting
action of carbon dioxide availability became complete at 400 ft-c,
whereas an increase of the carbon dioxide concentration to 0.111 per
cent resulted in an augmentation of photosynthesis at all illuminations,
with the curve still steeply rising at 1000 ft-c. Practical advantage of
this situation can sometimes be taken by artifically increasing the car-
bon dioxide in sealed
greenhouses. Improved growth of certain
plants is found for tensions of carbon dioxide of three to twenty times

the normal amount, but higher concentrations are harmful.
In nature some instances of local significant increases in atmospheric
carbon dioxide have been reported. In certain volcanic areas carbon
dioxide escapes from fissures in the rocks and, being heavier than
air, forms a layer near the ground. From these "death valleys," such
as the one in Java mentioned earlier, all animal life is excluded. The
carbon dioxide in soil air generally is increased as a result of decom-
position and respiration beneath the ground surface under conditions
of poor aeration. The air under dense vegetation may exhibit values
considerably above normal, and the air just above the soil of certain
cultivated fields has been found to show a tenfold increase in carbon
dioxide content, but no evidence is available as to any ecological ef-
fects that may have been produced. In an experiment conducted in
however, the carbon dioxide in the atmosphere was found
Georgia,
to be 25 per cent higher on certain foggy days, and during these pe-
riods the photosynthesis of plants under observation was increased
to as much as seven times the normal value (Wilson, 1948).
Our atmosphere receives carbon dioxide from geological and in-
dustrial sources. The supply is increased by carbon dioxide released
from the ocean and from respiration and decay. Carbon dioxide is
withdrawn from the atmosphere by the photosynthesis of the vegeta-
tion, The existing balance between supply and consumption provides
a very low concentration of carbon dioxide in the air, but, if the
minute amount of this essential gas were not present, all life on land
would come to a stop. The lives of terrestrial organisms thus depend
upon a very slender thread. In former geologic epochs the abun-
dance of atmospheric carbon dioxide may have been considerably
greater or smaller with the consequent possibility of significantly in-
creased or decreased photosynthesis. At present we realize that car-
bon dioxide is a general limiting factor in the sense that plant growth
might go on at a higher pace in the world as a whole if our atmosphere
contained more of this gas. However, since carbon dioxide is essen-
tially uniform over the earth's surface,
it does not cause
significant
differences in the distribution and growth of the terrestrial vegetation.
Carbon Dioxide in the Aquatic Environment

The amountof carbon dioxide present in the medium is crucially
important for aquatic organisms just as it is for terrestrial forms, but

the circumstances of the occurrence of carbon dioxide in the water are
much more complex. Carbon dioxide is readily soluble in water, but
since the concentration of carbon dioxide in the atmosphere is so low
(0.3 cc per liter), water in equilibrium with it will hold only about
0.5 cc per liter free CO
2 at 0C, or 0.2 cc per liter at 24C, in simple
solution. Actually, the total CO

2 in most natural waters is consider-
ably larger because additional amounts are present in the form of

carbonate and bicarbonate ions. Sea water at a salinity of 35%
normally contains a total of about 47 cc CO 2 per liter, or roughly 150
times the concentration in the Taking air. into account the relative
volumes of the ocean and of the atmosphere, it has been estimated
that more than 50 times as much CO 2 exists in the seas as in the air
( Rubey,
1951 ) The ocean is thus the great reservoir for the world's
.
supply of available CO 2 and this marine source tends to regulate the

amount in the air just the reverse of the situation with oxygen.
Reactions of Carbon Dioxide in Water. Dissolved CO 2 combines
with H 2 O to form carbonic acid, and this dissociates as follows:
C0 + H
2 2 ^ H C0 ^ H+ + HCOr ^ H+ + CO,-
2 3
The amount of CO in2 simple solution plus that in the form of 2 CO 3 H

is called the free CO 2 . The amount of the CO 2 in the bicarbonate
Reactions of Carbon Dioxide in Water 263
=
ions (HCO 3 ~) and in the carbonate ions (CO 3 )
is called the com-
bined CO
2 .
strong acid is added, the combined
If 2 will be con- CO
verted to the free form. The amount of acid
required to accomplish
this is a measure of the alkalinity, that is, of the amount of anions of
weak acids (chiefly 3 and HCO "
CO 3
=
)
in the water, and also of the
cations balanced against them. The alkalinity of sea water bears a
fairly constant relation to the chlorinity and hence to the salinity.
The relation of the equilibria of CO 2 to the alkalinity of sea water is
discussed in further detail by Rakestraw (1950); general accounts of
the reactions of 2 are CO
given by Harvey (1945) for sea water and
by Ruttner (1953) for fresh water.
When H CO 2 3
hydrogen ions (or more prop-
dissociates, it releases
erly, hydronium ions, H O).

affect the pH, which is the
3 These
negative logarithm of the hydronium ion activity. The pH is a meas-
ure of the degree to which the water is acid ( pH below 7 ) or alkaline
(pH above 7). If the equilibrium which is established in the water

is near
neutrality (pH
=
7), most of the CO 2 will be present as
HCO 3 ~ ion. At high pH values more CO 2 is present as CO 3 ion,
and at low pH values more is present in the free condition. Thus
pH and the relative distribution of CO 2 in its three components are
mutually interdependent, as indicated in Fig. 7.9. Addition or re-
100
7 8 9 10 11 12
pH
FIG. 7.9. Percentage of total carbon dioxide in each of its three forms in water as
a function of hydrogen ion concentration. (Emerson and Green, 1938.')
moval of CO 2 will affect pH; conversely, any other factor changing

the pH will affect the CO
equilibrium (Dye, 1952).
2
The bicarbonate and carbonate ions and other anions of weak acids
form a buffer system which tends to resist changes in pH. The buffer
of the water is determined by the abundance of these anions,
capacity
and it is therefore also directly related to the alkalinity. Sea water
and hard fresh water are relatively highly buffered natural media.
As a
result of buffering, the surface waters of the open ocean rarely
depart from a of 8.0 to 8.4, the deeper waters being generally

pH
confined to a pH
of 7.4 to 7.9. In the littoral zone, and particularly
in salt ponds and tide pools, the range is greater. The in marine
pH
muds may drop below 7.0 or rise above 8.5, and in dense growths of
sea weeds the pH may reach 9.0. Sea water and hard fresh water are
always on the basic side of neutrality. The pH of different fresh
waters varies from values below 3 to values higher than 10, although
the range of most streams and lakes is confined between pH 6.5 and
8.5. In soft waters with small amounts of combined carbon dioxide
and little buffering, the pH value may fluctuate widely within short
periods.
The
respiration of free-living aquatic plants and animals and the
decomposition of organic matter by microorganisms in the water add
to the carbon dioxide supply of the water medium. The photosyn-
thesis of the plants, on the other hand, tends to reduce the amount of
available carbon dioxide present. In poorly buffered ponds and lakes
variation in the relative rates of carbon dioxide production and con-
sumption produces significant changes in the abundance of free car-
bon dioxide andin the pH of the water. If these water bodies contain
a good growth of green plants, the fluctuation in carbon dioxide and

in pH displays a diurnal cycle as a result of the fact that photosyn-
thesis regularly exceeds respiration during daylight hours. This car-
bon dioxide and pH pulse is generally coordinated with, but recipro-
cal to, the oxygen pulse described above. In the Ohio lake observa-
tion illustrated in Fig. 7.1 the pH rose from 8 to 9 between sunrise
and sunset and dropped again during the night.
A
similar tendency for biological agents to change the takes pH
place in the ocean, but often the actual effect on is too small topH
be measured because of the high buffering capacity of sea water. In
a study of the English Channel throughout a yearly period, during
which photosynthesis varied from its maximum in the spring and
summer to its minimum in the winter months, the pH of the upper
water layer varied only from 8.16 to 8.25.
The relative constancy of pH values in the sea is one more instance
of the stability that characterizes the oceanic environment, and we
may pause for a moment to throw a backward glance over the topic.
We have noted the high degree of constancy in the open sea in re-
gard to salinity, temperature, and oxygen supply, and now in relation
to carbon dioxide, alkalinity, and pH. The ocean thus represents
the largest natural environment in which highly stable conditions exist
in respect to many ecological factors for weeks, months, or even years
Ecological Effects of Carbon Dioxide 265
together. R. E. Coker (1938) on the occasion of his retirement as
president of the Ecological Society of America expressed this thought

in the following
way:
seems to have been the work of a Divine Providence to cover the
It
greater part of the earth with the seas and at the same time to endow man-
kind with no means of making any great
change in them. "I will give you"
he might have said, "dominion over the lands, over the beasts of the field
and the birds of the air, and over the shrubs, trees and but over
grasses,
the oceans you shall have no control I trust
you with small things, but the
greater part of the surface of my beloved mundane sphere I shall keep
under my own lock and key. By means of the seas, which youcan not
measurably modify, I shall proteqt you against your own follies." So the
undisturbed high seas remain the great balance wheel of the terrestrial ma-
chine, the chief source of the rainfall necessary for the continuance of or-
ganic life on land, the real fountain head of the water whose return journey
to the ocean carries the power required for our industries, the
mainstay in
regulation of temperature, the molder of climates. Concerning the signifi-
cance of the seas to organic life anywhere, I do not need to say more to
climatologists, geologists, or biologists. Suppose that man could have put
his plow and machinery and his chemical reagents to work on the whole sur-
face of the earth. Might he not long ago have decided that such a great
expanse of brine was an error in Creation, as he has in effect concluded,
and perhaps properly, with reference to the great areas of forests and grass-
lands? Might he not have converted the saline waters into some other
chemical form? Think how much more tasty or more useful the sea wa-
ters might be were they turned into 20 per cent alcohol, for example, or
into gasoline; and we might have had now to form a committee for the
preservation of some small marine areas to be kept always as scientific "con-

trols"and as refuges for the native population of non-alcohol addicted and
non-gasoline tolerant diatoms, coccolithophores, Salpas, etc. very useless
things in the eye of the practical man,
but none the less interesting to the
field biologist.
But the designer of nature gave us no opportunity to bring such things to
pass. "No," he might have said, "you may change the virgin grasslands
into desert, you may pollute the waters and dam the streams as much as you
please. You may accomplish much that is good by doing these things
wisely. You mean well, but all the same, and for reasons that are quite sat-
isfactory to myself, I put quite
out of your reach the seas as the mainstay
of organic life on your planet." And this is not as facetious as it may
sound.
Dioxide. The most fundamental

Ecological Effects of Carbon
effect ofcarbon dioxide in the aquatic environment is the part that it
of green plants, just as is the case on land.
plays in the photosynthesis
We have seen that in most natural waters as in the atmosphere the
amount of free carbon dioxide is very small. In sea water with a pR
of 8.3, for example, less than 1 per cent of the total carbon dioxide
is in the free form (Fig. 7.9).
present However, both sea water and
hard fresh water contain a large reserve supply of carbon dioxide as
bicarbonates and carbonates. Plants living in water of high pH must
be able to get along by absorbing the tiny amount of free carbon
dioxide present (which is then immediately replaced by dissociation)
or else they must be able to use the carbon dioxide existing in com-
bined form. Many pond plants have been shown to absorb the
HCO :,
ion and to use it in their photosynthesis
(Steemann Nielsen,
1952). Although we do not know whether most marine plants rely
primarily on free or on combined carbon dioxide we find no evidence
that the lack of carbon dioxide ever acts as a limiting factor for plant
growth in the sea.

In soft-water lakes, in which the total supply of carbon dioxide is
small, extensive photosynthesis may reduce the concentration of this
material to such an extent that further growth of the plants is pre-
vented. Water is able to absorb carbon dioxide from the atmosphere
only at a slow rate; the supply from ground water and from respira-
tion and decomposition may be inadequate to meet the demand of
the green plants. The withdrawal of carbon dioxide from the water
also causes a concomitant rise in pH. If the
change in hydrogen ion
concentration is considerable, difficulty is
frequently encountered in
ascertaining to what extent the limitation of photosynthesis is due to
lack of carbon dioxide and to what extent to unfavorable pH values.
The growth of populations of planktouic algae in chemically fertilized
is believed sometimes to be curtailed
fish
ponds by the exhaustion of
the available carbon dioxide. Lime is
commonly added to correct
this situation. Improved growth of rooted vegetation in other soft-
water ponds has been produced as a result of liming.
The abundance of carbon dioxide also exerts certain specific effects
upon the animals of the aquatic environment. The rates of some

developmental and metabolic processes are increased at higher con-
centrations of carbon dioxide and are decreased at lower values, but
other animal reactions are inhibited by high carbon dioxide concen-
trations, so that no general statement of its effect can be made. In
vertebrates, and in some arthropods and mollusks the rate of respira-
tory movements is definitely raised by increase in carbon dioxide
tension, but in others this factor has little or no effect (Sheer,
1948).
The concentration of carbon dioxide in the water influences its
equilibrium in the blood of aquatic animals. An increase of the

carbon dioxide in the blood causes a decrease in the oxygen
affinity
of vertebrate hemoglobins and of some invertebrate
pigments (e.g.,
hemocyanin). In aquatic habitats with high carbon dioxide tensions
animals with the type of blood strongly affected by carbon dioxide
Ecological Effects of Carbon Dioxide 267
in this manner
would accordingly experience difficulty in obtaining
oxygen. Fish inhabiting tropical swamps with water heavily charged
with carbon dioxide have been found to a possess specially adapted
type of blood that is little affected
by the environmental carbon diox-
ide.
Species of fish dwelling in littoral areas where concentrations
of carbon dioxide are above normal have similar
adaptations en-
abling them to extract oxygen from water of wider carbon dioxide
range than possible for pelagic fish.
is
Changes carbon dioxide also tend to modify the pH and the

in
alkalinity of the blood. Alterations of these characteristics can be
compensated for by changes in the abundance of hemoglobin and of

cations such as Na and Ca, But these modifications of the blood
equilibria require time. Fish that can tolerate but little variation in
pH and alkalinity of the blood might well be seriously harmed by

swimming rapidly across a sharp gradient, as at the thermocline, from
water of low carbon dioxide to water of high carbon dioxide or vice
versa.
Another influence of carbon dioxide as an environmental factor is
in relation to orientation. The movementof certain aquatic animals

is affected by differences in concentrations of this gas. Reactions to
increased carbon dioxide may have arisen in relation to the harmful
effects of an excessive concentration of this gas mentioned above or
low oxygen tension commonly associated with it.
in relation to the
A search has long been made for the factor or factors that guide
anadromous fish migrating upstream to a specific tributary within
which their spawning beds are located, and differences in carbon
dioxide tension have been suggested as a possible orienting feature
for salmon. Experiments were conducted by Collins (1952) on the
alewife (Pomolobus), another anadromous fish, by dividing the
stream, in which the fish were migrating, into two channels and add-
ing carbon dioxide gas to the water of one channel. These tests
demonstrated that the alewife can distinguish between small dif-
ferences in free carbon dioxide content (down to 0.3 ppm) and that
the majority (72 per cent) "chose" the stream with the lower amount
of carbon dioxide. Supplementary tests showed that the fish were
oriented primarily by the difference in carbon dioxide itself rather
than by the concomitant change in pH. Whether a reaction of this
sort to naturally occurring differences in carbon dioxide is a chief
factor indirecting migrating
fish into one branch of a stream rather
than into another remains to be investigated. Differences in tem-
perature and other factors

have been shown to be influential in the
and in other observations, and evidence has been presented
foregoing
by Hasler and Wisby (1951) and Hasler (1954) that olfactory re-
sponses involving odor memory may play a part in the orientation of
fish.
Hydrogen Ion Concentration

To turn to a more specific consideration of the ecological influence
of pH in the free natural waters and in the interstitial water of muds
and soils, the hydrogen ion concentration is affected not only by the
reactions of carbon dioxide discussed above but also by other solutes
present, both organic and inorganic. Since any alteration of the pH
of natural waters is accompanied by changes in other physicochem-
the medium, the ecologist must constantly be on guard
ical aspects of
against assuming that the easily measured pH exerts the controlling

influence before determining what effect related changes in the
equilibria of the water may have. Edmondson (1945), for example,

showed that certain sessile Rotatoria are very likely excluded from
lakes by high bicarbonate concentrations, but not necessarily by high
pH.
Manyearly investigators believed that pH would prove to be an
ecological factor of major importance in controlling the activities and
distribution of aquatic plants and animals. Some later workers, go-
ing to the opposite extreme, suggested that the pH of the environment
has little or no importance. Present information indicates that the
effect of pH as a factor is real, but limited and highly variable in its
influence from group to group. As would be expected, pH is gen-
erally of minor significance in the ocean because of its relatively con-

stant value in the highly buffered sea water.
In bodies of fresh water the usual hydrogen ion concentrations en-
countered toward the middle of the pH scale seem to have little dif-
ferential effect on the majority of the inhabitants. The distribution

ofmany aquatic organisms is unrelated to pH over a wide range, and,
when tested in the laboratory, these species often exhibit a tolerance
to pH values well above and below those occurring in their natural
habitats. Speckled trout (Salvelinus fontinalis) are found naturally
in waters ranging from pH 4.1 to 8.5 and were subjected to values
as low as pH3.3 and as high as 10.7 without apparent harm. The
rotifer, Monostyla bulla, is reported to tolerate a range from pH 3.7
to 9.1; the water moss Fontinalis dalecarlica can withstand pH values
from 3.0 to 10.5. On the other hand, some plants and animals thrive
best under acid, others only under alkaline conditions, and others
seem to require a nearly neutral medium. A species of Euglena is
found in water of pH 1.8 draining from mines, and certain bacteria
Hydrogen Ion Concentration 269
and fungi tolerate values as low as
pH 1.4. Other species of animals
and plants live in highly alkaline waters: Lake Elmenteita in the
African Rift with a of 10.7 to 11.2
pH supports a considerable fauna
and flora and
(Jenkin, 1936), Soap Lake, Washington, with a pH
running up to 10 is found tremendous population of
to contain a
rotifers and claclocerans the summer. In contrast to these
during
examples of wide toleration, are instances of organisms that will live
only within a very narrow range of hydrogen ion concentration,
sometimes extending over less than a pH unit. The ciliated pro-
tozoan Stentor coeruleusis
reported limited to pH 7.7 to 8.0. These
aspects ofpH and other physicochemical factors of the aquatic en-
vironment are reviewed more extensively by Allee et al. (1949, Ch.
11), and by Welch (1952, Ch. 7).
The pH of mud and of soils is frequently one of the most important
characteristics of these substrata. A good
general account of the
ecological aspects of this highly complex subject will be found in
Daubenmire (1947, Ch. 2), and more detailed treatments are avail-
able in ZoBell (1946) and Waksman (1932). As we have seen, the
pH of these aquatic and terrestrial substrata varies widely, depending

on the nature of the parent material, the degree of weathering, and
the extent of biological activity, including decomposition. Organisms
inhabiting mud must be capable of withstanding pH values that may

be considerably different from those of the overlying water. The
floras of strongly acid soils, such as those developed from granitic
rocks in the cold temperate regions, are characteristically different
from the floras of alkaline limestone soils. Fungi are the chief or-
ganisms of decay in acid forest soils, whereas acid-sensitive bacteria
and earthworms are abundant only in soils with a more nearly neutral
reaction. The striking difference even in the optimum pH range of
is illustrated by the accompanying examples (Spurway, 1941).
plants
Bog rosemary (Andromeda glaucophylla) 3 0-5

. .
Blueberry (Vaccinium corymbosum) 4 0-5.0

Wake robin ( Trillium erectum) 7.0-7.5
Spleenwort (Axplenium parvulum) 7.0-8.5
Stinking chara (Chara vulgaris) 7 5-8 5
. .
Plants frequently modify the pH of their substratum by the action of

their root secretions and their decomposition products. This activity
accounts for the occasional occurrence in limestone regions of species
characteristic of acid soils. In such instances the pioneer plants have
probably established themselves in microhabitats of lower pH, and

subsequent generations have gradually reduced the pH in the ad-
jacent area. Thus we see that in mud and soils, as well as in the free
water of the aquatic environment, the pH may act as a specific limit-
ing factor for some species, but that it is most generally useful as an
index of the overall conditions.
Calcium Carbonate
Another way in which carbon dioxide exerts an effect upon environ-

mental conditions is
through its influence on the formation and dis-
solution of calcium carbonate (
lime ) . Carbon dioxide in water forms
carbonic acid which dissociates to form bicarbonate and carbonate
ions, and the latter reacts with calcium ions to form CaCO 3 :
C0 + H
2 2 ^H 2 C0 3 ^ H+ + HCOr ^ H+ + COj
CO 8
-
+ Ca++ ^ CaCOa (ppt)
The amount of these ions that will remain in solution
is
greater at
higher hydrogen ion concentrations (lower pH) and, unlike most
salts, at lower temperatures. Since a change in amount of carbon
dioxide affects the equilibrium between the carbonate and bicarbonate
ions, also influences the equilibrium between the precipitated
it
the Ca++ and CO 3 ions in solution. If CO 2 is added ( as

=
CaCO 3 and
by respiration), more H 2 CO 3 is produced, and this will dissociate to
form hydrogen ions, lowering the pH and reacting with the CO^ ions
to form more HCO 3 ions.
~
In nearly neutral solutions the equilibrium
is favorable to this reaction (Fig. 7.9). The reverse situation occurs
when CO 2 is withdrawn (as by photosynthesis), with the result that
more CaCO 3 is formed. In this way lime is often caused to deposit
on or in the tissues of aquatic plants.
As mentioned in Chapter 3, fundamental differences in the nature
of soils depend in part upon the abundance of CaCO 3 The amount .
of lime present isa result of the nature of the parent rock material as
modified by the combined action of climatic and biological agents.
In the pedocal soil-group division the ecological conditions are such
that CaCO 3 tends to be retained in the upper soil horizons but in the
pedalfer soil-group division carbonic acid and other acids, produced

by the metabolism of roots and of soil microorganisms and as by-
products of decomposition, tend to bring about the dissolution of any
CaCO 3 present and thus to allow it to be leached away. Ca++ and
other cations in the soil tend to be replaced by hydrogen ions, and
this causes the soil to become progressively more acid. As soil acidity
increases, ion replacement is accelerated and often results in a serious
loss of cations needed as plant nutrients, as will be considered further
in the next chapter. Carbon dioxide
thus a vital part of a com-
is
plex web of physicochemical interdependencies in the soil involving

carbonic acid, pH, Ca++ ions, and CaCO 3 each of which has
significant
,
ecological relations with the plants and animals living in the soil.
Deposits of CaCO
3 in fresh water are referred to as marl, although
the term is sometimes used for lime deposits in other situations.

also
Marl in fresh water is sometimes the result of the accumulation of
shells, but more often it is produced by the activity of plants. In
hard-water lakes which contain large amounts of Ca ++ ion, the removal
of carbon dioxide by photosynthesis readily causes the precipitation
of CaCO 3 usually deposited on or in the tissues of the plants them-
selves. The pond weed Potamogeton often feels gritty because of the
lime present, and Chara was named "stonewort" for the same reason.
As much as 30 per cent of the dry weight of the latter plant is ac-
counted for by CaCO 3 in its tissues. Certain bacteria and various
types of algae, especially the Myxophyceae, also cause the formation

of marl on the bottom of lakes and ponds. The alga Cladophora
forms a biscuit-shaped deposit of lime about its tissues, and these
"Cladophora balls," commonly attaining diameters of more than 8
cm, may accumulate like paving stones over the bottom or along the
shore.
In soft-water lakes and particularly those with an acid reaction,
we find the reverse situation. Ca ++ and CO-f ions are not only less
abundant but they also tend to stay in solution, with the result in
extreme instances that the deposition of CaCO 3 by the activity of
organisms is difficult or impossible. The Ca ++ ion plays an essential
role in membrane permeability and other physiological processes of
plants and animals.

CaCO 3 is a principal component of the shells
of mollusks and worms, and is incorporated in the exoskeletons of
arthropods where it serves to add stiffness. The skeletons of verte-

brates are composed of about 80 per cent calcium phosphate and a
considerable portion of calcium carbonate and other calcium com-
pounds. It is not surprising, then, to find that some animals are

limited by subminimal supplies of Ca
++
and CO 3 " ions in the water.
Robertson (1941) reports that Gammarus pulex requires at least 5
mg Ca per liter before it can harden its exoskeleton. As a general
rule few mollusks are found in acid lakes; those occurring are specially
adapted to withstand the unfavorable conditions and characteristically

display thin shells, often protected by a covering of chitin. How-
ever, the bivalves Pisidium and Campelorna were recorded by Morri-
son ( 1932) in water of pH 5.1 to 6.1 containing 3 to 5 mg Ca per liter
with shells nearly as heavy as in most alkaline waters. These animals
were also found in water of pH 5.1 with as little as 1 mg Ca per liter,
but the shells were poorly calcified.
Since the ocean tends to be relatively constant in regard to many
of ecological factors, we might expect to find little variation in
its
the conditions of lime formation in the marine environment. Ac-
tually, however, sea water so nearly saturated with Ca** and

is 3 CO =
ions that slight changes in temperature, pH, and carbon dioxide are
sufficient to throw the equilibrium just above or just below the satura-
tion point. The colder and less alkaline parts of the sea are under-
saturated in respect to Ca** and CO
3 ", and the water can take up ad-
ditional amounts; the warmer regions of the ocean are supersaturated

and CaCO readily formed, chiefly through the activity of various
3 is
biological agents. The circulation of water in the great ocean basins

is thus
accompanied by a cycle in which Ca and
*
3
=
ions tend
1
CO
to be taken up in polar or deep water, and lime deposits tend to ac-
cumulate in shallow areas of the tropical seas (Fig. 7.10). As surface
4
POLAR rCst and CoJ from rivers TROPICAL
FIG. 7.10. Schematic longitudinal section of northern half of Atlantic Ocean, in-
dicating the cycle of CaCO 3 The arrows indicate the north-south and vertical
.
components of the oceanic circulation.
water moves toward the poles becomes undersaturated in spite of

it
the new quantities of Ca ++

and C(V added by run-off and river dis-
charge, and the deep return flow remains undersaturated because of
the prevailing low temperatures and low pH values. But, as the
water toward the surface in the tropics, the increase in both tem-
rises
perature and pH bring about saturation and then Supersaturation.

These conditions favor the secretion of lime and the building of coral
formations in the tropical oceans.
Floating in the open water of the warmer seas are countless trillions
of planktonic organisms that use the
easily accessible Ca
++
and 3CO =
ions to build their shells. Prominent these are biflagellated

among
coccolithophorids, foraminiferans, and pteropods. When these or-
ganisms die, their calcareous skeletons rain down upon the bottom,
and, if the water not too deep, their remains accumulate to take
is
part in the formation of globigerina ooze and pteropod ooze. Globi-
gerina ooze is
particularly extensive in the Atlantic and Indian Oceans,
and this type of bottom deposit sometimes contains as much as 86
per cent CaCO In deeper regions of the ocean with lower tempera-
3.
tures and pH values the bottom deposits contain much less calcareous
material. These areas of the sea bottom are covered with red clay,
or have deposits of diatom ooze or radiolarian ooze formed from the
remains of plankton with siliceous skeletons that strongly resist solu-
tion (Kuenen, 1950, Ch. 5).
Calcareous formations around shores and on shoals in the marine
environment are the result chiefly of the activity of bottom-living or-
ganisms. Calcareous sediment resulting from wave action is de-
posited on the bottom and may possibly be supplemented in some
instances by direct precipitation from supersaturated water. The
plants and animals taking part in lime production range from micro-
scopic forms to large solitary species and those that form huge colonial
aggregations. Particularly notable are certain bacteria, specialized
representatives of the Protozoa, Coelenterata, Annelida, and Crus-
tacea, as well as the calcareous algae and a great many kinds of
mollusks (Sverdrup et al, 1942, Ch. 20).
Lime-forming organisms are present in all seas, but they are much
more prominent in warmAlthough the physiological proc-
waters.
esses involved in shell secretion are highly complex (Bevelander,
1952; Wilbur and Jodrey, 1952) and not thoroughly understood, we
do know that shell formation is greatly accelerated at high tempera-
tures. In really cold water shell development tends to be suppressed,
and exoskeletons of many invertebrates are thin or even absent, as in
the naked species of the pteropod Clione. Shells in tropical regions
are typically larger and thicker than those of related species in tem-
perate regions and often display extra spines, ridges, or other pro-
tuberances. The grandfather of them all is Tridacna, a bivalve mol-
lusk with a shell that may approach 2 m in length and weigh 250 kg
(Fig. 7.11). These animals, hidden among the coral formations, are
large enough to be a danger as a man trap since they may close on the
foot of a person walking on the reef or on the hand of a pearl diver
in deeper water.
Since higher temperatures facilitate the secretion of CaCO 3 and
accelerate the growth of organisms producing calcareous structures,
we are not surprised to find that coral reefs are most abundant in the
western portions of the tropical oceans where extensive areas of warm
water are found (Fig. 7.12). Coral animals themselves require a
temperature above 20 C for good growth, and many other organisms
taking part in reef formation flourish only in tropical seas. Foramini-
ferans, millepores, alcyonarians, barnacles, serpulid worms, and mol-
Photo J. I. Tracey, Jr., U. S. Geological Survey

FIG. 7.11. Underwater photograph of the giant clam Tridacna on the reef flat at
Bikini. The specimen is about 45 cm long.
lusks growing in unbelievable profusion among the corals add their cal-
careous parts to the formation. Certain green algae, such as Halimeda,
often contribute large amounts of lime to the floor of the lagoon. The
reef would not grow as an enduring structure, however, if it were not for
the cementing action of lime-secreting Bryozoa and particularly of cer-
tain red algae, such as Lithothamnion. These plants, known as encrust-
ing corallines, or nullipores, often produce a sort of pavement protecting
the new reef growth from destruction by the surf and thus contribute
enormously to reef formation although they are not as conspicuous as

the coral animals (Gardiner, 1931; Ladd and Tracey, 1949; Kuenen,
1950, Ch. 6). The lavish development of calcareous structures (Fig.
FIG. 7.12. World distribution of coral reefs

(Ekman, 1953, after Schott) in rela-
tion to the 20 C isotherm for the coldest month in the year. ( Hutchins and
Scharff, 1947, in the /. of Marine Research. )
Photo R. W. Miner, Am. Museum of Nat. Hist.

FIG. 7.13. View of coral structures at depth of 8 m on Andros Reef, Bahamas.
7.13) and the extraordinary variety and beauty of the reef community
will never be forgotten by anyone who has walked on a reef at low
tide, or, better,has explored one with a diving helmet.
These spectacular coral formations are the result of equilibria in
the circulation of Ca* + and CO 3 = ions that are dependent upon physical
chemical, and biological agents. In the words of Redfield (1941):
"The calcium cycle involves not only exchanges between land, sea,
and the bottom. It includes the consideration of the metabolic cycle,
the CO 2 cycle, the temperature cycle, and the cycle of movement of
water between the surface and the depths. One could not find a
better example of the necessity of considering the environment as a
whole as a unified system."
In the foregoing discussion of the ecological influences of oxygen
and carbon dioxide we have observed that these "sister" materials,
reciprocally involved in photosynthesis and respiration, exhibit a
great contrast in the circumstances of their occurrence and in their
variability. Oxygen is generously abundant in the atmosphere,
whereas the chief reservoir of available carbon dioxide is in the ocean.
We have found that in certain habitats either of these substances may
be so reduced in amount by the physiological processes of the in-
habitants as to curtail the life activities of the
organisms themselves.
We have also reviewed the particularly intriguing further complica-
tions brought about by the fact that carbon dioxide reacts with other
materials in the environment to influence the equilibria involving pH
and CaCO 3 All of these relations with oxygen and carbon dioxide
.
are thus further striking demonstrations of the mutual interde-

pendence of the organism and its environment.
8
Nutrients
NUTRIENTS AND THE ENVIRONMENT

A student once wrote in an ecology paper; "For all
organisms nutri-
tion par excellence" his way of stating that nutrients are a crucially
is
important ecological influence! Indeed, all living things are de-

pendent upon the environment for the supply of energy and of
the materials necessary for their nutrition. Green plants use sun-
light as a source of energy and synthesize carbohydrates from water

and carbon dioxide, obtained as discussed in earlier chapters. To
produce living tissue other materials must also be presentparticu-
larly protein.
Plants build their own proteins from carbohydrates,
nitrogen compounds, and other inorganic substances. These simple

inorganic building materials constitute the nutrients of the green plant.
Colorless plants and animals must have ready-made organic com-
pounds and they feed upon material which is, or

for their nutrition,
recently has been, part of another living organism. Energy is ob-

tained by the oxidation of this organic food. Nutrients for green
plants are thus obtained from the supply of inorganic substances in
the environment, whereas nutrients for animals and colorless plants
are represented by the organic matter derived from other organisms.
Nutrition is obviously and necessarily a reciprocal process. The
in the environment influences the growth and
availability of nutrients
distribution of animals and plants, and the activities of living or-
ganisms profoundly change the abundance of nutrient materials.

The oxidative metabolism of the living organism and the oxidative de-
composition of all plant and animal tissues after death result in the
return to the environment not only of water and carbon dioxide but
also of inorganic materials that can serve again as nutrients for green
plants. These relationships are indicated in the following scheme

which might be thought of as the "equation of all outdoors":
277
278 Nutrients
Green Plants +O 2
H O + C0 +
2 2 Nutrients + Energy /
\Animals
Colorless plants
Modes of Nutrition
The various modes of nutrition in the living world may be classified
according to the
accompanying scheme.
Autotrophic
Holophytic (Phototrophic)
Chemotrophic
Heterotrophic
Holozoic
Saprophytic
Parasitic
Mixotrophic
Autotrophic, or "self-nourishing," organisms can synthesize all es-

sential organic components entirely from inorganic substances; they
are therefore not directly dependent upon other organisms for food.
This group composed principally of holophtjtic forms which use light
as a primary source of energy through photosynthesis is represented
most prominently by the green plants, although the purple bacteria are
also included. The chemotrophic organisms, lacking chlorophyll, are
not able to make direct use of solar energy but instead derive their
energy from the oxidation of certain inorganic substances. This spe-
cialized mode of nutrition is limited to relatively few kinds of organ-
isms. Among those encountered frequently in natural environments
are the sulphur bacteria, which oxidize hydrogen sulphide first into
free sulphur and then into sulphate compounds, and the iron bacteria,
which oxidize ferrous salts to ferric salts.
Heterotrophic organisms require already formed organic com-
pounds as a basis for their nutrition and hence are dependent upon
autotrophic organisms for food directly or indirectly. The most
prominent subdivision of this group is
composed of animals that dis-
play the holozoic mode of nutrition. These free-living animals char-
acteristically ingest solid food and digest it
internally. Saprophytic
organisms commonly lack a digestive cavity and absorb organic food
directly from the environment, usually employing external digestion.
Included in this category are most bacteria, many fungi, some flagel-
lates, and a very few higher plants. The parasitic mode of nutrition
Modes of Nutrition 279
utilized by some species in almost
every taxonomic group of plants
is
and animals. Parasites live on or in other living organisms and obtain

organic food directly from the bodies of their hosts.
Mixotrophic organisms are capable of both autotrophic and hetero-
trophic modes of nutrition of one sort or another. Insectivorous
plants possess special anatomical adaptations for entrapping insects
and digesting them (holozoic nutrition) and at the same time carry
on photosynthesis (holophytic nutrition). Many green flagellates
similarly display the capability of utilizing either of these types of
nutrition, and, since they are also motile, it is thus a constant source
of discussion whether
to classify these forms as plants or as animals.
When light available
is these flagellates can photosynthesize like
typical green plants, but at other times they live saprophytically.

Euglena, for example, is found living in this manner in the decompos-
ing organic matter on the bottom of a pond. Other green flagellates,
are provided with a functional mouth and are capable of the in-
gestion of solid food particles like a typical animal. The green alga
Chlorella is another Dr. Jekyll and Mr. Hyde, since it is capable of
either phototrophic or saprophytic nutrition, and other members of
the phytoplankton may be found to lead this double life.
The fact that most animals and colorless plants require for their
nutrition the organic matter of other organisms means that the food
of these heterotrophic forms has an ecology of its own. The food
organisms themselves grow, change, move around, and die. Further-
more, many of these prey species not only serve as a source of nutri-
ment for the predators but may also influence them in other ways as
part of their environment. For animals, then, our consideration
of
nutrients as an ecological factor overlaps the general consideration
of the presence of other organisms as environmental influences which
will be taken up in subsequent chapters.
But with green plants the situation is quite different. Here the
nutritional needs are substances that are relatively simple in the
chemical sense. The absorption of nutrients from the environment
by plants a relatively direct and much less complicated process than
is
the food getting of animals. This fact is no doubt one reason why
certain aspects of ecology were first developed by botanists. With
the holophytic organisms we can often see more clearly the limitation
of growth and distribution by the influence of nutrients. We shall
the ecological aspects of the nutritional
accordingly consider first
requirements of green plants since these are by far the most important
of the autotrophic forms. Then we shall discuss the significance of
the nutrient factor for animals and other heterotrophic forms, includ-
280 Nutrients
ing the saprophytes involved in the "return" processes of decomposi-

tion and regeneration.
Influence of Nutrients on Green Plants
Nutrients Required. The nutritional requirements of green plants

embrace several major materials and a longer list of minor substances.
The principal elements that go into the construction of plant tissues

are: carbon, oxygen,
hydrogen, and nitrogen. Wehave already dis-
cussed the sources of carbon from 2 , of CO hydrogen from H 2 O, and
of oxygen from O2 , 2 CO
and FLO. An enormous supply of nitrogen
,
exists in the air but atmospheric nitrogen is unavailable to most
plants. In this respect nitrogen presents a complete contrast to 2 CO .
We have seen that the quantity of CO 2 in the atmosphere is

extremely
small but that it is
directly and readily available to plants. Since
nitrogen makes up 79 per cent of the atmosphere, terrestrial plants

would seem be practically surrounded by this gas yet only a few
to
special types of bacteria can use free nitrogen. Aquatic plants are
similarly unable to utilize

directly the large amount of nitrogen gas
dissolved in natural waters. Green plants must obtain their nitrogen
from nitrogen compounds; these are not generally plentiful and are
derived primarily from organic decomposition. Most plants grow
best when supplied with nitrate many can obtain their nitrogen only
in that form but others such as some green
algae can assimilate
nitrite,ammonia, and amino acids (Algeus, 1951).
Twoother elements necessary in moderate amounts for the growth
of holophytic plants are sulphur, which is abundant in the soil and in
the sea as sulphate, and phosphorus, which occurs as phosphate but
is not at all
plentiful. Although some phosphate is derived from the
disintegration of parent rock, the decomposition of organic matter is
the immediate source of this material for most green plants. Essen-
tial in smaller quantities are potassium, calcium, and magnesium.
Potassium originates from potassiferous silicates found in almost all
rocks; the occurrence of calcium has been discussed in the previous
chapter. Magnesium, the keystone in the structure of chlorophyll,

is
usually sufficiently available in the soil and in natural waters.
Other elements known to be necessary for the growth of plants at
least in trace quantities include: iron, manganese, copper, zinc,
molybdenum, and boron (Stiles, 1946). The fact that plant growth
can be affected by zinc in the amount of one part in 200,000,000
illustrates the minuteness of the quantities of these elements that may
be effective. Silicon is used by diatoms in the construction of their
Law of the Minimum 281
shells; other elements such as aluminum, fluorine, and bromine are

taken up by certain plants but seem to have no nutritional significance.
Although sodium and chlorine are commonly present in plant tissues,
these substances are probably not
necessary for most species. Cer-
tain elements found in plants and also in animals have been concen-
trated hundreds or even thousands of times more than
they are in
the environment. This condition is
strikingly true of trace elements
and is where the chemical composition of
easily apparent in the sea
the surrounding water is highly uniform. Silicon and iron are greatly
concentrated in the bodies of diatoms, and titanium, known to occur
in certain marine organisms, has not
yet been detected in free sea
water (Sverdrup et al., 1942, Ch. 7). Vanadium may constitute al-
most 0.2 per cent of the dry weight of Ascidia mentula. Since vana-
dium is present in sea water in a concentration of only 0.3 to 0,6 mg
per cu m, the element has been concentrated roughly four million
times in the body of this animal.
Several other elements, that may or may not be essential to the
plants, are nevertheless significant for the animals feeding upon the
plant material as being either beneficial, e.g., cobalt, iodine, and
nickel, or harmful, e.g., selenium and molybdenum. Few if any of
these minor nutrients exist as elements in soil or in the water; they
mostly occur and must be absorbed as salts or ions in chemical com-

bination with other elements. Above-normal amounts of some of the
trace elements are definitely injurious to plants often with only a
narrow range between minimal, optimal, and harmful concentrations.
For example, an increase of the boron concentration in the soil to
1
ppm will kill some plants, whereas for other species 1 ppm is optimal
and 5 ppm are toxic,
Law of the Minimum. If any necessary nutritive element is com-
pletely lacking, the growth and eventually
the maintenance of a plant
will obviously be prevented. In some habitats all the essential sub-
stances may be present, but one or more of them may exist in con-
centrations so low that certain species cannot absorb them rapidly
enough to satisfy their nutritional needs. Under these conditions

the growth of the plants will be limited in conformity to Liebig's law
of the minimum. This law states that growth is limited by the sub-
present in minimal quantity

stance that is in respect to the needs of
the organism. Liebig's law was originally delineated, and is best
limitation by nutrients, but it is sometimes
applied, in relation to
used in a broader sense to include limitation by other factors of the
environment.
The reader should understand that the limiting substance is the
282 Nutrients
nutrient that is minimal relative to requirements and not necessarily

the one that occurs in the smallest absolute amount. In this sense
the law of the minimum in
ecology is
analogous to the law of com-
bining weights in chemistry. The following equation gives the pre-
cise amounts of Na and Cl that will combine to form NaCl:
23g Na + 35g Cl -> 58g NaCl

If
lOOOg of Na were added to 35g of Cl, no more than 58g of NaCl
would be formed. Under these circumstances the 35g of Cl limits
the total amount of NaCl produced. If, however,
23g of Na were
allowed to combine with 34g of Cl, somewhat less than 58g of NaCl
would be obtained. The 34g of Cl is now the limiting substanfce,
although, in this instance, it is not the smaller quantity of the two
components.
The same sort of limitations occur in the growth of plants, but the
amounts of nutrients absorbed are not nearly as rigidly fixed as the
combining weights in simple inorganic reactions. Much has been
learned from water culture methods regarding the variations in the
quantities of nutrients used by higher plants in normal growth and

also regarding minimal requirements (Hewitt, 1952). Sometimes
the amounts or the ratios of nutrient materials used are completely
altered at different seasons. For example, a certain group of roses
tested in July, took up 450 ppm N and 250 ppm K, whereas in Decem-
ber they absorbed 150 ppm N and 750 ppm K (Turner and Henry,
1948). It has also been learned that a surplus of some nutrients
can in certain instances make up for the lack of others another illus-
tration of the principle of partial equivalence. Experiments have

shown that Na is necessary for the growth of the sugar beet, regard-
less of the amount of K
that may be present, although Na is not
essential for most plants. Furthermore, if
sugar beets are grown
with insufficient K, the symptoms of K deficiency are practically elim-
inated by supplying extra Na (Lehr, 1942).
Similar information on the variation in nutrient requirements has
been obtained from cultures of algae. The amount of one nutrient
taken in sometimes found to depend upon the amounts of other
is
nutrients present,and the ratio may change radically according to

circumstances. Ketchum (1947) found that the rate of assimilation
of phosphorus by the diatom Nitzschia closterium was related to the
concentration of both phosphate and nitrate in the medium, whereas
the uptake of nitrate was independent of the concentration of phos-
phate and related only to the concentration of nitrate. Various indi-

vidual cultures grown in the same medium varied by as much as 50
per cent in their phosphorus content (Ketchum and Reclfield, 1949).

Some cultures multiplied threefold in deficient media and contained
only y5 the normal amount of phosphorus. These algae not only
could make up their deficiency in one
day when again supplied with
phosphate, but also would absorb more of this material than neces-
sary excess phosphate were available.

if
Although a certain degree of flexibility exists in the use of nutri-

ents by plants, in their ability to produce some growth under deficien-
cies, and in the possible partial substitution of one nutrient for an-
other, an ultimate minimum is eventually reached for the availability
of the required elements. If the
plants or the nutrients are mobile,
the lack of nutriment will affect the whole population at about the
same time.
Urban sewage and certain other pollutants discharged into rivers
and lakes provide extra nutrients that frequently cause the growth
of excessive quantities of algae that are unsightly and odoriferous.
Practical application of the law of the minimum has been suggested
in controlling this objectionable growth. Since the treatment of all
the sewage effluents is costly, consideration has been given to the
removal of one essential nutrient only. After the elimination of phos-
phate from the sewage in pilot tests in Wisconsin the growth of algae
was greatly curtailed, although all other nutrients were present in
great abundance. The feasibility of using
this procedure on a large
scale is
being investigated.
Pollution from duck farms bordering Great South Bay, New York,
cause the water of the bay to be rich in reduced nitrogen compounds
and to furnish a low ratio of nitrogen to phosphorus. This unusual
situation in regard to the nutrient factor, in combination with other
has resulted in the suppression of
special environmental conditions,
the usual type of phytoplankton in Great South Bay and the produc-
tion of tremendous quantities of small green algae (Ryther, 1954a).
This alteration in the phytoplankton population appears to have pre-
vented the oysters in the bay from feeding and growing normally,
with consequentserious reduction in the oyster fishery.
Limitation by Nutrients in Nature. In the terrestrial environment
nutrients exert a control over the growth and distribution of plants
a result of deficiencies, although excessive quantities of
primarily as
harmful substances occasionally exclude plants from particular areas.
Availability of suitable nutrients is, of course, only one aspect of the
soil that determines whether a given species can grow successfully;
physical texture,
moisture, pH, and other chemical aspects of the
environment, as well as the climate, must also be suitable. Emphasis
284 Nutrients
should again be placed on the fact that these factors are often related
and their effects mutually interdependent to a considerable extent.
The most commonly deficient nutrients in the soil are phosphate,
nitrate, and potassium. A general discussion of the factors tending
to conserve or to deplete these materials in the terrestrial substratum
was presented in Chapter 3. The existing concentration of these
nutrients depends upon the composition of the parent rock material
and the extent of the modification of it
by leaching and upon the
decomposition of biological products. Our information on the limita-
tion of plant growth by lack of phosphate, nitrate, or potassium in the
soil derived chiefly from agricultural research. A consideration
is
of this vast body of knowledge is obviously beyond our present scope,

but the reader will find further discussions in the Year Books of the
U.S. Dept. of Agriculture, in Meyer and Anderson (1952), and in
Lyon, Buckman, and Brady (1952).
Under natural conditions the phosphate removed from the soil by
plant growth is in part replaced by the breakdown of the parent rock,
although this process is extremely slow and some rocks contain little
orno phosphorus. Of much greater significance in the restoration of
phosphate to most the decomposition of organic matter. Since

soils is
this material is derived from the bodies of dead organisms and from
animal excreta (Hutchinson, 1950), the process represents a critical
instance of the reciprocal action of organisms on their environment.
Replenishment of nitrate and of potassium in the soil is similarly

dependent primarily on the decomposition of organic material, but
the latter element is also derived from potassiferous silicates found in
most rocks. Supplementary supplies of nitrate are derived from am-
monia formed in rain water by electric discharges in the upper air
(Hutchinson, 1944) and from the fixation of free nitrogen by certain
soil bacteria.
Not only must the essential nutrients occur in the soil in sufficient
quantity, but, even more important, they must be present in available
form. Iron, manganese, magnesium, zinc, and sometimes phos-
phorus remain in essentially unavailable states in soils that are too
alkaline. On the other hand, under strongly acid conditions phos-
phorus forms insoluble phosphates with iron and aluminum, nitrates
cannot be readily formed from ammonia, and certain elements may
become so soluble as to attain toxic concentrations. One of the chief
reasons for adding lime to agricultural soils is to correct unfavorable
acidity and thus

to render nutrients available for absorption
by the
plants.
Calcium has important effects as a nutrient in addition to the
part
it
plays in influencing pH and other aspects of the environment. Al-
though some plants do not germinate if more than a trace of calcium
is
present, many more species are dependent upon a generous supply
of this element. Plants such as Aster amellus and Libanotis montana
that require a high percentage of calcium in the soil are referred to as
obligate calciphytes or calciphiles, and their occurrence is limited to

limestone or dolomitic regions. Many species are facultative calci-
phytes. Other plants such as (heather) and Vaccinium
Calluna
(blueberry) that occur where the soil contains less than 3 or 4 per
cent calcium, are known as calciphobes. Investigation shows that
these correlations may be primarily due to the physical structure of
the general richness, or its
soil, its as
indirectly influenced by the
pH
presence of lime, rather than to a direct nutritive dependence upon
calcium. The occurrence of Sphagnum, formerly always mentioned
as a strong calciphobe, is now believed to be
adversely affected by high
concentration of hydroxyl ions rather than of calcium. This compli-
cated subject is discussed more fully by Lundegardh (1931, Ch. 7).
Soils containing large amounts of gypsum (hydrated calcium sul-
phate) or of
serpentine (hydrated magnesium silicate) support
peculiar or impoverished floras. Some kinds of plants require gyp-
sum but others are intolerant of it. Johnston ( 1941 ) reported that in
the deserts of northern Mexico the complex pattern of distribution in
certain plants was controlled with remarkable rigidity by their de-
pendence on gypsum coupled with very successful powers of dis-
semination. Haploesthes Gregii (a grass) and Namn Stcwartii, for
example, were often found in widely separated patches of gypsum

soils no more than 2 or 3 sq m in extent. Within the same genus one
species, Fouquieria shrevei,
was confined to gypsum areas whereas
another species, F. splendens, occurred only on non-gypsum soils.
Serpentine can be tolerated by only
a restricted group of plant
of sparse vegetation growing on soils rich in
species, and the areas
this known as "serpentine barrens." South of Phila-
material are
barrens of this type may be seen as areas about half a mile
delphia
wide and several miles long in which pitch pine, black-jack oak,
scrub oak, and cat briars are the most prominent species, whereas
of large red and white
surrounding regions support good stands
oaks and gums with few if
any pines (Wherry, 1932). A Venezuelan
exhibits a similarly interesting as-
orchid, Epidendrum O'Brienanum,
sociation with iron ore deposits; the plant grows within areas con-
not found beyond their peripheries
taining this ore but
is (Buck,
1949).
In other localities deficiencies in trace elements control plant
286 Nutrients
growth. In certain parts of Australia hardly any production of clover

could be obtained although general growth conditions and the supply
of common nutrients in the soil seemed to be adequate. After a long
study it was discovered that required trace amounts of molybdenum
were lacking in these areas. When the astonishingly small quantity
of 5g of molybdenum per hectare ( % 6 oz per acre ) was added to
the soil, a thick growth of clover more than 30 cm high was obtained.
As a result of generations of study of the nutrient needs of agricul-
tural plants we have accumulated a considerable knowledge regarding
the use of commercial fertilizers for our common annual crop plants.
Bags of chemical fertilizers that we see being applied to farms bear

designations such as 6-8-4 or 5-10-5. These numbers refer to the per-
centages of total nitrogen, available phosphoric acid (P 2 O 5 ), and
water-soluble potash (K 2 O) that are contained in the fertilizer.
Great success has been attained in supplying these commonly de-
by means of such chemical fer-
ficient materials to agricultural lands
tilizers. However, we still know extremely little about deficiencies in
trace elements and in the more complicated organic compounds.
These materials also are leached from the soil by rain, and removed
from the area when crops are harvested. In the movement known
as "organic farming" the plowing-in of green manure and other types
of organic materials is especially recommended with the hope of re-
plenishing as many as possible of the trace elements and organic

constituents that are removed from the soil as the result of farming.
Crops grown with a full supply of all nutrients, minor and major, will
be more vigorous, more resistant to disease and provide more com-
plete nourishment for man and the domestic animals using them
as a source of food.
We have much less information about the needs and deficiencies
of nutrients for natural vegetation in uncultivated areas. For forest
land the time scale is entirely different, and the critical nutrients may
also be different. When a crop of trees is harvested after 40 years'
growth, we do not know what loss of nutrients the soil has sustained
during this long period, nor do we know how to treat forest land to
maintain its productive capacity.
Aquatic plants have the same general nutrient requirements as ter-
restrial plants,and, interestingly enough, two of the elements that
are critical on land, nitrogen and phosphorus, are also likely to be
seriously scarce in the water environment. Potassium is ordinarily

sufficientlyabundant in fresh water, and reference to Table 3 will
indicate that it is one of the six most abundant ions in sea water.
Other materials occurring in small or trace quantities may sometimes
act as limiting factors for the The
growth of aquatic plants. availa-
bility of nutrients in fresh water and in the sea is complicated by the

following (1) the
facts:
supply and the availability of these materials
are involved with circulation and various chemical
equilibria in the
water, (2) the amounts of the nutrients needed by the plants vary
according to their physiological condition, and (3) the growth of the
plants significantly depletes the supply. For a more detailed discus-
sion of these complicated interrelations than is
possible here the reader
should refer to such treatments as those of Ruttner (1953) or
Sverdrup
et al. (1942, Ch. 16).
As would be expected, the amounts of nutrient salts in fresh water
vary widely from lake to lake and also may change greatly within
the same body of water from season to season, as rates of plant growth
and organic decomposition wax and wane. Differences in availabil-
ity of plant nutrients in the water is the principal criterion upon which
European limnologists have divided lakes into three major types:
oligotrophic, eutrophic, and dystrophic. Although difficulty exists in
applying this classification universally (Welch, 1952), the following
general distinctions can be made. Oligotrophic lakes, typically very

deep, are poor in phosphorus, nitrogen, and calcium; electrolytes and
organic materials are low, but oxygen is abundant at all depths and
in all seasons. Eutrophic lakes are relatively shallow, and are typi-
cally rich in plant nutrients and in organic materials; electrolytes are
variable in abundance; oxygen is depleted seasonally and may be en-
tirely absent in the hypolimnion. Dystrophic lakes, occurring prin-

or old mountains, are abundantly supplied
cipally in bog surroundings
with phosphorus, nitrogen, calcium, and organic materials, but the
growth of most lake organisms is limited by the occurrence of high
concentrations of humic substances.
Electrolytes low, and are
oxygen almost
is or entirely absent in deeper water.
In view of the stress laid upon the constancy of the ocean as an
environment in many respects including its salinity, it might be sup-
posed that all nutrient salts in sea water are uniformly abundant.
Such is far from the truth, however. The salinity of the ocean is de-
termined almost entirely by the abundance of the salts listed in Table
3, and these are measured in parts per thousand. Phosphate and
nitrate are not found in this table at allthey occur in quantities of
the order of parts per billion. The paucity of these nutrients is known
to be for the curtailment of plant growth in
frequently responsible
the sea.
Other materials occurring in small or trace quantities may act as
Silicate has been reported upon occa-
limiting factors in the
sea.
288 Nutrients
sion as insufficient for the

production of the siliceous shells of diatoms.
The importance of manganese for the
growth of phytoplankton is
discussed by Harvey (1949). In general, however, we know alto-
gether too little about the occurrence of trace elements in natural

waters in relation to the amounts needed for the healthy growth of
aquatic organisms. Since artificial sea water prepared from the
list of common constituents will riot
support life
indefinitely, we
re-
alize that other materials in minor quantities are essential for con-
tinued growth.
In both the fresh-water and marine environments in-
particular
terest has centered on the of and nitrate not
availability phosphate
only because these nutrients commonly become critically
scarce but
also because their often brought about
depletion is
by the growth
of the plant population itself. We
have little precise information on
the rate of use of nutrients by phytoplankton under natural condi-
tions or their minimum requirements, but a certain amount of data is
available from laboratory experiments. In one carefully controlled
vulgaris of 70 million cells per

set of tests a of Chlorella
population
literreduced the phosphorus content of its culture medium from 1.6
microgram-atoms (/Ag-atoms) per liter to 0.4 /xg-atom per liter and in
another test from 0.8 /Ag-atorn per liter to zero in 24 hours. The
diatom Nitzschia frustndum in a population of 40 million cells per
liter reduced its
supply of phosphorus from 1.6 /xg-atoms per liter to
zero in a similar length of time (Rice, 1949). The growth rate of
these fresh-water species was sharply reduced at the lower concentra-
tions of phosphorus. The multiplication of the marine diatom
Nitzschia closterium was found to be independent of phosphorus
concentration at values above 0.5 /xg-atom per liter, but below 0.3
and especially below 0.15 ^g-atom per liter division rate was dras-
tically curtailed. By summer

the concentration of phosphorus in
many natural waters has been reduced to critically low values for
example to less than 0.1 in the English Channel and to less than 0.25
/xg-atom per liter in western Lake Erie.

A
classical illustration of the seasonal exhaustion of nutrients by
the marine phytoplankton is shown in Fig. 8.1 that represents the
seasonal changes in phosphate and nitrate in the English Channel.
The considerable supply of these nutrients present during the winter
rapidly reduced each spring by the growth of
is the phytoplankton
sometimes to indeterminably small quantities. The concentration of
and phosphate commonly remains very low during the sum-
nitrate
mer when no effective stirring of the water takes place. Meanwhile
the growth of the diatoms is curtailed
by the very depletion the
diatoms have caused. Further cannot take place until the
growth
autumnal breakdown of thermal stratification allows a new
supply of
nutrients tobe brought up from the deeper water layers
by vertical
turbulence. Sometimes a second period of diatom growth occurs
during the autumn, temporarily reducing the concentration of nutri-
ents again.
Eventually reduced light and the more effective stirring
caused by lower temperatures and stronger winds bring this autumnal
growth to an end, and allow the surface layers to regain the higher
concentration of nutrients characteristic of the winter period.
u
I I I ! II I I I I f I I I I I I
1925 1926
FIG. 8.1. Seasonal changes in availability of nutrients for marine phytoplankton as

of phosphate and nitrate in the English Channel.
represented by annual cycle
( Harvey, 1928, Copyright, Cambridge Univ. Press. )
In bodies of water that are suitable in regard to size and drainage,

such as inland ponds, the seasonal depletion of nutrients may
many
be prevented by adding fertilizer to the water and thus
artificially
removing this limiting factor for the growth of the plants. Organic
290 Nutrients
manures have been supplied to fish ponds in the Old World for cen-
turies, but in the 1930s the use of chemical fertilizers was begun on
an expanding scale in the United States. By broadcasting fertilizer,
such as "6-9-2" to "12-9-2," over the pond from the shore (Fig. 8.2),
or from a boat, the concentration of nutrients is maintained continu-
ously at such a level that phytoplankton can flourish through the warm
FIG. 8.2. Broadcasting commercial chemical fertilizer over a pond in Belmont,

Mass., to supply nutrients to the phytoplankton at the beginning of the food chain.
season of the year. The thick growth of planktonic plants provides

abundant food for copepods, cladocerans, and other types of zoo-
plankton and for the bottom fauna, among which chironomids are
particularly prominent, The resulting increase in the abundance
of these invertebrates stimulates the growth of fish such as the blue-
gill
sunfish that feed upon them, and the enlarged population of
these "forage" fish furnishes a rich food supply for predatory species
such as the bass. In this way the production of pan fish or sport fish
in the pond is significantly improved (Edminster, 1947; Meehean,
1952 and succeeding articles in the Symposium on Farm Fish Ponds
and Management). Similar augmentation in the growth of marine
fish by the use of commercial fertilizer to stimulate the development
of phytoplankton and subsequent links in the food chain has been
demonstrated in Scottish sea-lochs (Gross, 1947; Raymont, 1950).
In fertilized Loch Craiglin plaice accomplished 2 years' normal
growth in 1 year (Fig. 8.3), and flounders
accomplished 5 to 6 years'
growth in less than 2 years.
Growth of phytoplankton
resulting from the fertilization of ponds
may produce further
importance. Dense
of ecological
effects
"blooms" automatically cause the of benthic plants by
disappearance
Apr. June Aug. Oct. Dec. Feb. Apr. June Aug. Oct. Dec. Feb. Apr.
Age 1 year 2 years
FIG. 8.3. Diagram showing the greater growth of the flounder, Pleuronectes flesus,
in Loch Craiglin (upper curve) after fertilizer was added compared with normal
growth in an unfertilized loch (lower curve). (Data from Gross, 1947.)
cutting off the light from the bottom. Bottom vegetation is generally
undesirable in farm ponds that contain both prey and predator species
of fish because its
presence allows too many small fish to escape
capture by the large fish. On

the other hand, the growth of the
must not be allowed to proceed to such a point dur-
planktonic algae
ing the summer that the decomposition of the algal material after the
growing season will exhaust the oxygen supply and kill the fish during
the winter. For this reason the addition of fertilizers must be cur-
tailed in northern regions where the winter is long. Carefully con-
trolled experiments carried out in relation to practical fish farming
have valuable quantitative data on nutritional and other
provided
involved in the aquatic environment. At
ecological relationships
the same time the failure of hasty or ill-conceived fish farm practices
form an admirable illustration of the necessity for a thorough under-
as a background for successful con-
standing of ecological principles
servation or cultivation of biological resources.
292 Nutrients
Influence of Nutrients on Animals
As the great majority of animals display the holozoic mode of nutri-

tion, thistype of heterotrophism will be considered first, together with
certain nutritional aspects of parasitism. The nutritional relations of
saprophytes particularly the fungi and bacteria will be included
in the discussion of
decomposition in the succeeding section.
The
nutrients required by animals are proteins, carbohydrates, fats,
salts,and accessory substances such as vitamins. Limitation by the
nutrient factor may result from the shortage of necessary foods in
the environment or from the inability of the animal to obtain and
digest them. The deficiency of the food itself may occur in relation
to abundance or its composition. In some situations the total
its
amount of nutriment is sufficient but the particles of food are so

scattered that they cannot be gathered fast enough to satisfy daily
nutritional needs.
The main bulk of the animal diet consists of carbohydrates, fats,
and proteins. Considerable flexibility exists in respect to the use of
these materials as sources of energy, but at least some protein food is
required in the growth and repair of animal tissue. Insufficient
supply of food to meet energy requirements or deficiency in amount

of protein, minor mineral nutrients, or necessary trace elements may
act as a limitation on the growth, reproduction, or distribution of an
animal.
Free-living animals and parasites obtain these nutrient materials

by feeding on the tissues of other organisms in their environment.
The bodies of the organisms preyed upon as a source of food all con-
tain carbohydrates, fats, and proteins, but in widely varying propor-
and they may or not furnish all the minor and trace ma-
tions, may
terials that are needed. Furthermore, the possible food organisms
must be caught, killed, and dealt with mechanically, in the process
of ingestion, and chemically, in the process of digestion.
Herbivorous animals may be either larger or smaller than the plant
food on which they feed, but carnivores must always be larger,
quicker, or stronger than their prey, or be able to overcome them by
the use of poison glands or some other adaptation. Parasites are
smaller than their hosts. Predators must also be equipped with ap-
pendages and mouthparts that are capable of breaking through the

bark of trees, the shells of invertebrates, the tough skin of verte-
brates, or other types of protective outer covering, and capable of
chewing up the food sufficiently for swallowing. Apparently soft

grass, forexample, contains spicules of silica so hard that only rumi-

nant animals with
specially adapted teeth can feed on it as a regular
diet.
The necessity for alacrity and for the possession of suitable ana-
tomical adaptation for
dealing mechanically with the prey is suffi-
ciently evident, but not so immediately apparent is the necessity for
the animal to
possess the digestive equipment required to deal chem-
ically, with the food once it is inside. Carnivores do not live on hay
nor do ruminants live on meat. In studying the food sources of
animals the possible lack of enzymes for digesting materials that ap-
pear to be abundant and available in the environment is a difficulty
sometimes overlooked, especially in relation to lower animals. For a
long time it was supposed that clams, oysters, and other common
bivalves obtained their nourishment chiefly from the larger, more
conspicuous species of diatoms filtered from the plankton. Most of

these larger types of phytoplankton are now known to pass through the
intestines of the shellfish quite undigested. Physiological investiga-
tion has revealed the fact that these mollusks have no extracellular
cellulase or protease. Hence, they are quite incapable of digesting
relatively large food particles that are enclosed in an intact cellulose
or protein cell wall ( Coe, 1948 ) The nutritional needs of filter feed-
.
ers like clams and mussels appear to be satisfied by the ingestion of
organic detritus or of a type of nannoplankton small enough to be en-

gulfed by digestive phagocytes.
Many specializations have arisen
in relation to the mechanical and
chemical requirements of food getting in natural habitats ( Allee et al.,
1949, Ch. 17). Although most animals are clearly either herbivorous
or carnivorous, these categories are not absolute and some species
like theopossum are omnivorous in their diet. Some carnivores eat
considerable plant material. Herbivores
occasionally consume
animal food, as is exemplified by the fact that reindeer have been
known to eat fish when no other food was available. The diet of
the black bear ranges all the way from sizable mammals to tiny ants,
and from blueberries and honey to salmon.
Many species of animals are highly modified for feeding on special
parts of plants.
The aphids and scale insects have sucking mouth-
parts enabling them
to obtain the sap of their hosts. Some insects
are adapted for boring into and digesting, the wood, the bark, or the
cambium of trees.Not only do certain insects feed exclusively on
the leaves of plants, as do many mammals and a few reptiles, but in
addition the "leaf miners," which live between the leaf surfaces, are
further specialized to feed exclusively on the softer interior tissue.
294 Nutrients
Many animals, among which birds are prominent, show a predilection

for the seeds or fruits of the flowers,
plants, whereas other types eat
or more certain or secretions of the flowers,
frequently, only parts
such as the pollen or the nectar. Hummingbirds, butterflies, and
bees are well knownin the latter
category. Finally, the roots of plants
form the chief or the exclusive food of some burrowing animals. As
a further development of are re-
specialization, certain herbivores
stricted to a food from
plants belonging to a particular taxonomic
group, that is, to one family, one genus, or even one species. An ex-
ample of selective feeding is shown in Fig. 6.15.
Carnivores also may sometimes display an epicurean fastidiousness
in their diets, but specialization on single animal foods is not as elab-
orately developed as it is
among Certain species of bats,
herbivores.
leeches, insects, mites, and ticks, to be sure, are specifically adapted
for bloodsucking, and other instances of preference for certain animal
tissues exist. Animals specially adapted for subsisting on dead or-
ganic matter include the dung beetles and the carrion feeders such as
vultures, flesh flies, and sylphid beetles. Various species of scaveng-
ers are restricted to the lipoids, proteins, keratinoids, or tendinous tis-
sues of the dead animal. For the most part, however, carnivores eat
all
parts of their prey and are guided largely by availability in their
selection of food species.
The distribution of free-living animals, parasites, and other hetero-
trophic forms is
obviously closely related to the occurrence of the or-
ganisms upon which they depend for food. The nutrient factor limits
geographical range in a general way or in a narrowly specific manner,

according to the degree to which each species is omnivorous or re-
stricted in its food habits. Grazing animals
like the
antelope are
found in grasslands; browsing animals like the deer live in forested
regions. Similar broad control of distribution occurs in the aquatic

environment. Off the New England coast cod are caught chiefly
on the banks, such as Georges Bank, where mollusks, crabs, and other
organisms upon which the cod feeds grow abundantly on the firm
bottom. In contrast, the redfish is found chiefly in the deeper water
off the banks where the soft muddy bottom supports populations of
shrimp which form the principal food of this fish (Fig. 8.4).
Control of distribution on a smaller scale is seen in those hetero-
are limited to one type of food.
trophic organisms that Monophagous
animals are those that eat only one species of food organism. Para-
a limited group of species sometimes no
sitesgenerally attack only
more than one species. Saprophytes are often capable of deriving
nourishment from only one type of organic material, and, if this "sub-
T
FIG. 8.4. Contrasting distribuiton of cod,

Gadus callarias (upper figure) and red-
Sebastes marinus (lower as indicated by the catch of the New Eng-
fish, figure)
land commercial fishery in 1936. Each dot indicates a catch of 25,000 kg; broken
line indicates 90 m contour. (Ackerman, 1941, Copyright, Univ. of Chicago
Press.)
296 Nutrients
strate" is
highly restricted in its occurrence, the distribution of the
saprophyte will be correspondingly curtailed.
Many illustrations of the control of distribution by the nutrient
factor for these various types of organisms will occur to the reader,
but perhaps the richest source of examples is found in the insect
world, as fully discussed by Brues (1946). Our common North
American walking stick, Diapheromera femorata, feeds generally on
the leaves of oaks, whereas a giant East Indian member of this insect
group eats the leaves of only a single species of Eugenia found in
Sumatra.
The degree to which insect pests on crop plants are monophagous
is an important consideration in the introduction of new crop species
and in the procedures to be adopted in pest control. The Colorado
potato beetle, which spread northward from its native home in
Mexico, feeds almost entirely on the foliage of the potato plant, and
it sooner or later moves into
every new area in which potato farming
is
begun. The cotton boll weevil crossed the Rio Grande from
Mexico about 1892, and by 1894 it had spread to six counties in south-
ern Texas. Advancing 40 to 160 miles a year, the weevil had infested
more than 85 per cent of the Cotton Belt of the United States by
1922. In view of the terrific destruction caused by this pest to the
cotton crop, it is fortunate that the insect is prevented by its mono-
phagous feeding habit from spreading to other crops or to native

vegetation.
Insects also provide examples of the high degree of specificity of
many parasite-host relationships a matter in which nutrition is prom-

inently involved. The Mallophaga (biting or bird lice) generally re-
strict themselves to hosts of one, or of closely related, species. The
physiological basis for this specificity is strikingly illustrated by the
taxonomic relations of the lice that feed upon the bodies of the cow-
bird. This bird lays its eggs in the nests of other birds belonging to
no less than 158 species, where the young cowbirds would have every
lice of the foster species if the infestation
opportunity to acquire the
depended primarily upon chance contact. Actually, however, the
lice of the cowbird are not those of the foster species but belong to
genera found on other blackbirds taxonomically related to the cowbird.

The absence of certain minor constituents from the diet may have
far-reaching ecological consequences. Lack of sodium chloride in
the food causes deer, elk, and other ruminants to travel long distances
to salt licks. Big Horn sheep, driven by the advance of civilization
into parts of the Rocky Mountains where water is soft, must be pro-
vided with extra salt. A rich supply of lime in the food is
required
Decomposition and Regeneration 297
by snails, mammals, and certain other land animals. As mentioned in
Chapter especially fine race horses are raised in the bluegrass region
3,
of Kentucky. Here the abundance of calcium and phosphorus in the
grass makes possible the healthy bone growth of the young horses.
The firesalamander is an example of an animal that avoids calcium
soils. The special need of aquatic organisms for lime has been dis-
cussed in the previous chapter.
Certain elements and vitamins are required by animals in only
trace quantities, but, if these minute amounts are not present in the
diet, serious deficiency diseases and eventually death result. In re-
gions such as the belt from Washington east to Montana and south
to Colorado, in which iodine is deficient in the soil, the plants, al-
though themselves healthy, do not contain the amount of iodine

needed by animals feeding upon them. As a result goiter develops
in live stock, and also in human consumers, unless extra iodine in
the form of iodized salt is added to the diet.
Zinc, iron, copper, and cobalt are other elements needed in trace
amounts by animals. Zinc is essential in one of the enzymes involved
in respiration, and the other three elements mentioned are concerned
with formation of hemoglobin, the deficiency of which produces
anemia. In certain parts of Scotland and of Australia all attempts
at sheep raising were unsuccessful until the difficulty was finally
traced to the lack of cobalt in the soil. Only very small amounts of
cobalt given the sheep directly, or applied to the grazing areas as
fertilizer, were needed to bring about a remarkable improvement in
the growth of the animals.
The similar need of animals for vitamins obtained from the food is
too well known to require extended comment here. Only minute

quantities of vitamins
are consumed, but these are absolutely neces-
sary and must be obtained directly from the plants that manufacture
them or in some other way, if the animals are to survive. Control of
the lives of animals and of other heterotrophic forms by the nutrient
factor may thus involve the sufficiency in amount of the common food
stuffs,the availability of essential accessory substances in extremely
small quantities, or some other necessary aspect of nutrition.
DECOMPOSITION AND REGENERATION

Since the food of animals and of other heterotrophic forms is de-
rived from green plants or from the bodies of organisms that are di-
the nutrition of
rectly or indirectly dependent upon green plants,
these phototrophic forms is clearly of basic importance in the natural
298 Nutrients
community. We have seen earlier in this chapter that for both ter-
restrial and aquatic run short
plants the nutrients that are likely to
particularly phosphate and nitrate are derived in large measure
from the decomposition of dead organisms. The destructive part of
the biological cycle is thus revealed to be just as critical as the con-
structive part in keeping the wheel of life turning. This idea can
perhaps be expressed no more graphically than in the words of a

student who wrote on an examination: "If it were not for the decom-
position process, the whole world would become choked with the
dead bodies of plants and animals, and this shocking situation would
bring all life to an end."
Processes of Decomposition and Transformation
The release of the nutrients that have been built into the bodies of
plants and animals involves two steps: first, the organic matter must
be decomposed into soluble form and subsequently into inorganic
form, and second, the resulting inorganic material must be trans-
formed into compounds that can be absorbed by phototrophic plants.
Carbohydrates decompose into carbon dioxide and water; fats break
down into these materials and also release phosphate. The decom-
is more elaborate,
position of protein involving hydrolysis into pro-
teoses, peptones, and polypeptides, and then break down into amino
acids, ammonia, carbon dioxide, and water in addition to minor con-
stituents. Special complexes present in living organisms, such as
cellulose, hemicellulose, chitin, agar, and bone, decompose much more
slowly but eventually to the same products.
Living organisms are required to carry out practically every one
of these steps in decomposition. Fungi are especially active in de-
composing organic matter on land; bacteria are prominent in damp

soil and in the water environment. These microorganisms not only
work on the surfaces of solid material but also attack dissolved or-
ganic matter.
The necessity for the transformation of the products of decomposi-
tion from one inorganic form to another is especially apparent in
nitrogen compounds. Since most plants can obtain their nitrogen
only in the form of nitrate, the ammonia or ammonium salts resulting
from the decomposition of protein must be oxidized first to nitrite
and then to nitrate. These transformations, comprising the process
of nitrification, are brought about in both the land and the water en-
vironments by the activities of autotrophic ( chemotrophic ) bacteria:
the oxidation of ammonia to nitrite is
accomplished by bacteria of the
Decomposition and Transformation 299
genera Nitrosomonas and Nitrosococcus, and the oxidation of nitrite

to nitrate is carried out
by bacteria of the genus Nitrobacter.
The supply
of available
nitrogen in the soil and in the water may
be further augmented as the result of nitrogen fixation
by specialized
bacteria belonging to the genera Azotobacter (aerobic) and Clos-
tridium (anaerobic), which are free
living, and Rhizobium, which
live
symbiotically in root nodules on certain higher plants, notably:
peas, and other legumes. These
locust trees, alfalfa, clover, beans,
bacteria take in free nitrogen, build it into their bodies and, in the
Nitrate ( N0 3 ) salts -*-
Plant proteins Animal proteins

and other com- and other com-
plex nitrogen plex nitrogen
compounds compounds
Nitrogen compounds
in dead organisms
also urea and
other wastes
Proteins and
other complex
organic com-
pounds of pro
toplasm
NH 3
ammonia
FIG. 8.5. Diagram of the nitrogen cycle, showing the principal components and
processes. Compounds not included in circles are free in the environment.

(Modified from Principles of Modern Biology by Marsland and Plunkett. By
permission of Henry Holt and Co., Copyright, 1945.)
case of the symbiotic forms, pass it on to their hosts. After these

bacteria or the tissues of the host plants die and decompose, the sup-
is made available
ply of fixed nitrogen that they contain through the
activities of the nitrifying bacteria. Free nitrogen can also be fixed
by certain photosynthetic bacteria, sulfate-reducing bacteria, and
blue-green algae. Other types of bacteria cause denitrification, or
the loss of fixed nitrogen from the environment, by reducing nitrate
to nitrite and nitrite to nitrogen, or even to ammonia (Frobisher,
1944, Ch. 25; Welch, 1952, Ch. 10).

Thus nitrogenous material derived from dead animals and plants
300 Nutrients
is
decomposed and transformed by different microorganisms into in-
organic compounds that can be utilized once more in the growth of
phototrophic plants. The various stepssome of them reversible-
involved in this vital nitrogen cycle are indicated in 8.5. An Fig.
example of the quantitative changes in successive nitrogenous prod-
ucts is shown in Fig. 8.6 depicting the course of an
experiment in
400 r-
5 10 15 20 25 30 35 40 45 50 55
Days
FIG. 8.6. Decomposition of nitrogenous organic matter in mixed plankton under
laboratory conditions, showing the successive appearance of soluble nitrogen com-
pounds. One portion inoculated with diatoms after 45 days; dotted lines represent
uninoculatcd portion. (Von Brand, Rakestraw, and Renn, 1937, Biological Bull)
which marine plankton decomposed and a new growth of the plank-

ton took place after regeneration of the nutrients. Phosphorus, car-
bon, sulphur, and other elements entering the living complex take
part in similar cyclic movements between the organism and the en-
vironment. The carbon cycle is represented diagrammatically in
Fig. 8.7.
Place of Decomposition
In the terrestrial environment the decomposition of organic matter

takes place on and in the soil. Material in particulate form is carried
beneath the surface by organisms and by rain to depths varying ac-
cording to the ecological conditions as discussed in Chapter 3. Mate-
rial in solution may be transported
still further
by percolating water.
The decomposition of this organic matter thus may take place at a
variety of levels within the soil profile, although most of it goes for-
ward in the A horizon. As we have mentioned earlier, a good por-
tion of the products of decomposition may be retained in the soil
when the vegetation is suitable and the rainfall moderate, but some
Place of Decomposition 301
loss of mineral salts takes place under the best of conditions. How-
ever, serious amounts of these valuable nutrients are leached from
soils that are acid and subject to excessive rainfall or that exist un-
der other unfavorable circumstances. Farm or forest practices that
permit soil erosion to take place have resulted in accelerated and often
loss of mineral nutrients and humus from the land.
irreparable
Carbon dioxide
in air and natural waters
Respiration Respiration
Carbon (organic)
compounds of
plant protoplasm
Organic compounds
of animal proto-
carbohydrates Absorption Metabolism
proteins plasm
lipins
etc.
Carbon (organic)
compounds in the
environment
in dead organisms
?aras/fe s
and their
remnants
Organic compounds Organic compounds

of protoplasm of protoplasm
FIG. 8.7. of the carbon cycle, showing the principal components

and
Diagram
circles are free in the environment.
processes. Compounds not included in
Marsland and Plunkctt. of

(From Principles of Modern Biology by By permission
Henry Holt and Co., Copyright, 1945.)
sink
In the water environment dead organisms tend eventually to
the
even though they may have a period of buoyancy. After
or-
has gone into solution, no further tendency to sink

ganism's substance
the decomposing material may
exists. During the period of sinking
have been displaced horizontally by currents for considerable dis-
matter has been decomposed
tances. By the time that the organic
and transformed, the resulting nutrient materials may be very far re-
moved both and vertically from the spot originally in-

horizontally
habited by the organism from whose body they were
derived.
In deep areas of lakes or of oceans the bodies of many dead plants

and animals sink to depths below the euphotic zone before they
302 Nutrients
decompose. Under these circumstances, when

the phosphate and
nitrate are released inform suitable for absorption by green plants,
they will be at levels where light is too weak for photosynthesis.
These nutrients cannot be used again for plant growth until they have
been returned to the euphotic zone by vertical currents and mixing,
as will be discussed in the next section. A cycle thus exists involving
this movement of nutrient materials to the deeper parts of water
bodies and the restoration of the decomposition products to the upper
1000
IE
- 2000
fc
&
3000
4000 LL .// i
1.0 2.0 3.0 4.0

P0 4 -P, Mg-atoms/Z,
FIG. 8.8. Vertical distribution of phosphate in the Atlantic, Pacific, and Indian
Oceans. (From The Oceans by Sverdrup et al., 1942, Copyright, Prentice-Hall,
N. Y.)
layers by vertical transport. In deep lakes and in the ocean the con-
centration of nutrient salts is much greater in the lower strata than it
is near the surface, as may be by the condition in the
illustrated
Pacific (Fig. 8.8). The rates of upward and downward transport for
the year as a whole are presumably in approximate balance, and the
a steady state. However, mineral salts are con-
cycle tends toward
Stagnation in Cycles 303
added to the ocean by run-off and river
tinually being discharge a
process constituting a loss of nutrient material from the land, These
nutrients enter the marine cycle and the
major portion of them is
eventually distributed to the huge reservoir of nutrients in the deep
sea. These important materials are thus being drained
critically
away inexorably from the land environment and
being added to the
accumulation in deep water which is unavailable, except for that
por-
tion brought to the surface again by vertical circulation.
Stagnation in Cycles. Materials move from the environment into

the bodies of plants and animals as they grow, return to the environ-
ment when they die and decompose, and in some instances undergo
complicated transformations and translocations in the environment
before they are again taken up by living organisms. The study of
the geographical distribution of materials used by plants and animals
and their cycles is known as biogeochemistry. A further discussion
of these circular causal systems in
ecology is presented by Hutchin-
son (1948). Materials are not equally abundant in the different
phases of these cycles, nor do the various steps take place at uniform
rates. Biogeochemical investigations show that materials of critical
concern to plants and animals have accumulated in certain places and
represent points of stagnation in the cycles. The gradual augmenta-
tion of nutrient materials in the deep sea is an instance of this phe-
nomenon. Nutrient substances may thus be withdrawn from circula-
tion for longer or shorter periods.
A relatively temporary stagnation is represented by the organic
matter in the soil or on the bottom mud of natural water bodies. The
nutrient minerals contained in this organic matter are unavailable for
In many soils this re-
plant growth until the material decomposes.
tardation of the cycle is beneficial because, as explained earlier, the
formation of humus in the soil has valuable physical effects besides
providing a slow, steady release
of nutrients. Guano deposits, such
as those on the islands off Peru, illustrate a long-term accumulation
of nutrient materials (Hutchinson, 1950). The many cubic miles of
peat, lignite,
and coal buried in the earth's crust represent stagnation
in the carbon cycle for hundreds of millions of years. A
similar
in the calcium cycle is represented by the deposits of chalk,
stagnation
limestone, and coral material.
Great differences exist in the total supply of the various building
materials needed by plants and animals as well as in their availability.
When stoppages occur in cycles of elements likely to be scarce in

available form, such as nitrogen and phosphorus, critical conditions
may arise. An inexhaustable supply of nitrogen exists in the earth's
304 Nutrients
atmosphere, but this represents a reservoir that can be drawn upon

only slowly by certain physical and biological agents. Phosphorus,
on the other hand, is a rare material: only about ôo f * ne earth's
crust is
composed of this element. The only ready
sources of phos-
phorus are the products of decomposition of the bodies of organisms.

We have seen that the supply of in the rocks and in the
phosphate
deep sea can be obtained at only a very slow rate. Therefore the
acceleration of the loss of this element from the land by soil erosion
isa critically serious matter for life in general, and for our agriculture
in particular, since our food
supply is chiefly derived from the ter-
restrial environment. Wells, Huxley, and Wells (1939) have said:
"Phosphorus is the weak link in the vital chain on which man's civili-
zation is
supported."
Regeneration
The return process in the cycle of materials in natural environ-
ments, by means of which nutrients once used are made available
again for the further growth of organisms, is known in ecology as
regeneration. There are two aspects of regeneration: first, nutrients
must be rendered available chemically through the processes of de-
composition and transformation that we have traced in the preceding
sections; second, nutrients must be rendered available spatially, that
is, restored to zones where green plants can grow and resynthesize
organic material. Critical minerals that have been carried below the
level of plant growth must be brought up again before they can be
used.
Mention has been made earlier of the favorable situation in prairie
soils inwhich rainfall is not sufficient to cause serious leaching, and
in semiarid soils in which an upward movement of the ground water
takes place. These effects, augmented by the activity of the roots
of grasses, account to a considerable degree for the restoration of
nutrients to the surface layers in pedocal soils. Regeneration of
nutrients in the pedalfer soils is less complete. In the deciduous
forest region, however, some restoration of materials to the surface
is
accomplished by the fall of leaves and other litter, and by the min-
ing activities of earthworms
and other animals. In the podzol re-
the rainfall produces excessive leaching with the result
gions greater
that nutrient salts are carried beyond the reach of plant roots.
Here we have an analogous situation to that in the deep parts of
the aquatic environment, where decomposition is often completed
at depths below those at which green plants can grow. In water
Rate of Regeneration 305
habitats the return portions of the same currents that
carry oxygen
down to the deeper levels in lakes and in the ocean bring nutrients
up from the richer reservoirs in the lower strata. This upward trans-
port of nutrients by wind stirring, eddy conduction, or mass circula-
tion which also affect other
properties of the water is another mani-
festation of the principle of common
transport.
Rate of Regeneration. The rate at which
regeneration takes place
varies very widely and depends not
only upon the speed of decom-
position but also upon the rapidity of restoration of nutrients to the
growth zone. The rate of regeneration is rapid in some instances
and slow in others. The rapidity with which the processes of decom-
position and transformation go forward depends upon temperature,
supply of oxygen, and other conditions that influence the industrious-
ness of the various kinds of bacteria required. In very cold regions
and in poorly aerated soils and muds the chemical steps in regenera-
tion take place at a sluggish rate and sometimes come to a standstill.
Under these circumstances undecomposed, or partially decomposed,
organic matter accumulates and, if time and circumstances are right,
deposits of peat, coal, or oil
may be formed.
When decomposition and transformation to inorganic compounds
have taken place, the rate at which nutrients are restored to surface
layers depends on
the physical conditions in the soil and in the water.
In oceanic areas of active upwelling of water from deeper layers the
regeneration of nutrients may take place as fast as the plants can use
them. The extensive regions of permanent upwelling off the west
coast of Africa and the west coast of North and South America are
characterized by water rich in nutrients, and consequently they sup-
all seasons. In most temperate waters
port a vigorous plant growth at
vertical circulation is retarded or stopped completely during the sum-
mer. The deep stirring in temperate seas during the winter period
nutrient-rich water to the surface and gives rise
brings a new charge of
to the expression:"Once a year the sea is plowed." In deep lakes of
the temperate region phosphate and nitrate are restored to the euphotic
zone during the spring and fall overturns.
During periods when virtually no vertical circulation takes place
waters must depend upon the nutrients fur-
plant growth in natural
nished by decomposition within the surface layers and by the rela-
tively slow eddy
conduction from deeper strata often further retarded
by the presence of a thermocline.

An adequate rate of replacement
is
just
as necessary as an adequate concentration of nutrients. A
of the phosphate and nitrate in the water might
single measurement
indicate the presence of adequate amounts of these substances, but
306 Nutrients
in view of the rapidity with which the nutrients are used by a grow-
ing plant population, the supply of nitrate and /or of phosphate
will
soon be exhausted unless regeneration keeps pace with utilization.
Quantitative data on the rates of decomposition, regeneration, and
assimilation of phosphate by phytoplankton are available from con-
trolled experiments in outdoor tanks (Pratt, 1950) in addition to lab-
oratory tests (Rice, 1949). Analysis of the seasonal cycle of phos-

phate in the Gulf of Maine by Redfield, Smith, and Ketchum (1937)
for the fractions of this
gives the following approximate estimates
nutrient assimilated by the phytoplankton from the various sources
of supply: 2 per cent from the original inorganic phosphate in the
surface water layer, 73 per cent from vertical transport, and 25 per
cent from decomposition and animal excretion within the surface
layer (Ketchum, 1947). Similar experiments in the terrestrial en-

vironment have been carried out principally in relation to cultivation
of crops, and relatively little is known about the rate of regeneration
in wild areas. For further information on this subject for cultivated
areas the reader may consult the Yearbooks of the United States De-
partment of Agriculture.
Ratio of Regenerated Materials. The
proportions of the materials
are limited by the relative abun-
provided by the regeneration process
50 100 50 100 50 100 150 200

Phosphate (P 2 6)
FIG. 8.9. Correlation between concentrations of nitrate and phosphate in waters

of the Atlantic, Indian, and Pacific Oceans expressed in milligrams per cubic meter
( Redfield, 1934, Copyright, Univ. Press of Liverpool. )
Ratio of Regenerated Materials 307
dance of the elements in the
organic matter that has decomposed.
Only those nutrients will be provided by decomposition that were
in the dead organisms. The fact that chemical fertilizers
present
added to the soil do not replace all the nutritive substances, nor fur-
nish themin the same proportions as those removed
by the growth
of the farm crops, has already been mentioned. Procedures recom-
mended by "organic farmers" in which the unused portions of the
crops are returned to the soil as a green manure and in which animal
manure is used extensively go far toward maintaining a proper bal-
ance of organic matter and the various mineral nutrients, but further
correction of proportions or the addition of trace elements may also
be necessary.
The interdependence of the ratios of materials regenerated and
those of nutrients assimilated is also demonstrated in the aquatic en-
vironment. Analysis of samples of sea water from many localities in
the Atlantic, Indian, and Pacific Oceans reveal a striking parallelism
phosphate and nitrate in spite of the great fluctuations

in the ratio of
in the concentration ofeach (Fig. 8,9). The ratio of carbon to the
other two elements also tends to remain constant, giving the follow-
ing average values:
C:X:P = 41:7:1 grams - 106:15:1 atoms
The ratios of these three elements in the plankton are found to be

very closely the same. Plankton is thus known to take up nutrients
in the ratio in which they are provided in the water; when the plank-
ters die, these materials are restored to the water in the same ratio
(Redfield, 1934). It is not easy to decide how
this unique situation
arose. Do these ratios represent the composition of the primitive

ocean in which species of phytoplankton evolved that could assimilate
nutrients in just these proportions? Or do the ratios represent an
equilibrium reached and maintained by

the activity of nitrogen-fixing
and denitrifying bacteria in the sea? The answer to these questions
must await further investigation.
Let us cast a backward glance over the web of interdependencies
involved in the nutritive relations between the organism and the en-
vironment. We have seen that the lack of nutrients in quantity or
in chemical or mechanical availability, limits the growth
quality, or
and distributionof both plants and animals. As living things grow,
of nutrient materials in their environment;
they reduce the supply
but, when they die and decompose, their substance adds to this sup-
In situations in which either the organ-
ply directly or indirectly.
isms or the nutrients are mobile the shortage of food affects the whole
308 Nutrients
population, not merely the individuals most recently added to the

population. For a replenishment of certain critical nutrients the chief
source is the dead bodies of the animals and
plants themselves.
The
presence of other sets of organisms the decomposers and transform-
ersis required to convert the organic materials to forms suitable for
the growth of green plants, and frequently physical agents are neces-
sary to restore the nutrients to zones where growth can take place
once more.
9
Relations within
the Species
Our discussion of nutrients in the previous chapter leads directly to

a consideration of the presence of other organisms as a part of the
environment. No individual animal could live by itself because it is
dependent upon other organisms for food as well as for other require-
ments. Certain green plants could conceivably exist in isolation for
a period, but, if the
species is to be maintained, progeny must be
produced. The plant would soon be surrounded by young with
which it would be in competition, or in some other relation, so that
it would no
longer be living as an isolated individual. As the popu-
lation of each species increases and as groups of animals and plants
are formed, new relations appear in ecological situations as well as
in others the whole is more than the sum of its more obvious parts.
In this chapter we shall deal with intraspecific relationships and in the
next chapter with interspecific relationships.
ORIGIN OF GROUPS
Groups of individuals of the same species may arise in several ways:
(1) as the result of reproduction, (2) as the result of passive trans-
result of active locomotion.
port, or (3) as the
Reproduction
Some groups are the result of the breeding activity of one breeding
uniteither an individual in the case of asexual reproduction, or a
If the
pair in the case of sexual reproduction. progeny of this breed-
ing unit stay near together, a group will be formed. In some types
of animals and plants the young remain attached to the adults. This
situation is seen commonly in the cryptogamic plants, but instances
are also found among the higher plants. Strawberry plants, for ex-
309
310 Relations within the Species
ample, send out runners that take root, producing new plants, and
these in turn send out more runners. In this way dense aggrega-
tions of the species are formed. Unfortunately for man, briars and
poison ivy also produce thick clumps of individuals by this very suc-
cessful vegetative reproduction.
Similar attached aggregations are formed by certain sessile organ-
isms of the animal kingdom. Many sponges, bryozoans, tunicates,
hydroids, and other invertebrates grow in colonies, and coral animals

produce extensive formations as a result of colonial development.
Animals displaying this growth habit are sometimes very numerous,
but they are not capable of free locomotion, and they have not at-
tained as high a degree of development as non-colonial animals.
In other groups formed by the progeny of a breeding unit the in-
dividuals are unattached but remain together. The parents and their
immediate offspring constitute a family among animals, and the same
term is sometimes applied to the progeny of a single plant. A few
relatives may join the family unit, as has been observed in a den of
wolves (Murie, 1944). Larger aggregations result from a further
extension of family groups. An animal and "his sisters and his
cousins and his aunts" may remain together as a clan, and unrelated
members of the species may subsequently join the group to. form a
larger herd, pack, or flock. Seals, sea lions, wolves, monkeys, prairie
dogs, many ungulates, and various kinds of birds are examples of ani-
mals that form units of this sort. More complex groups of related in-
dividuals are represented by colonies of social insects, such as ants*
bees, wasps, and termites. In the plant kingdom the progeny of
neighboring adults may join to form larger groups of the same

species.
If such a
group is invading a bare area, it is termed a plant
colony, as will be discussed more fully in Chapter 12 in relation to
ecological succession.
In other instances groups resulting from reproductive activities are
not necessarily the descendants of one breeding unit. Aggregations
of young are frequently formed by the simultaneous release of eggs
or larvae within a restricted area. The coincident production of
young by neighboring adults is often set off by some common en-

vironmental stimulus, such as the occurrence of a critical illumination
or temperature. The
nearly simultaneous spawning of oysters, trig-
gered by temperature conditions as described in Chapter 5, results in
the sudden appearance of swarms of oyster larvae in estuaries of the
Atlantic coast.
Passive Transport 311
Passive Transport
Asecond, and entirely different, mode of origin of groups of the

same species results from the passive transport provided by the
medium. Wind often sweeps mosquitoes and other insects from ex-
posed terrain and thusindirectly concentrates them in sheltered places.

Most commonly this mechanical action of the medium in bringing
about aggregations is seen in the aquatic environment. Inhabitants
of streams tend to be swept together in eddies, and the currents of
lakes and oceanic areas often concentrate phytoplankton and zoo-
plankton in the same way, but more slowly. The current system in
the Gulf of Maine, for example, plays a major role in the accumula-
tion of certain types of plankton in the water mass overlying Georges
Bank. The clockwise circulation of water around the margin and
the relatively quiet eddy of homogeneous water over the center of the
Bank have produced conditions in the central area that favor the de-
velopment of Sagitta and of the copepod fseudocalanus but are un-
favorable for Catanus, a closely related copepod. The current sys-
tem tends to concentrate the two former species in the central part
of the Bank and to exclude immigrants from populations of Calanus
and other species developing beyond the margins of the Bank ( Clarke,
Pierce, and Bumpus, 1943).
Another example of the concentration of animals by passive trans-
port is the occurrence of
"plankton traps" found particularly in Scandi-
navian fjords but also to a lesser extent in other estuaries. Charac-
teristically,
a considerable amount of fresh water enters from rivers
at the head of the fjojd, and this flows out at the surface over the
more saline water beneath. Water from moves
offshore in at the
bottom, gradually rises at file head of the and mixes with the
fjord,
outward flowing water. Plankton organisms that tend to remain in
deeper water layers because they are positively geotactic or negatively

phototactic are drawn into the fjord by the deep circulation but are
not carried out again by the surface flow. The result is that a me-
chanical concentration of the plankton takes place in the lower layers
of the f jordy In any species in which tactic responses cause the young
stages to move toward the surface, a horizontal separation of the age

groups results from this current actiqn. The transport of the larvae
of sessile invertebrates to the upper portions of tidal estuaries, as de-
scribed in Chapter 2, is another manifestation of the concentrating
action of this type of differential water movement.
Active Locomotion
Groups of the same species brought about by the active locomotion

of the individuals
may arise ( 1 ) from the guidance of the organisms
towards the same area by oriented
responses to inanimate features
of the environment or (2) from the attraction of the
organisms to
others of their own kind. The formation of groups by active locomo-
tion of course, found most
is,
frequently in the animal kingdom, but
certain motile algae and the swarm
spores of aquatic plants also exhibit
this behavior.
Common Orientation. If the individuals of a species react in the
same way some physical stimulus in the environment, their locomo-
to
tion will bring them to the same
locality with the result that an aggre-
gation will be formed. A familiar example is the clustering of insects
about a source of light at night as a result of their positive phototaxis,
or the attraction of fish and invertebrate animals to a torch held over
the water (Maeda, 1951). Since land isopods, such as the wood
louse, move more slowly, or stop creeping entirely, under moist con-
ditions they tend to collect in damp places. A dead fish on the shore
or a dead deer in the forest acts as a lodestone to which a large num-
FIG. 9.1.
Aggregation of mud snails (Nassa obsoleta)
exposed at low tide in
Barnstable Harbor, Mass.
Common Orientation 313
her of scavengers will be attracted. Other sources of food or shelter

similarly serve as a focal point at which animals from the surrounding
areas tend to congregate (Fig. 9.1). In these instances each individ-
ual reacts independently, and the group is formed as a secondary
consequence.
Sometimes the aggregation is the result of more
complex and long-
continuing reactions. Several kinds of marine worms form aggrega-
tions by reacting individually to some aspect of the lunar cycle and
swimming to the surface of the sea, as described in Chapter 6. When

the animals have formed a huge swarm in this way, they initiate their
breeding activity. Since the eggs and sperm are thus discharged
into the water at the same time and at close quarters, the chances of
successful fertilization are much improved. Consequently, the re-
action of the animals to the physical stimulus that leads to the forma-
tion of the aggregation has survival value in this instance.
The dense schools of salmon preparing to breed in the headwaters
of streams have come into being following the orientation of each in-
dividual separately during the previous weeks while the fish was find-
ing its way for hundreds of miles up from the ocean (Fig. 9.2).
Photo International Pacific Salmon Fisheries Commission

near bank
FIG. 9.2. Sockeye salmon in Adams River, British Columbia, schooling
'
before moving onto the spawning beds. Note dead, spawned -out salmon washed
up on the bar.
When a large number of salmon have arrived at a suitable spawning
place, the fish react more

definitely to each other, pair off,
and begin
their breeding activity. After spawning the adults of the Atlantic
salmon eventually return to the sea, but in the Pacific species the
adults die in the stream and huge numbers of dead fish
may choke
the narrow waterways. The great quantity of decomposing organic
matter thus added to the streams may deplete the oxygen supply
locally and produce further ecological consequences. Nutrient mate-
rials
resulting from the decomposition may possibly stimulate the
growth of plankton on which the young fish depend. When the eggs
hatch, a large number of young fish appears simultaneously in the
headwaters of each salmon stream and represents another occurrence
of aggregation in this species. A further discussion of the ecological
problems involved in the life
history of the salmon will be found in
a symposium sponsored by the American Association for the Advance-
ment of Science (1939).
Mutual Attraction. Another manner in which aggregations of the
same species are formed is by an initial direct attraction of the in-
dividuals to each other. Mutual attraction of individuals is found
among lower animals, but frequently the reaction is largely non-
specific.
If
isopods are distributed over a surface of uniformly low
moisture, the animals stop against the first individual to come to a
halt and soon a dense cluster will be built up. Brittle starfishes placed
in a bare aquarium will move to form closely entwined aggregations,
but, the aquarium contains eel grass, or even glass rods simulating
if
eel grass, the starfishes will remain spaced out in contact with these
objects. In these instances the other individuals present are merely
satisfying physical needs of moisture or of contact. In schools of

fish, flocks and herds of mammals the origin of the group
of birds,
results from mutual attraction on a highly or a completely specific
basis. "Birds of a feather flock together" because of a definite attrac-
tion to others of the same species. The members of these groups
are not necessarily from the same parents and are usually quite un-
related,but during the breeding season units are formed within some
populations in which the members are breeding partners and family
relatives, as discussed above.
Sometimes the reaction to keep in close contact, or to follow others
of thesame species, is very strong indeed. The manifestation of this
in groups of ungulates, such as sheep, has given rise to our term
"sheep-minded." Anyone who has visited sheep country knows the

very great strength of the tendency of these animals to keep close
together. So strong is this reaction that if the leaders of a flock
Mutual Attraction 315
stumble in a ditch, the
remaining sheep will continue crowding for-
ward causing a
"pile-up." Sheep men greatly fear the occurrence
of these pile-ups in which as
many as 500 sheep may be killed within
a few minutes.
Animals forming groups recognize others of the same
species most
commonly by vision, but various other means are also used. Birds
migrating in darkness or fog, or moving through thick vegetation,
probably rely on call notes for keeping in touch with one another.
Sound probably used in the aquatic environment also, to a much
is
greater extent than has been realized. Sound travels through the
water medium much more effectively than through the air medium
quite the reverse of the relationship for other orienting stimuli, such
as light and odor. Various crustaceans, a great many fishes, and cer-
tain cetaceans, including blackfish and porpoises, produce grunts,
whistles, squeaks,and other noises that can be heard under water

and are undoubtedly useful for recognition among
for great distances
aquatic animals (Kellogg, Kohler, and Morris, 1953).

Some species of ants which are totally blind nevertheless manage
to keep close together by the use of their "contact odor" sense. In
tests with bullhead fishes each individual was found to move toward
any object of the same size and color as itself. The fish would then
touch the object with its barbels, and, if the sense organs encountered
a paraffin model, the fish would move away. In this instance the first
reaction causing the fishes to move together was a visual one; other
reactions came into play subsequently. Among land vertebrates
many mammals use odor for recognition.
Sometimes the reaction to school or to form flocks is so strong that,
if no others of the same species are available, the animal will join a
group of another species. Thus we commonly find a few isolated
gulls flocking with a group
of terns on the shore, or a single sander-
the beach in company with a flock of least sand-
ling hurrying along
pipers. Examples among other types of animals will occur to the
reader. An amusing illustration of this tendency to associate with
a group, even of another species, was observed in the large basement
tank at the laboratory of the Woods Hole Oceanographic Institution.
A number of squid had been kept in this tank for some time, and
these animals always moved together in a dense school. One day a
small mackerel of about the same size was placed in the tank. Since
no other mackerel were present, this fish immediately joined the ranks
of the squid, swimming along with them at the same rate in close
formation. As the observer approached the side of the tank, the
shot backwards. Since the mackerel was
squid were startled and
quite unable to move in this direction, he was left isolated and ap-
parently bewildered, turning to right and left. Never was there such
a frustrated mackerel!
The groups that have arisen in the various ways described above
show all gradations of integration and permanency. In brief, we
may note that some groups are brought together mechanically and
often the individuals have no direct relation to one another. In other
groups, such as insect colonies, a definite organization of the mem-

bers exists. Some of the aggregations are of a temporary nature,
lasting for only a few hours, days, or weeks. Starlings are commonly
observed to gather in noisy flocks in the evening and to roost together
in certain
large trees, belfries, or other favorite spots; but the birds
usually disband at daybreak, flying about independently or in small
groups during the daylight hours. Bats follow a reverse daily sched-
ule, coming together to roost during the day and dispersing during
the night. Some
aggregations during the breeding season,
exist only
whereas in other species flocking or herding takes place for the period
of migration only. On the other hand, many herds of large mammals
and many insect colonies are essentially permanent outlasting the
lives of individual members and remaining in existence until de-
stroyed by some unusual environmental change. For a fuller dis-

cussion of this subject and of other aspects of animal aggregations,
the reader is referred to the more extended treatment by Allee
(1931).
EFFECTS OF INCREASED NUMBERS

An increase in the abundance of a species originating in any of the
ways discussed in the previous section results in consequences of

concern to the species itself and also to other interdependent species
of the habitat. The repercussions from the numerical increase of one

will be taken up in subsequent
species on other species chapters.
Here we shall consider the results of population increase within the
species, dealing first
with relations that are generally harmful and
latej*
with beneficial relations. Our present discussion will be cen-
tered on contemporary effects, that is, on effects within the life span
of the individuals or of the populations considered. Long-term ef-
fects on the species may be quite different. Competition, for ex-
ample, is usually harmful to the individuals concerned, but its selec-
tive action in guiding the course of evolution may result in an eventual
benefit to the species as a whole.
Harmful Effects 317
Harmful Effects
An numbers means an increase in competition for the

increase in
necessities of Rivalry between members of the same species is
life.
typically keener than that between members of different

species.
In the words of Darwin (1859), "the
struggle will almost invariably
be most severe between the individuals of the same species, for they
frequent the same districts, require the same food, and are exposed
to the same dangers."
Overpopulation therefore results in a serious
interference of one individual with another sometimes in a passive
or indirect way, at other times as direct aggression or even cannibal-
ism. The accumulation of metabolites often curtails the further
growth of the species within the area, and the simple matter of oc-
cupying available space imposes a mechanical limitation on some

populations. Since established land plants are fixed in position, the
sphere of influence of each individual, and the area from which it

must draw its necessities, are sometimes easier to observe than those
of active animals. When a population of plants has increased so that
the individuals are growing close together, their roots compete for
nutrients and for water and their tops compete for light. As com-
petition becomes more intense, growth rate is

correspondingly re-
tarded.
A ready-made record of the effects of competition was found in
the cross section of the trunk of a locust tree that had grown in Bel-
mont, Mass., and was blown down

in the hurricane of 1944 (Fig.
the date of the first year of the tree's

9.3). By counting growth rings
life was determined as 1929. At that time an open field was aban-
doned and a few trees seeded themselves in the area at widely spaced
intervals. For the first few years thereafter, the tree grew rapidly,
but, as size increased, competition with neighboring trees for nutrients,
for water, and particularly for light became progressively more se-
rious. The result may be observed
in the diagram as progressively
smaller growth rings in 1934, 1935, and 1936. In the spring of 1937
the land was cleared of most of the trees for the construction of a
house. With competition removed the growth of this locust tree
was "released," to use the term of the forester. More wood was
added during the growing season of 1937 than had been added in
any of the previous 4 or 5 years, and growth continued at a high rate
for the remainder of the life of the tree.
The form of development of plants as well as their growth is af-
fected by competition. The growth form

of a solitary tree is com-
pletely different from that of a tree of the same species that has de-
veloped in a stand closely surrounded by other individuals. In the
latter situation the lower branches are killed by the reduction of light,
or the growing buds and branchlets are knocked off by the branches
of trees near by, as they whip about in the wind. The smaller,
neighboring trees that bring about this
pruning action are referred
to as "trainers" byforesters, since they cause the tree to develop
without lateral branches in a form that will later be suitable for saw
timber.
Â^ ^tet ^
.Growth ring for
the season of 1944
Inner bark
Outer bark
Land cleared
spring 1937
Growth
released
Progressive
overcrowding
Competition
retards growth
FIG. 9.3, Cross section of locust tree (Robinia Pseudo- Acacia), 17 cm in diam-
eter, grown in Belmont, Mass., showing effects of early overcrowding and sub-
sequent "release."
Many examples of the harmful effects of overcrowding could be

cited for other types of plants and for animals. find the same We
of at work among populations of microbes as
principles competition
among forest trees. Bacteria deplete their supply of nutrients as they
multiply and produce an accumulation of metabolites, until the fur-

Harmful Effects 319
ther growth of the colony is
prevented. Organisms continuing to live
in the same medium tend to
change or to "condition" it. Condition-
ing of flour by laboratory populations of the flour beetle Tribolium
confusum has been shown to cause reduced fecundity, extended dura-
tion of metamorphosis, and increased
mortality (Park, 1941). The
same kinds of harmful changes are brought about by overcrowding
under natural conditions. In some situations aquatic animals
may
aggregate to such an extent as to exhaust the oxygen supply; else-
where they may exhaust the food supply, produce harmful metabo-
lites, displace one another mechanically. Of several hundred
or
oyster larvae originally setting on an old shell or other small object,

only a few will find sufficient space to develop into full-sized
oysters;
all the others will eventually be killed by crowding. Where com-
petition for space is keen, oysters grow in a long slender form, un-
desirable for the market. Oystermen in cultivating their beds com-
22
I I I I I I I I I
20
18
16
14
$ 12
I
I
10 20 30 40 50 60 70 80 90
Mean flies per bottle
FIG. 9.4. Curve showing decrease in rate of reproduction in Drosophila as cul-
tures become more crowded. ( Pearl and Parker, 1922. )

monly break clusters of young oysters

apart to insure that the animals
develop in the well-rounded form characteristic of solitary oysters.
The harmful effect of excessive numbers of animals is sometimes
manifested as an interference with breeding. The progressive re-
duction in the reproductive rate of Drosophila with increased density
of the population is shown in Figure 9.4. As the flies become more
crowded, interference with feeding and possibly also with oviposition
is
experienced.Frequent collision and interruption of feeding re-
sult ininadequate nourishment and lowered fecundity although the
supply of food may be ample (cf. Robertson and Sang, 1944). In
some kinds of animals the ratio of males to females is also significant
in controlling success of
reproduction. Although many animals do
best when the two sexes are present in equal numbers, a one-to-one
ratio is detrimental for some
polygamous species. Among
pheasants,
for example, each cock normally maintains a harem of about five hens,
and, if more males are present in the population, the consequent fight-
ing and disturbance of the incubating females greatly reduces the
success of the breeding. With species requiring highly specialized
places for breeding or nesting, the first effect of an enlarged popula-

tion may be a critical shortage of breeding sites.
Photo by Swem
FIG. 9.5. "Deer-line" formed by denuded branches of Douglas fir and Ponderosa
pine, caused by browsing of overabundant deer in Oregon.

Harmful Effects 321
Perhaps the most generally harmful effect of

increasing numbers
among animal populations is the competition for food. In the photo-
graph, Fig. 9.5, a "deer-line" is distinctly visible on the vegetation at
a height of about 2 m. This denudation of the trees has been caused
by the browsing of an excessive number of deer. With the shortage

of food the growth of the deer
population will be curtailed, and, if
all the edible vegetation
throughout the area is removed, the deer
will be in danger of starvation. In extreme instances the
vegetation
may be permanently injured with the result that the area will sub-
sequently not support as many deer as previously.
The effect of competition for food on the growth of bluegill sunfish
is well demonstrated
by records of farm fish ponds. In one test a
pond was stocked in March with fish
averaging 5.7
g.
When weighed
again in June of the same year, the average size of the fish had in-
creased to 76 g. During this month each pair of sunfish produced an
average of 4000 young. Since the young were not removed either
artificially or by a predatory species of fish, the new generation en-

tered into direct competition with the adults for food. When the
pond was drained in November of the same year, the average weight
of the parent group had decreased from 76 g to 54 g, showing that
Swingle and Smith, 1942

FIG. 9.6. Effect of overcrowding on the growth of bluegill sunfish. Both fish are
1
year old: upper specimen from a pond stocked with
3750 fish per hectare (1500
per acre); lower specimen from a pond stocked with 450,000 fish per hectare
(180,000 per acre).
these fish had not only failed to gain but had actually lost weight
as a result of excessive competition. Another instance of the curtail-
ment of growth of sunfish brought about by overpopulation is shown
in Fig. 9.6;
Beneficial Effects
In the previous section we have seen that, if the population of a
species within any area continues to increase, a point will eventually

be reached at which harmful effects are produced. Before this point
is reached, the members of the population may receive definite bene-
fit from the presence of others of the same species. An isolated in-
dividual or a single pair of organisms is often not as able to deal
successfully with the environment as a group. This subject has been

extensively discussed by Allee (1931 and 1951). Moderate increase
in abundance of a plant or an animal may afford protection from
enemies or from physical features of the environment. It may ac-
celerate reproduction and improve survival. The degree and nature
of activity is also influenced by the density of the population ( Schuett,
1934). Learning by fishes and other animals has been shown to be

more rapid when other individuals are present than when they are
isolated (Welty, 1934). Groups of the same species also make pos-
sible a certain division of labor and the beginnings of social organi-
zation. Some of these effects of increased numbers that are bene-
ficial within the species will be considered in the following sections.
Protection. Animals and plants in groups often protect each other

against harmful features of the environment without any special
group organization. In a thick stand of trees a higher humidity
will be maintained and a better resistance to wind and water erosion
will be experienced than if the same individuals were growing in a
widely spaced manner. Furthermore, since the blowing away of

fallen leavesis
largely prevented in a dense grove, the humus and
moisture content of the soil is increased. Groups of animals have
been shown be less susceptible to poisons, oxygen lack, extreme
to
temperatures, and other environmental dangers than single indi-

viduals.
The
large number of honeybees living close together in a hive
enables these insects to modify the temperature inside their micro-
habitat to an extent that would be utterly impossible were the bees
living as solitary individuals. As winter conditions arrive, the bees
are able to maintain the temperature within their cluster in the hive
at 25 to 30C by increased muscular activity. During hot days in
summer the bees force air through the hive
passages with their wings,
thus increasing evaporation and
lowering the inside temperature.
Owing to this primitive "air-conditioning" temperature inside the hive
fluctuates only
slightly in spite of extreme external conditions ( Uvarov,
1931).
A group of animals is often protected against its predators by the
simple effect of numbers. A flock of small birds does not attempt to
fight off an attack by a hawk and probably could not do so; hawks
generally obtain their prey by picking off stray individuals. Perhaps
the hawk is less likely to be successful in
catching his prey from a
flock because of the confusion of numbers,
just as a tennis player is
distracted by having his opponent throw three balls to him at once,
with the result that he does not succeed in catching any one of them.
A favorable effect on survival of an optimal degree of crowding
has been found in laboratory cultures of Drosophila and of Daphnia
and in natural populations of the sessile rotifer, Floscularia. At popu-
lation densities of 5 and 10 per 50 cc of culture medium Daphnia lives
considerably longer than at densities of either or 25 per culture
1
unit in tests at 25C. At 18C mortality appears to be least at a

density of about 75. Lower metabolic rate or better control of harm-
ful bacterial contaminants by the supraminimal populations are sug-
gested as possible explanations (Pratt, 1943). The mean length of

life of Floscularia is practically doubled if the young rotifer, instead
of living a solitary existence on a water plant, attaches to the tube of
an older Floscularia and thereby establishes a small colony. No ex-
planation of this striking fact has come to light

(Edmondson, 1945).
Influence on Reproduction. Since most animals and plants re-
is obvious that too
produce by sexual methods, it great a scarcity of
individuals of the opposite sex will reduce the rate of breeding. In
an earlier chapter we have called attention to the probability that
this constitutes a serious limitation for deep sea fish and perhaps
also for whales and other aquatic animals of low population density.
Luminescence and underwater sound have been suggested as possibly
serving as aids for males
and females to find each other in the huge
expanses of the ocean. The same limitation affects land animals and
distributed.
plants that are very sparsely
A similar situation on a much smaller scale
is
presented by flour
beetles dispersed in a large uniform volume of flour.
Investigations
with Tribolium confusum have shown that when only two individuals
are present in a "microcosm" of 32 g of flour, the number of young
less than when four individuals
produced per female is considerably
inhabit the same volume. The explanation is that fecundity is in-
creased by recopulation, and recopulation increases with the density

of the population. At densities higher than four, however, the fact
own eggs progressively curtails the success
that the beetles eat their
of reproduction. This cannibalism eventually sets the upper limit of
the population (Park, 1941). In experiments with ciliates the re-
productive rate was found to increase with the size of the initial popu-
lation. This effect may be due to the release into the medium of a
beneficial substanceproduced by the protozoans, to the low absorp-
tion per individual of some inimical material originally present, or
to more adequate control of the bacterial population ( Mast and Pace,
1946; Alice, 1951).

Another effect of numbers on success in reproduction involves re-
actions, some of which may be considered to be in the realm of
group psychology. many animals breeding activity is initiated

In
only when an aggregation of more than a certain minimum size has

been formed. The necessity for the existence of a sizable group in
order that reproductive behavior will be called forth is
clearly seen
among social insects and among certain fishes, birds, and mammals,
as well as among other types of animals.
This effect of numbers is particularly striking in birds that breed
in colonies ( Darling, 1938 )
Anyone who has visited an island where
.
terns are nesting or a cliff where gannets have established a rookery
(Fig. 1.2) will have obtained an indelible impression of the atmos-

phere of excitement caused by the sight of thousands of wheeling and
darting birds and by the sound of their screaming calls. The visual
and auditory stimuli this uproar are necessary, or at
produced by
least valuable, in preparing the birds psychologically for mating as
well as for setting off and integrating the elaborate behavior patterns
involved in nest building, incubation, and feeding the young. If
there are too few birds in the colony, the reproductive cycle will not
be successfully completed. Furthermore, the large concentration of
birds often induces synchrony in reproduction, and this provides a
certain amount of protection since predators cannot destroy a serious
proportion of the eggs and young if they all appear within a short
period. Even without synchronous reproduction large colony size
has the advantage of a relatively smaller periphery where young may
wander away or be attacked by enemies. This need in certain species
forgroup stimulation accounts for the failure of isolated pairs to breed
and for inefficient breeding as seen, for example, in small colonies of
gannets (Fisher and Vevers, 1944). An extreme illustration of these
principles is the report that a minimum of 10,000 birds, nesting at a
mean density of 3 nests per sq m, is necessary for the establishment
of a successful
breeding colony of the guano-producing cormorant
on islands off Peru (Hutchinson, 1950). Among mammals, popula-
tions of muskrats smaller than about 1
pair per linear mile of stream
or per 35 hectares of marshland
appear not to breed successfully
(Errington, 1945).
Degree of crowding can influence the sex and structure of some
species and the coordination of these features with the natural life
cycle. Crowding has been shown to influence the sex ratio in certain
cladocerans, in Bonellia, and in Crcpichda. A/own, a common clado-
ceran inhabiting temporary ponds, reproduces parthenogenetically
under ordinary conditions with the production chiefly of females. If
the population becomes crowded, however, males begin to appear.
This reaction is
obviously related to the reduction of space attending
the drying up of a pond and necessitating the production of resistant
eggs if the population is to survive. Since resistant eggs are produced

only sexually, males must be present in the population. The produc-
tion of males
by crowding is thus nicely attuned to the needs of these
animals under the exigencies of life in temporary ponds.
In the aphids, or "plant lice," which attack vegetation, the winter
eggs hatch into wingless females; these reproduce parthenogenetically

under ordinary circumstances, with the result that the population on
each plant grows rapidly. If the plant host becomes overcrowded
and consequently withers, the destruction of the whole group of
aphids is threatened. However, crowded conditions react on the
aphids in such a way as to bring about the production of winged
females. This extraordinarily neat adaptation makes possible the
migration of many of the aphids to other plants, relieving the con-

gestion at home and establishing new centers of population growth.
Later in the season males appear and sexual eggs are produced that
tide the population over the winter period and complete the cycle.
Another effect of the size of breeding population is in relation to
of a
genetic elasticity and, hence, the adaptability of the progeny
species to varied conditions. Most species of plants and animals in
nature are composed of a great many biotijpes, that is, types of indi-
viduals that grow and reactdifferently because of different genetic
constitutions. Owing to varying environmental conditions certain
biotype groups become established in different ecological regions
of
the range of each species. These ecological subdivisions of the
species are known as ecotypes

and are genetically distinct races.
Since the ecotypes are interfertile, they are placed in the same taxo-
nomic species. Ecotypes are sometimes recognized as subspecies,
but in other instances they are not sufficiently distinct morphologically
to warrant that designation. Each ecotype is the result of selection

by its environment and has become specially adapted for a particular
set of conditions. in different
Wide-ranging species are represented
parts of their ranges by different ecotypes.
A
small population of an ecotype, particularly if it is isolated, will
tend to become less variable genetically because of inbreeding; that
is, will come to contain fewer biotypes.
it With less adaptability
the population will be less likely to survive bad conditions and will
fail to
respond quickly to the occurrence of good conditions. In con-
trast, a larger population with more biotypes is more likely to include
some individuals that can withstand adverse circumstances and that
can take advantage of new variations in the environment. These
relations probably account, in part at least, for the fact that certain
reduced populations fail to spread widely or to recover a former
abundance although environmental conditions appear favorable
(Cain, 1944, Ch. 16; and Allee, 1951, Ch. 4). The isolation of seg-
ments of a varying population also plays a major role in influencing
the course of evolution. This large topic, which is beyond the scope
of the present book, is considered in the works referred to above and
also by such authors as Elton (1930), Mayr (1942), and Lack
(1947).
An example of the failure of a small population of an ecotype to
spread readily isfurnished by the distribution of wild irises in Canada
(Fig. 9.7). Iris setosa has considerable morphological variation and
isdistributed widely in western Canada and Alaska. The subspecies

Irissetosa var. canadensis, however, exhibits high morphological con-
stancy and is limited to the Gulf of St. Lawrence region. This sub-
species survived the ice age, during which other luxuriant types may
have succumbed, but as a relict ecotype it emerged so uniform genet-
ically that during the intervening centuries it has been able to repopu-
late only the immediate area and is slow in adapting itself to other
environments.
The disadvantage of low numbers may also be illustrated from ex-
perience in the oyster fishery. Around the shores of Great Britain,

and elsewhere, oyster populations have been greatly reduced by over-
fishing. However, after restrictions were placed on the amount of the
oystermen's harvest, or in other areas after fishing had been aban-
doned entirely because of unprofitable yields, the populations often
failed to rebound to their previous large sizes. Gross and Smyth
(1946) believe that the explanation is to be found in the lack of
these sessile animals.
genetic flexibility in the small isolated groups of
A small population not only may be unable to grow rapidly and to
spread but also, after it has been reduced below a critical size, it may
be unable to hold its own. This latter
possibility introduces the prin-
ciple of the minimum population which states that in order for a popu-
lation to survive
indefinitely in an environment its numbers must be
maintained above a critical minimum.
A classic illustration of this
principle is furnished by the fate of the
heath hen, a bird that was
formerly abundant in Massachusetts and
may have been distributed from Maine to Delawo.re (Gross, 1928).
By 1880 the heath hen was restricted to Martha's
Vineyard Island,
and a realization of its low abundance led to the establishment of a
large reservation for the birds on the island. The heath hens in-
creased to about 2000 in 1916, but a fire, a
gale, and a hard winter
with a great flight of goshawks decimated the
population, leaving
fewer than 50 breeding pairs. Numbers continued to decline
irregu-
larly until only 20 birds were counted in 1927, and the last bird was
seen in 1932. In spite of elaborate
protection this species could not
be saved from extinction once the population had been reduced
below its critical size.
Division of Labor. Beneficial effects of increasing numbers
may
also be brought about by a division of labor made possible within
FIG. 9.7. The discontinuous range of Iris fietosa (open circles) and its varieties:
canadensis (small black dots) and interior (large black dots). Hatched lines in-
dicate the maximum extent of the Pleistocene glaciations. (Cain, 1944, modified
from Anderson, Copyright, Harper and Bros., New York.)
the group. In some sessile colonial animals, such as certain of the

coelenterates, different individuals are specialized for definite func-
tions. This differentiation of zooids permitting a division of labor
within the colony reaches its highest development in siphonophores
such as the Portuguese man-of-war. Different zooids of this animal
are specialized for flotation, protection, nutrition, or reproduction.
In some siphonophores zooids are also specialized for locomotion and
produce a movement of the whole colony by rhythmic ejection of

water from their cavities. Certain colonial Protozoa and Protophyta
also show a division of labor. Although this development among
unicellular colonial forms very primitive, its advantages may have
is
led in the past to the evolution of multicellular organisms.
Among organisms that form groups but in which the individuals re-
main separate we find other types of division of labor. The most
fundamental and widespread of these is the differentiation of the
species into two sexes. This division of labor has arisen in both the
animal and plant kingdom, and interestingly enough the appearance
of sex occurred long after plants and animals had evolved from their
common ancestral unicellular form. We have here a remarkable case
of parallel evolution in which the same adaptation arose indepen-
dently on at least two occasions.

A variation in activity among members of a group without any
special morphological adaptation is seen in the phenomenon of social
dominance. This
the establishment within the group of a social
is
hierarchy in which an animal of higher position outfaces or drives

away an animal of lower position ( Allee, 1951 ) Such "peck orders"
.
were first
recognized in flocks of fowl, but now dominance-subordina-
tion relations are known to exist among certain groups of fishes,
lizards, rodents, ungulates, carnivores, and other animals (Collias,

1944, 1952). The peck order among hens establishes the social posi-
tion of the birds in the flock. A dominant hen has attained the right
to peck a subordinate hen without being pecked back. When the
become established, the group func-
social organization of a flock has
tionsmore smoothly and less fighting occurs, since protocol is recog-
nized by all members for all group activities.
Another type of division of labor in a group is leadership. Within
a flock of goats one animal will become recognized as the leader in the
wanderings of the group; but this animal is not necessarily the domi-
nant individual in the sense of the term used above. The existence
of a definite leadership was also demonstrated in a band of
monkeys
inhabiting a tropical island. One individual always led the way when
the band moved through the aerial pathways of the treetops. When
the leader was shot, the band was
reported to remain in a confined
area of the island, and did not take its usual trips to other parts of
the island.
Cooperation without any special morphological modifications among

the individuals and with a minimum of organization is another ad-
vantage often resulting from the formation of groups. When at-
tacked, musk ox form in a circle with their horns extending outward
and thus secure for themselves a protection that would not exist if
each animal attempted to defend itself individually. A group of
beavers working together can dam a stream that could not be success-
fully dammed by animals operating singly. In a penguin colony
some adults guard not only their own young but also the young of
others while the remaining parents leave the area in search of food.
Feeding cormorants are observed to maintain a

rough line as they
swim and dive and thus presumably improve the effectiveness of their
fishing (Bartholomew, 1942).

A somewhat more elaborate division of activity within the group
is found in certain species during the breeding season. In sea lions
(Fig. 9.8), fur seals, elk (Fig. 9.9), deer, and other mammals, as well
as various game birds, the population of a region becomes organized
into breeding units each consisting of a male, his harem, and sub-
sequently their young. After the mating season the harem groups are
broken down, and the animals may reorganize themselves into sepa-
ratebands of males, females, and young for the remainder of the year.
Group organization in the fur seals (Callorhinus ursinus) may be
taken as an example. The bulls arrive first
early in May at the
breeding grounds on the Pribilof Islands, and each stakes out a

breeding site for himself which he jealously guards. When the cows
arrive in mid-July, each of the larger and more senior of the bulls
collects a harem for himself within his own territory. The remain-
ing bulls, kept away by ferocious fighting, form a group of disap-
pointed bachelors. Bulls with harems are continually being chal-
lenged by the bachelors, and, although each established bull drives
off the intruders for one or more seasons, eventually each bull ages
and after losing a fight or a series of fights he is forced to relinquish
his harem to a new master. Soon after the females are organized
into harems the pups resulting from the previous breeding season are
born, and subsequently mating with the bulls takes place. When
breeding has been completed, the harems disintegrate. After a
period of feeding and of nursing the pups, the seals reorganize them-
selves in groups consisting of males only, and other groups made up
of females and pups. In these groups the seals leave the Pribilofs
..
'",.. ,. ;*% îit?^ V T
Photo Alfred M. Bailey, National Audubon Society

FIG, 9.8. Breeding groups of sea lions on San Benitos Island off the west coast of
Mexico. Each bull sits erect guarding his territory and presiding over his harem
and pups.
fhoto U. V. National I'arK Service

FIG. 9.9. Bull elk and his harem in Rocky Mountain National Park in late
September,
in October and travel to their winter quarters along the California

coast (Allen, 1870). With this social organization in operation
among the fur seals, the non-breeding males can be killed for their
pelts without reducing the productivity of the herd; hunting is re-
way by international agreement. Similar organization
stricted in this
of the herd into harems during the breeding season and into larger
non-family groups during other parts of the year is found among the
European red deer, Cervus elaphus (Darling, 1937) and among the
American elk or wapiti, Cervus canadensis (Murie, 1951).
The most complex types of division of labor are exhibited by man
himself and by certain insects. The division of labor in man is a
learned behavior and occurs without structural modification. In com-
plete contrast, the behavior exhibited by members of an insect colony

is instinctive and the individuals
performing various functions in the
colony are specialized morphologically and physiologically (Fig.
9.10). This differentiation of members of the same species is devel-
oped to its
greatest extent among the termites and ants (Wilson,
1953). Since the division of labor in an insect colony is accompanied
by structural specializations and instinctive behavior, it is less flexible

than that occurring in mammalian groups, and especially in man's so-
ciety.
This fact and their size limitations have no doubt largely pre-
vented the insects from attaining a more dominant position in the
world.
Amonggroups of animals all degrees of integration exist from

aggregations with no organization to highly elaborated societies.
Allee (1951) has developed the theory that societies have evolved
from aggregations. If the existence of a group confers definite sur-
vival value on the individuals, the group will tend to persist. Those
reactions that lead to the formation of the group will also persist,
and these will constitute the beginning of social behavior. The co-
operation that we now observe among the individuals may have
resulted as a consequence of the formation of the group. The
division of labor appeared as chance variation and persisted because
of its survival value; it is not necessarily conscious nor purposeful.
According to the foregoing ideas the elaborate social organization

found among a few species may therefore have evolved through the
following steps. First, others of the same species are tolerated in a
restricted space and then definite reactions to their presence are de-
veloped. If survival values result either through behavior or physio-

logical adjustments,
a tendency for an increase in the permanency of
the group will exist. Following this, a further development of the or-
ganization of the group take place with the appearance of lead-

may
the division of labor. It is impossible to
ership, dominance, and/or
say just when an aggregation becomes sufficiently organized to
be
termed a society all
gradations exist. The evidence is strong, how-
ever, that the beneficial effects of increased numbers have played a
considerable part in the development of organized groups and fully
differentiated societies.
FIG. 9.10. The various forms of the termite Kalotermes fiavicollis and their de-
velopmental origins. The eggs (bottom) hatch into young nymphs which after
5 to 7 molts reach the pseudergate stage (individual in center). From this stage
the termite can change into a winged reproductive (top) by way of two wing-
padded nymphs. At intermediate stages environmental influences may cause the

nymphs to change into supplementary reproductives (left) or, by way of soldier
nymphs, into soldiers ( right ) Most of the nymphs do not differentiate and these
.
function as workers. All these stages are present in the termite colony, and their
activities are integrated in its maintenance. (Liischer, 1953, Set. American,draw-
ing by E. Mose. )
Natality and Mortality 333
POPULATION DEVELOPMENT
The organisms inhabiting an area at a given time constitute a
population. the
If
organisms all belong to one species, they form a
single-species population. In most natural situations several to many
kinds of plants and animals coexist in the same habitat. The in-
habitants may then be regarded as composing a corresponding num-
ber of single-species populations that are intermingled, or alternatively,
as forming one mixed or multi-species population. The interspecific
relations of mixed populations will be treated in the next chapter
Here we shall discuss the quantitative relations that arise during the
growth and fluctuation of single-species populations under the in-
fluence of the harmful and beneficial interactions considered in the
foregoing sections.
Principles governing population dynamics are found to apply
equally to special situations, such as laboratory cultures, in which
only one species is present, and to natural areas in which many popu-
lations exist together. The presence of other species acts as part of
the environment in influencing the changes in the population of the
species under consideration.
Natality and Mortality
The abundanceof a species in an area tends to increase because of
reproduction, and to decrease because of death. The rate of repro-

duction depends upon the birth rate or natality, and the rate of death
is referred to as mortality. The dispersion of members of the species
also affects abundance positively in the case of immigration, and
negatively in the case of emigration. We shall consider first the sim-
plest situation in which no dispersion is taking place, leaving a discus-

sion of the influence of migration into or out of the area to a later
section. The maximum possible rate of reproduction for a given
is termed the
species under optimal conditions potential natality. In
natural situations the potential natality is rarely, if ever, attained be-
cause the birth rate is inevitably reduced by one adverse circumstance
or another. The actual birth rate under the existing conditions is
referred to as the realized natality. In parallel fashion the lowest pos-

sible death rate for a given species in the best of circumstances is the
potential mortality, and the actual death rate is the realized mortality.
Natality and mortality vary not only from species to species but
also according to the age of the individuals. Natality is
usually
highest during the middle of the life

span after the individual has
become mature and before it has become senile. Variation of mor-
with age or different kinds of
tality life
stage differs greatly among
animals and plants. In some species extremely high mortality is ex-
perienced in the egg, larval, or seed stages; in others high mortality
does not occur until late in life.
The and animal is greater

potential natality of every species of plant
than potential mortality, and hence, under favorable conditions,
its
every species always has the capacity to increase. If under existing

conditions the realized natality also is greater than the realized mor-
tality,
the population will actually increase. If the two rates are
equal, the population will be stationary; but, if the realized mortality

is
greater, numbers will diminish. A birth-death ratio defined as
ir
100 ..
,
is known as the vital index.
deaths
Biotic Potential and Environmental Resistance. The maximum
possible rate of increase (highest vital index) for a population of a
species occurs under ideal conditions in which the birth rate is the
highest possible for that species (potential natality) and the death
rate is the lowest (potential mortality). Maximum birth rate is de-
termined by the largest number of viable progeny (spores, eggs,
young, or seeds) that an animal or a plant can produce and the
frequency of reproduction. Minimum death rate is determined by
internal factors controlling survival when environmental factors are
all
completely favorable. The values of maximum birth rate and
minimum death rate are thus fixed by life
processes inherent within
the organism, and the maximum rate of population increase, or the
biotic potential, is an innate characteristic of each species. The
value of the biotic potential, or potential increase as it is sometimes
called, differs widely from species to species; contrasting examples of
differing rates were given in Chapter 1. As will be more fully dis-
cussed subsequently, evolutionary processes have established certain
relations between the biotic potential of each species and the exigen-
cies of its existence.
Under natural conditions the full biotic
potential of an animal or a
plant population is
ordinarily not realized since conditions are rarely
completely favorable, Harmful climatic changes, attacks by preda-
tors anddiseases, and other external circumstances curtail the growth
of the population. As the population grows, the increase in numbers
itself
produces changed conditions. A moderate increase in density
may sometimes have an ameliorating effect, as we have seen, but
sooner or later the detrimental effects of overpopulationscarcity of
Biotic Potential and Environmental Resistance 335
food supply or of breeding sites, accumulation of metabolites, and the
like will appear. The combined effect of these factors tending to
curtail population growth is called environmental resistance. The
capacity to increase resides within the species, but the degree to
which it is realized is determined by the environment, including the
changes in the environment brought about by the species itself, and
aspects of the environment consisting of other members of the species.
Thus the actual rate of the increase of a population is determined by
the balance struck between biotic potential and environmental
resistance,
Certain features of the environment are largely or entirely unaf-
fected by changes in the density of the population. These are density-
independent factors. For example, an increase in the abundance of
a species in a marine area does not affect the temperature or the
salinity of
the water, but changes in these factors, harmful or other-
wise, are brought about by agents unrelated to the density of the
population. Other changes in the environment are directly related

to the abundance of the animals or plants concerned; these are
density-dependent factors. Scarcity of such necessities as food, oxy-

gen, or breeding sites may become increasingly
acute as a population
of animals grows; lack of nutrients, excessive pH values, or other
inimical condition may be brought about bythe growth of a plant
population. In addition, the susceptibility of organisms to disease

as well as the ease of transmission is often increased as the density of
the population grows. Generally speaking, the physical features of
the environment tend to be density-independent and the biotic in-
fluences are often density-dependent factors, but the reverse is some-
times true, and certain factors may change from one category to the
other according to circumstances. For example, the predation of a
tawny owl ( Strix aluco ) on a population of mice in an English wood-

land was found to be a density-independent factor since the owl was
observed to eat only 4 to 6 wood mice (Apodemus sylvaticus) each
the size of the prey population (Miller, 1951), But
day, regardless of
in other situations predation will be a density-dependent factor if
each predator kills more prey when the prey are abundant. During
periods of plenty many predators are known to kill more than they can
eat. Increased density of prey may also cause increase in the pred-
ator factor by making possible increased reproduction and growth of
the predator population as well as by attracting predators from neigh-
boring areas.
The question just discussed
of whether the density of a population
controls or modifies a certain environmental factor must be carefullv
distinguished from the question of whether the influence of

the factor
on the population is related to the size of the population. For ex-
ample, the winter temperature in most habitats is a density-independ-
ent factor since its value is not affected by the number of organisms
present, but the effect which the temperature has may or may
not be
related to the size of a population of animals. In some habitats ex-
treme cold will kill the same percentage of a large population as of
a small one, with the result that the number of animals succumbing
is
proportional to abundance. In other instances the habitat will
provide adequate (from extreme temperature or other danger)

shelter
for a small number of animals but inadequate shelter for a large num-
ber. As a consequence a much greater fraction of the large popula-
tion would be harmed than of the small population. If in another
situation a factor killed off a constant number of animals in an area,
it would have a
greater relative effect on a small population than on
a large one.
Form of Population Growth
When plants or animals reproduce, they add more individuals to

the population, and the enlarged breeding stock then has the capacity
to produce a still
larger number of progeny. Thus the population
tends to grow at an ever-accelerating rate. Sooner or later, however,
harmful density-dependent influences begin to take effect. If we
assume for a moment that no other interfering factors are present, the
growth of the population will follow a mathematically prescribed
form.
Let us take a simple numerical example. Suppose we assume that
under the most favorable conditions a certain pair of animals can pro-
duce 6 young during a year and that the resulting population suffers
a mortality of 2 during the year. We then have:
N + A - M = tf ,
S + (5
- 2 = 6
in which N and N, represent the populations, at the beginning and

at theend of the year, respectively, A is augmentation, is
mortality, M
and R is the biotic potential or rate of potential increase per
genera-
tion. The generation time example is one year. At the be-
in this
ginning of the second year our sample population stands at 6, and,

since there are more individuals to breed, a larger number of progeny
Logistic Curve 337
willbe produced during the second year than during the first. As-
suming the same rates of reproduction and mortality, 18 young will
be produced, 6 will die, and the total population at the end of the
second year will be:
Ni + A - M = N*
6+ 18 - 6 = 18
or
RN - N, l
3 X 6 = 18
If the same rate of increase continues
unimpeded, the population will
have grown 54 at the end of the third year, 162 at the end of the
to
fourth year, and so on, as shown in Table 18. The population, which
under these circumstances is exhibiting a geometric or "logarithmic"
increase, is
represented by the equation:
N= #'
where f in years starts at 0, This is indicated graphically by curve
(A) Fig. 9.11. The rate of change of population size is given by the
equation :
dN = A/
N loge R7) ,
^
which means that growth rate is
proportional to size of population.
TABLE 18
GROWTH OF HYPOTHETICAL POPULATIONS IN WHICH INCREASE Is

(1) UNIMPEDED AND (2) SELF-LIMITED
Years
Total population
01234
26 18 54
Unimpeded Increase
102
5
486
6
1458
7
4374
Numbers added 4 12 3(> 108 324 972 291(5
Years
Total population
0123
26 14 27
Self-limited Increase
4 5
39 46 48
6 7
49
Numbers added 4 8 13 12 7 2 1
Logistic Curve. Since the full biotic potential of a species is not

realized under most natural conditions, the population does not in
fact increase as fast as it could if its
growth were completely unim-
peded. However, the natality rate remains above the mortality rate,
if
the population will continue to grow and will increase at an accelerat-
ing rate. Nevertheless, the increase in the population will eventually

produce conditions harmful to itselfdensity-dependent factors will
come into play. The rate of growth will then be progressively cur-
tailed until it reaches zero when the population reaches the largest
size possible for it within the area concerned. If the harmful effect
of crowding increases proportionally, the rate of change of population

size can be expressed as follows:
dN _ (K - N
N log, R
dt V K
where R the hiotic potential per generation, N is the size of the popu-
is
lation atany moment, and K is the ultimate maximal size possible for
the population in the given area. In words, the equation simply says
that the rate of increase of the population is
equal to the potential
80
70
60
Asymptote
50
40
30
20 LA "-Inflection point
10
20
10
1234 Years
FIG. Upper figure: graphs showing theoretical increase in total numbers

9.11.
during (A) unimpeded and (B) self-limited population growth (logistic curve).
Lower figure: graph of numbers added to population in each time interval during
the logistic growth shown by (B) above (values plotted at midpoint Of each time
interval).
*
For further discussion of these relations reference may be made to H. C.
Andrewartha and L. C. Birch. "The Distribution and Abundance of Animals"
1954, Univ, of Chicago Press.
Logistic Curve 339
increase limited by the degree of realization of maximal size. When

the growth of a self-limiting population is
expressed graphically, a typi-
cal S-shaped curve is obtained known as the logistic curve, shown as
(B) in Fig. 9.11. The curve is

truly logarithmic only at its
begin-
ning, departs from the logarithmic increase as impeding factors be-
come effective, reaches an inflection point at which the acceleration of
growth becomes negative, and approaches an asymptote representing
the limiting size (K) of the population.
If we now
apply to our previous numerical example the more real-
isticcondition that increase is progressively curtailed as the popula-
tion grows, we obtain the growth values, rounded to integers, indicated
in the lower part of Table 18 with a maximal population size in this
case assumed as 50. Besides the population totals at the end of each
year (or other period) we are also interested in knowing the numbers
added to the population during each unit of time. With unim-
peded growth the annual increment becomes larger and larger in-
definitely; but with self-limited growth the annual increment passes
through a maximum at the time when the logistic curve reaches
the
inflection point. In our example the numbers added per year in-
crease to a maximum of 13 during the third year and then drop off
nearly to zero in the seventh year. A curve showing these changes

in
the increments to the population and their relation to the logistic curve
presented in the lower portion of Fig. 9.11. The fact that

is the
population has approached its asymptote with annual increment ap-
proaching zero does not necessarily mean that little or no reproduction
is
taking place in the populationit simply means that births are com-
pletely offset by deaths, natality is equaled by mortality, or A =M .
Under these circumstances reproduction may continue to be high ac-

companied by high mortality, or reproduction may be low with low
mortality; as long as A and M are equal the size of the population will
not change.
Early in the history of the population only small numbers are added
each year because the breeding stock is small. During the middle
period annual increments are large, but, as the population reaches its
maximum size, small annual increments again occur either because
breeding is
sharply curtailed or because the young produced suffer
severe mortality. The reader should particularly notice that the
largest annual increment is not found when the population is at its

maximum, but occurs at the inflection point of the logistic curve, that
is, at the time when
the population is growing most rapidly.
Populations of awide variety of organisms, ranging from bacteria

to whales, have been found to follow the logistic curve in their growth
form. Illustrations of such curves of growth for laboratory cultures

of Protozoa, yeasts, Drosophila, flour beetles, and water fleas, and for
natural populations of bees, ants, thrips, sheep, and other animals are
discussed in further detail by Allee et al. ( 1949, Ch. 21 ). The growth
of man's population follows a similar pattern whether examined in
individual regions or in the world as a whole. A plot of the census
records for the United States for the years up to 1940 is shown in Fig.
9.12, and the curve has been extrapolated by fitting
the logistic func-
Year
FIG. 9.12, Growth curve of the population of the United States, showing census
counts from 1790 to 1950. The logistic function has been fitted to the counts
from 1790 to 1910 and extrapolated to 2100. The agreement of the extrapolation
with the counts for 1920 to 1950 is shown, and a cessation of growth about the
year 2100 is indicated. (Modified from Pearl, Heed, and Kish, 1940.)
tion. The agreement of the 1950 census figure of 151 million with
the extrapolated curve and the indication of an asymptote at about
184 million in the year 2100 may be observed.
Equilibrium and Fluctuation

The logistic
curve discussed in the previous section applies only to
periods of population growth (when A>M)

and to situations in
which the rate of increase is controlled only by density-dependent
factors. Since the inhabitants of a natural area have mostly been
present for a long time, we see the initial stages of population growth
only in special instances.

The early part of population increase is
observed when a bare area is invaded, when a new species is intro-
duced, or when a check holding a species to a low number is sud-
denly removed. Aside from these special situations the changes ob-
served in population size involve the action of both density-dependent
and density-independent factors usually long after the original growth
of the population. In both laboratory cultures and in natural habi-
tats, after the initial attainment of maximal size, a population will
(1) maintain itself at about the same level for a long period, (2)
decline and eventually become extinct, or (3) fluctuate regularly or
irregularly.
the population approaches its asymptote in such a way that the
If
supply of food, and other necessities, and the removal of harmful by-
products keeps pace with growth, then the population will maintain
itself ator near this equilibrium level (A M) until outside condi-
tions are altered. Under these circumstances the reproductive rate
may be high or low, but, as long as it is
exactly offset by mortality,
the population size will not change. Nevertheless the magnitude of
A and M exert an important ecological effect on evolution.
may If
A is
very large, as is true of
many invertebrate animals, the accom-
panying high mortality will generally take place when the progeny
are young and usually before reproduction has occurred. On the
other hand, if A and M are small, a much larger proportion of the
young animals may live long enough to reproduce before they are
eliminated from the population. In the former circumstance, most
of the mutations appearing in the population will be lost, but in the
latter case a larger percentage will be retained long enough to affect
the next generation. Wefind that the possible effect of this difference
corresponds to the generally more rapid evolution of mammals, for

example, than that of invertebrates.
In other populations the harmful conditions that were produced by
increasing numbers and that brought the growth of the population to

a stop may progressively intensify. Under these circumstances the
greater and greater scarcity of food, the accumulation of metabolites,
or other inimical change will cause a decline in the population either
immediately or after a period of time. If the changes in the environ-

ment brought about by overcrowding are irreversible, extinction of
the population will eventually follow. This result is commonly seen
in laboratory cultures of bacteria, Protozoa, and other organisms. In
nature a similar fate may overtake a population developing on a small
island as a result of the introduction of breeding stock by natural
processes of dispersal or by man. Rats, goats, rabbits,

or other ani-
mals, escaped from explorers' ships or introduced by colonists, often

find conditions suitable for rapid growth and reproduction. Unim-

peded, geometric increase in the population ensues until all suitable
food or other necessity on the island is exhausted. Thereupon starva-

tion, and often disease, decimate the population, and the death of all,
or nearly all, of the animals quickly follows.
More usual than either of the foregoing is the third type of situa-
tion in which the population overshoots its equilibrium level, but,
after a reduction in numbers, conditions are ameliorated sufficiently
for the population to increase again. The repetition of this process
causes fluctuations. If these are small, the population curve is desig-
nated as flat; if the amplitude is large and regular, the curve is termed
cyclic, and,
if
irregular, especially with sudden periods of great in-
crease, it is called irruptive (Fig. 9.13). Some ecologists refer to
Irruptive
Cyclic
* '>
Extinction
Time
FIG. 9.13. Diagram of types of fluctuations in populations.
regular changes in abundance as oscillations and to irregular changes

as fluctuations, but the more specific terminology given above will be
employed here. The reader should understand that all gradations ex-
istbetween these types of population change.
Fluctuations after a population has approached its equilibrium level
may be caused in whole or in part by changes in physical features of

the environment or in biotic influences such as the abundance of
predators, diseases, or food organisms. These population changes

due to the interaction of different species will be considered in detail
in later
chapters. At this time we wish to emphasize the fact that
conditions within the population of a species can themselves be re-
sponsible to a greater or lesser degree for the fluctuation in addition

to any
disturbing influences that may come from the outside, The
causes of certain fluctuations can be traced to the
reciprocating ef-
fects of
natality and mortality within the population itself.
A neat illustration of the mechanisms involved in self-induced fluc-
tuations of this sort is found in
experiments with laboratory cultures
of Daphnia magna (Pratt, 1943). Single females were placed in
bottles containing 40 ml of water, and as
they and their progeny re-
produced parthenogenetically the population in each bottle grew
in accordance with the
logistic function. The culture medium was
renewed each day, insuring ample food supply and removal of me-
tabolites, and other external conditions were kept uniform. Follow-
ing the initial attainment of maximum size, the population sank to a
low ebb and then continued to fluctuate in extreme fashion in spite of
constant environmental conditions (Fig. 9.14). The explanation is
100
I 80 A
o 60
I
E 40
A
20
10 20 30 40 50 60 70 80
Time in days
FIG. 9.14. Changes in the abundance (solid line) of Daphnia magna in a labora-
tory culture. The variations in number of births (--)

and of deaths ( )
that underlie the fluctuations of the population are indicated. (Pratt, 1943.)
seen in the changes in the birth and death curves indicated in the
diagram. Although the number of births declined after the tenth

day, young Daphnia continued to be produced for 18 days with the
result that more young were added to the population than could sur-
vive when they reached adult condition. As overcrowding continued,
the death rate rose rapidly and the birth rate dropped to zero. Even
after the size of the population was considerably reduced, no repro-
duction took place at first and in fact no new young animals appeared
until the fortieth day. Animals born in adversity thus did not pro-
duce young immediately upon reaching maturity, and the reduction
in their reproductive capacity persisted for a long period after fav-
orable conditions had been restored. As a consequence the popula-
tion continued to overshoot and undershoot a possible equilibrium

level. When experiment was repeated at a lower temperature,
this
the population changed more result that after one or
slowly, with the
two oscillations an equilibrium value was reached and maintained.
Delay in the manifestation of excessive natality and prolongation of
the effects of overcrowding similarly act on natural populations and
add their influence to that of external factors in causing fluctuations.
The relative abundance of individuals of various ages in the popu-
lation known as the age distribution of the population. Differences
is
in age distribution depend upon the species and whether the popula-
tion is changing in size or stationary (Petrides, 1950). rapidly A
growing population usually contains an especially large number of
young individuals, whereas a declining population includes a rela-
tively high proportion of old individuals. In a stationary population
the distribution of ages is more uniform and tends to approach a
stable pattern. Since natality and mortality vary with the age of the
individuals, the age distribution of the population influences the birth
and death rates for the population as a whole. Species that produce
large numbers of young generally suffer high mortality during the
young stages. A complete description of the mortality of a popula-
tion furnished by a life table such as has been constructed for man
is
and has long been used by life insurance companies. Life tables and
survivorship curves have now been worked out for a number of
natural populations and show characteristic differences in mortality
patterns among various species (Deevey, 1947 and 1950). For ex-
ample, mortality in the oyster is extremely high during the larval stage
and becomes much lower later in life, in hydra mortality is nearly
constant at allandages, in man mortality tends to be low for a long
period during youth and to become high rather abruptly in old age.
Optimal Yield
The foregoing analysis of population development is of interest
not only in relation to theoretical considerations but also in connec-
tion with practical applications. In the exploitation of natural
popu-
lationseither plant or animal a harvest is desired of the largest
number of individuals per unit of time that is possible without per-
manently impairing the breeding stock. In other words, we wish to as-
certain the size of the largest sustained yield that can be obtained and
learn at what level the population should be maintained to
produce
this yield. An answer to this question the formation of the
is
theory
of the optimal yield. The basic idea underlying this
theory is that in
Optimal yield 345
the simplest situation with no modifying conditions the population

should be maintained at the inflection point of its growth curve since
at that point the largest increment is
being added per unit of time
and the taking of this increment would represent the largest harvest
possible without inroad upon the breeding stock. We shall return
later to a discussion of special circumstances applying to particular
species that modify this basic relation and cause a different density
level of the population to be more favorable for the practical exploita-
tion of these species.
To illustrate the basic principle let us imagine a flock of geese from
which we wish to obtain as
many birds as possible for food. Suppose
that the flock originated from the establishment of one pair in a lim-
ited area and that the growth of the population followed the logistic
curve (Fig. 9.11) with values similar to those indicated in the lower
part of Table 18. The largest single yield would obviously be ob-
tained by allowing the flock to grow for 5 or 6 years and then shoot-
ing all or most of the birds. For a sustained harvest, however, only
the annual increment to the population should be taken; in this ex-
ample, 13 birds could be taken every year if the flock were allowed
to grow to 27 before harvesting. If a hunter considered exploiting
this flock without knowing its
existing si/e and found that only 7 or 8
birds were added to the population each year, he would have to de-
termine whether the low growth rate was due to the population being
smaller or larger than the optimum. Low yield in any such popula-
tion thus be caused by overexploitation resulting in too small a
may
breeding stock, or in underexpolitation resulting in harmful crowding
of the breeding stock. The best conservation procedure often calls
for the reduction in size of a natural population. Recognizing this
fact,rangers now
regularly cut back the populations of deer in national
parks whenever numbers become too large because of the absence of
wolves, panthers, or other natural enemies that otherwise would keep
the deer in check. Consequently, in exploiting a population, it is de-
sirable to ascertain whether density is above or below the level at
which increase is most rapid and, if possible, to adjust the size of the
population accordingly.
As already suggested, several important modifying considerations
must be added to the underlying idea of the theory of optimal yield.
Up to this point we have discussed populations in terms of numbers
of individuals. Fishermen do not measure their catch by counting
the but by weighing them; foresters evaluate their timber by its
fish,
volume, not by the number of trees. Thus, the growth in size of the
individuals must be taken into account as well as the increase in num-
bers. For such species the growth of the population is measured in

terms of weight or volume, and, when
plotted in
corresponding units,
again follows a logistic curve, although the numerical values will, of

course, be different. Accordingly, in attempting to ascertain the
optimal tonnage of fish to catch we must consider the growth rate of
the population in terms of increase in weight rather than numbers.
If there are no further modifying considerations in respect to a par-
ticular fishery, the optimal catch may be obtained when the popula-
tion is maintained at the level at which the largest weight increment
will be added per unit of time, this increment being due both to an
increase in numbers and to the growth of the individuals (Fig. 9.15).
Other practical considerations are the availability of the population
for exploitation and its suitability for the market. If game in a forest
or fish in a pond are scarce, more time, effort, and expense will be re-
quired to harvest the same number than if the population were dense.
With sparse populations the yield per unit effort is lower, In specific
instances, it
may be
better to allow the population to increase to a
somewhat greater density than that which represents the level of most
rapid growth in order to increase availability. Often minimum limits
and sometimes maximum limits of size exist in relation to market-
ability. Accordingly, the size composition of the catch must be con-
sidered as well as its weight or volume.
Two additional considerations that also may modify the basic plan
for obtaining the optimal yield in respect to certain species are: the
number of breeders necessary to provide sufficient young, and the
rate of natural mortality. In species with a low natality such as most
large mammals and birds, the recruitment, or number of

young an-
nually added to the population, is
closely dependent upon the size
of the breeding stock. For each new young whale there must be one
adult female; but one fish, one oyster, or one termite can produce mil-
The spat released by only a
lions or billions of young in a season.
few clams would provide an ample set for a large area, and, if al-
lowed to grow up, the young produced in one year could repopulate
a whole bay under completely favorable conditions. In such species
the recruitment usually depends more on the environmental condi-
tions during the early life of the new generation than on the number
of breeders, and the optimal level at which the population should be
maintained is determined by other considerations.
We have seen that allowing populations to build up by avoiding
overexploitation sometimes has advantages in the greater size of in-
dividuals, the more concentrated populations, and the larger number
of breeders. On the other hand, allowing too great development of
Optimal Yield 347
1 1
C+-4O
o __
03 o 3
a
i
S
.
2
!
!^
!fr
0-3
the population will result in the disadvantages of overcrowding.

Another consideration is the fact that the longer an animal or a plant
remains unharvested, the longer it will be exposed to environmental
die from other
dangers and the greater the possibility that it will
causes or be removed by predators other than man. If natural mor-
tality is very high, the crop may

have to be harvested at an earlier
age than would be necessary should the reverse be true.
The optimal
level for the population thus varies widely according to the ecological
relations of each species. The optimal yield is obtained when the
rates of reproduction, growth,and mortality are so adjusted as to pro-
duce the greatest annual increment with due regard to availability and
other modifying considerations. More detailed discussion of both
the theoretical and the practical aspects of this important topic is
available in Russell (1942), Trippensee (1954), and the publications
of the United States Department of Agriculture.
SPATIAL RELATIONS OF POPULATIONS
Space Requirements
As a population grows in numerical strength and as its members in-

crease in size, the individuals tend to come closer and closer together
as long as the population occupies the same area. In many instances
the habitable area sharply limited, as it is for aquatic populations
is
developing in a pond, or for land organisms multiplying on an island.

Although in other instances populations can spread out at their mar-
gins for a time as they develop, eventually the unoccupied area is
completely taken up and a condition of overcrowding begins to ap-
pear.
Every organism requires a minimum amount of space within which
it can carry on its necessary exchange with the external world. This
minimum space must be sufficient to provide food and other necessi-
ties, to absorb metabolites, and to permit reproduction. A relation
obviously exists between the size of the organism and the minimum
size of the space that it can inhabit. Whales do not live in ponds.
The space required by some organisms is quite small and may be
no larger, in one plane, than the actual dimensions of the body. The
number of adult barnacles that can inhabit a rock, for
example, is
determined simply by the number of barnacle bases for which the
area of the rock affords room. Barnacles can grow with their sides
closely appressed
to their because water currents
neighbors bring
food particles and oxygen and take away metabolites and progeny.
Space Requirements 349
Most species require more space than their physical dimensions.

The actual amount of room needed varies
greatly from species
to
species, but frequently the space required is very large either because
some necessity is sparse in its distribution or because of certain, often
poorly understood, psychological relations. A census of the fishes in-
habiting the shore zone of Morris Cove, New Haven, Conn, revealed
the presence of representatives of 32 species, but few specimens
longer than 100 mm

were found. Although the adults of small species
were taken, only the young of the large species were present. The
depth of water was ample for more sizable fish to move about in, but
fish larger than the critical size were
definitely excluded from this
habitat regardless of age although the mode of action of the space
requirement in this instance is not known (Warfel and Merriman,

1944).
The manifestations of the space requirement may also occur in re-
lation to breeding, hostility, or other reaction. Insects such as the
Canna Leaf Roller (Calpodes ethlius) have been found to breed sat-
isfactorily under laboratory conditions if placed in containers of cer-

tain size. They will not breed in smaller containers, nor in much
larger confined areas such as greenhouses, which appear to be other-
wise suitable. Cannibalism among insects often occurs when they
are crowded together, whereas members of the same species do not
attack each other when they meet at other times under uncrowded
conditions. The English sparrow, a partially domesticated bird, will
breed regularly in large aviaries but will not breed in small cages.
The foregoing examples suffice to illustrate the point that definite
space requirements exist not only in relation to the physical needs of

the organism but also in relation to reactions essential in their life
cycles.
For such as the majority of plants and non-motile
sessile forms,
animals, the provision of minimum space is automatic to some extent.

As crowding continues to increase, competition brings about the
stunting and eventually the death of a portion of the population.
This process can be readily seen in the development of a stand of trees
from a bed of seedlings. Root competition, shading, and diseases
gradually thin out the population. Young white pines in New Eng-
land, for example, sprout from seeds to form a bed with a density of
perhaps 75,000 seedlings per hectare (30,000 per acre). When these
of 60 to 80 years, they will form a mature
pines have attained an age
stand in which the density has been reduced by natural processes to
about 750 trees per hectare (300 per acre), or even less, without any
treatment by the forester. Similar examples may be found among
oysters, mussels,and other sessile animals growing on substrata of

limited dimensions; a few individuals, growing faster than the rest,
This
gain the upper hand and smother their more retarded
brothers.
automatic reduction of the population by the growth of its own mem-
bers produces at least the minimum spacing necessary for survival,
but it usually does not provide the most favorable spacing for the best
development of the individuals. Man
can often improve growing
conditions for particular species in which he has a special interest by
further thinning and this is done regularly, of course, by professional
growers, such as oystermen, farmers, and foresters.
For motile animals the danger exists that during favorable condi-
tions the members of a species inhabiting a region will move too close
together, with the result that wholesale destruction will take place
when conditions become unfavorable again. In some species we find
an instinctive reaction causing the animals to space themselves out to
a certain extent at least; and the existence of this response helps to
avoid overcrowding, as will be discussed in the ensuing section.
Home Range and Territory
Many kinds of animals are known to establish a center of opera-

tions for themselves and to confine their roamings within certain
boundaries. The area within which an animal tends to stay is known
as its home range. Sometimes other members of the same species
are not allowed to enter the area, or a portion of it, and
any trespassers
are forcibly driven out. The inner sanctum that is actively defended
called the animal's territory. The territory may be established
is
by a
social group, such as a hive of bees or a colony of ants, by a
breeding
pair, or sometimes by a single individual. The existence of territories
with "exclusive rights" was recognized in an old Chinese proverb
that states: "One hill cannot shelter two tigers." A dog's own
yard
is his
territory,
and he drives other dogs out with a self-assurance that
is familiar to all of us. Within their own territories animals
typically
acquire a heightened position of dominance in relation to other mem-
bers of the same species.
The concept of territory was first delineated for birds. It was ob-
served that in many species each male established a territory at the
beginning of the breeding season. The male then isolates himself,
and confines his movements to his own territory He becomes hostile
to other males of the same species, with which he was on amicable
terms only a short time previously. This bird drives others of his kind
out of his territory, but pays little or no attention to birds of other
Home Range and Territory 351
species that wander by unless they actually intrude on his intended
nesting site.
Characteristically, the male bird chooses a singing
perch, makes himself conspicuous, and, when a potential mate comes
to the area, goes
through the courtship display. In some species, as
for example the house wren, the male
may even start building a nest
before the female arrives.(No doubt he has to do it all over again
when she appears on the scene!) After the nest is completed and
during the period when eggs or young are in the nest, the territory
is defended with particular fierceness.
carefully observing the movements of a pair of birds for a

By
period of time, the ecologist can determine the position of the unseen
boundary lines of their territory. If the birds build a second or a
third nest during the same season, these later nests are placed some-
where within the original territory. The division of a region into
territories by pairs of breeding birds is illustrated in the diagram of
Figure 9.16. It will be noted that the territories of birds of the same
species do not overlap indeed, a neutral zone usually exists between

them but the territory of one species may overlap that of another
species if no reaction of antagonism occurs between them.
In a 16-hectare tract of spruce-fir forest in northern Maine 148 male
birds of various species were found to have established breeding ter-
ritories by a census conducted during a week in June. During the
ensuing 3 weeks a total of 302 territorial males were removed from
the tract by shooting. Study of the area indicated that new males
from a "floating" unmated population moved and established ter-
in
ritories as rapidly as the former inhabitants were removed. Thus
thenumber of sites suitable for the establishment of territories was
shown to be the chief factor limiting the number of birds that could
breed in the area (Stewart and Aldrich, 1951).
The dimensions of the territory or home range may be measured
in kilometers in the case of large mammals or birds of prey or in meters
or smaller units in the case of smaller animals. Bears may roam over
distances of 30 or 40 km, whereas porcupines confine their home
activities to trips of a kilometer or so and the home range of beavers
is
ordinarily less than a kilometer in diameter. The home ranges of
the whitetail deer vary in size from about 80 to 120 hectares ( 200 to
300 acres), of the cottontail rabbit from 1 to 3 hectares, and of some
kinds of mice from less than %
to more than 1 hectare. The daily
roamings of prairie deermice in southern Michigan were found to be
confined within 2 or 2% hectares, but on different occasions during a
breeding season both male and female deermice may travel over an
area as great as 4 hectares (Howard, 1949). The home ranges of box
Homing 353
turtles in Paxtuxent Research Refuge, Maryland, investigated by

Stickel (1950), were found to average 100 m
in diameter and to over-
lap considerably. Chameleon lizards were reported to establish ter-

ritories of 20 to 25
sq m
in area. The arrangement of territories of
the wood ant
illustrated in Fig. 9.17.
is In this instance the ants*
nests were found to be only a meter or so apart. The ants foraged
for distances of 70 m
or so, but in no instance did the members of
one colony encroach upon the territory of another colony. In col-
onies of breeding birds the private domain around each nest site may
be very small indeed, but this may be jealously guarded (Fig. 9.18).
Wood mice apparently do not establish territories that are actively
defended, but home ranges are delineated. Some avoidance response
keeps the animals confined to their home ranges. The reality of this
action of population pressure has been demonstrated by trapping ex-
periments. When an area cleared of one species thus creating a

is
"social vacuum" animals from neighboring areas will move into the
vacant area within a few days. Obviously then, the animals on the
periphery of the area in question did not fail to enter the area because
it lacked food or other
necessity, or because they lacked the inclina-
tion to move the distances involved. The neighboring mice were
kept out of the area by "social" pressure of some sort resulting from
the presence of other members of the same species. Although animals
avoid entering the territory of others, peripheral contacts with neigh-
bors are presumably a normal condition. The rapid entry of the mice
into a cleared area may be'clue in part to a movement of the animals
"as if an attempt to encounter again the stimuli produced by
in
neighbors" (Calhoun and Webb, 1953).

The size of the territory or home range is usually larger than that
which would be needed for the food supply of the individual estab-
lishing it. In the case of territories staked out at the beginning of

the breeding season, it is obvious that the food supply must be ade-
quate for the young that are to be raised. Apparently during the
course of evolution reactions for establishing the territory have come
into existence that cause the breeding pairs to space themselves out
to anticipate the food needed by the new family. The
sufficiently
size of thehome range or territory has probably come to be based
primarily on the food supply, but in some instances it may be de-
termined by available breeding sites or other needs of the species.
Homing. Frequently the attraction of an animal to the location of
its home is very strong; this is particularly true if the territory is
established in relation to a breeding site, and the reaction is further
intensified if
young are present. The return of an animal to its ter-
10 20 30m
FIG. 9.17. Wood-ant trackways and nests at Picket Hill, Oxfordshire, England.
= nests, o = former nests, B = birch trees, G = gorse bushes, x = nests of wil-
low wrens. ( Elton, 1932, Copyright, Cambridge Univ. Press. )
Photo R. H. Heck, Am. Museum of Nat. Hist.

FIG. 9.18. Territorial dispute in penguin colony, Bleaker Island, Falklands.
Return Migration 355
ritory or breeding site is referred to as homing. The reaction is

par-
ticularly familiar among birds, but other types of vertebrates, many
species of insects, certain Crustacea, and some other invertebrates are
known to home. Even the pulmonate snail Onchidium, after leaving
its home crevice in the rocks to feed in the
surrounding area, returns
to its own crevice and cannot be induced to enter any other
particular
(Arey and Crozier, 1921). Bees fly unerringly back to the hive from
distances greater than 6 kilometers.
Homing is especially well developed among birds. The distances

covered are often spectacular, and the precision with which the home
territory is located is little
Although small perch-
short of miraculous.
ing birds may travel only 100 or so to mprocure food, hawks and
eagles forage for many kilometers, and sea birds often range widely
over the ocean before returning to the home site. When investigators
have carried parent birds great distances away from their nests and
released them in unfamiliar surroundings, the prompt return of the
birds to their breeding sites has demonstrated the extraordinary
strength of the homing reaction as well as the existence of some highly

successful method of orientation. The fact that the performance of
domesticated homing pigeons improves with training and familiarity
with the terrain indicates that these birds use landmarks to some
extent, but other methods of orientation must also be employed
(Matthews, 1950).
In nature birds will home across unfamiliar country and over water
where no landmarks exist. Perhaps the most spectacular instance of
homing is that of a Manx shearwater that was taken from its nesting
burrow on the Island of Skokholm, Wales, banded for identification,
and transported across the Atlantic by airplane. This bird was re-
leased from the Logan International Airport, Boston, Mass., on June
3, 1952, and arrived back at its burrow in Wales 13 days later, having
traversed 3000 miles of ocean. We do not know whether birds and
other animals use the same methods of orientation for homing as they
do for seasonal migrations. The whole fascinating subject awaits
further study. A general discussion of orientation was presented in
Chapter 6.
Return Migration. Many birds, mammals, fishes, and other animals
move seasonally from one habitat to another, sometimes traversing
this sort in which at least some of the
great distances. Journeys of
population completes a round trip are known as return migrations.
The most spectacular movements of this sort are found among the
vertebrates, but return migrations also occur in the invertebrates, as
exemplified by certain crabs

and by butterflies (Williams et al, 1942).
the instinct to return to a certain area is exceed-

Among many species
fish
ingly strong, as will be realized, for example, by anyone observing
a river. to an colonial in New
migrating up According early report
England: "Alewifes continue to move up the stream, yea though ye
beat at them with clubs." Perhaps some of these migrations may be
thought of as movements out from and back to a

home territory and
hence an extension of the homing reaction. Return migrations as
or
adaptations to (1) climatic conditions, (2) feeding needs, (3)
breeding activity, have been discussed in previous chapters.
In some of these migrations members of the population come back
to the same general area or even to the same locality. Birds return
to the same region after journeys of several hundred or several thou-
sand miles and after absences of 9 months or more. Frequently a
pair of banded birds uses the identical nest site for two or more sum-
mers although the birds have journeyed to the tropics
in succession
during the intervening winters. The greater shearwater roams widely

over the North and South Atlantic Oceans during most of the year,
but its
only known breeding place is on the Island of Tristan da Cunha
about 2000 miles south of the equator (Murphy, 1936). In some
similarly mysterious way slender-billed shearwaters ("mutton birds")
from all over the Pacific Ocean locate two tiny islands between Aus-
tralia and Tasmania and descend on the islands on the same day
all
in a huge flock to begin the breeding season (Griscom, 1945).

Alaskan fur seals breed only on the Pribilof Islands in the Bering
and they manage to locate in some unknown way these tiny
Straits,
rock in the vast ocean after swimming for a 1000 miles or more
bits of
from regions off the coast of California where they spend the winter.
In some of these breeding migrations the reaction to the home ter-
ritory may persist over the winter and stimulate the animal to return
to the breeding area at the next season. It is difficult to believe, how-
ever, that after several years in inland streams adult eels retain any
homing reaction to the area in the Sargasso Sea where they were
hatched yet they do return there. The same might be said of the
Pacific salmon that ascend the rivers to spawn after spending 4 to 7
years in the ocean, but something does stimulate the salmon to under-
take the migration and something orients the fish to the very tributary
in which
they developed as larvae (Hasler, 1954). Perhaps migra-
tions taking place after long intervals and also those correlated with
climatic changes and feeding activities are unrelated to homing or only
secondarily related to it. In these instances, external or internal fac-

tors may stimulate the animal to migrate at a certain time, and condi-
tions along the way may orient the animal back to the region of its
Emigration 357
origin with the retention of no specific horning reaction or of one that

is
operative only after the animal has arrived within the general area
of its
original home territory.
Emigration
When animals leave their home range never to return, the move-
ment spoken of as emigration. This type of migration may take
is
place either as a drift of individuals or of small groups, or as a mass

movement. The young of many species of birds and also to some
extent the adults wander in all directions after the breeding season
and before the autumnal migration to the winter range begins. An
example of this centrifugal movement is obtained from the returns of
banded ducks (Fig. 9.19). Drift emigration of this sort is very wide-
spread in its occurrence, but it often escapes notice because the
animals or birds move inconspicuously in small numbers. Some kinds
of animals wander about for long periods of their lives, seeking pri-
marily perhaps food or water, without definite return to any given

place; such irregular roaming is spoken of as nomadism. This and
other types of animal travels are discussed fully by Heape (1932).
In contrast to the foregoing, mass emigration is often spectacular
although it occurs in relatively few species. At more or less regu-
larly recurring intervals such animals as the mouselike lemmings of

Scandinavia and Canada, the springbucks of Africa, and the locusts
of Egypt and India undertake mass movements involving millions
or billions of individuals. The causes of these mass emigrations are
not clearly understood: in some instances they may be due to climatic
changes; in others they may be due to lack of food which may or may
not be the result of overproduction of the species itself. The onset of
acute hunger, nervous disorder, or other conditions occurring at the
time of mass emigration often profoundly alter the behavior of the
animals. For example, according to Heape (1932), some such in-
ternal change "transforms the shy and timid springbuck into a fearless
creature with no regard for danger of any kind, and converts the no
less
shy and timid lemming into a truculent swashbuckler."
An instance of a mass emigration of lemmings was witnessed in
1937 by the manager at the Perry River post of the Hudson's Bay
Company, Northwest Territory, Canada, and an excerpt from his ac-
count follows ( Gavin, 1945 ) :
at the old post site on April 27 and for about four

While camped
days subsequently, the onset of the migration was noticed. The whole
tundra was a mass of moving lemmings and each time we went into the tent
there would be a dozen or more inside. The migration went on night and
Lincoln, 1933
FIG. 9.19. Radial emigration of ducks after the breeding season, plotted from
reports of banded birds.
Emigration 359
day; my 13 dogs outside the tent could be heard killing them at frequent
intervals during the night.
When we started for thenew post, the sea ice was covered with a moving
mass of lemmings, all headed in an easterly direction. They stopped at
nothing. Untold thousands plunged over the ice into the water of a lead,
about a foot below, and swam the 10 or 15-foot channel, but were unable to
climb up the sheet ice on the other side. They perished in large numbers
in these leads, but here and there,
they found passages up tne ice and
blindly continued their journey. Around the ends of the leads they pressed
on without interruption going the pools of water, lying on the surface
through
of the ice without deviation and without the slightest hesitation. This scene
extended as far as we could see in any direction. The natives later in-
formed me that at Kol-gyuak-a river 45 miles east of Perry the same thing
was going on at the same time. I do not know how far west it extended,
but it was the same for at least 15 miles west of the post. This mass
migration lasted for about 10 days and reached its peak about May 3 or 4.
In travelling from the old post to the new, my dogs grabbed up and ate
so many lemmings while they were running that their stomachs distended
to a noticeable degree.
They were so surfeited that they were useless for
further work until they had gotten over their abnormal feeding. I had to
rest them for 24 hours to allow them to get over their
gluttony. An esti-
mate of the average density of lemmings during this migration would be
one to the square yard.
Studies of the migratory locust have shown that under

ordinary
conditions this insect exists exclusively in a form known as the soli-
tary phase. Particular conditions of temperature or dryness cause a
great increase in the numbers of the species, and this is followed by

the development in the population of the
swarming phase of the
locust, differing in color and in structure from the solitary phase.
With the appearance of the swarming phase, emigration of great num-
bers of locusts begins (Fig. 9.20). Huge swarms of locusts, such
as were recorded in Biblical times, spread into surrounding areas
and do untold damage to crops and other vegetation. Since the
newly invaded regions are less favorable for the maintenance of the
species, fewer of the young survive, and numbers are also reduced
by enemies that have been attracted by the plentiful food. As the
density of the locust population diminishes, the swarming phase dis-
appears and the solitary phase reappears. Gradually the locust dies
out of the invaded areas, and the species shrinks back to its
original
range (Uvarov, 1931).
In this chapter we have considered the wide variety of relations
among members of the same species. Other individuals of the same
kind become abundant in an organism's environment as a result of
reproduction, of passive transport, or of active locomotion. We have

seen that, as numbers of a species increase, eventually harmful effects
Photos I). L. (i/ww. refixxIiHtd h\ firnnission of the Anti-Locust Research Centre
FIG. 9.20. (Upper) A small swarm of migrating desert locusts in Kenya (density
estimated at 14 insects per cu m). (Lower) Desert locust hoppers (fifth instar)
marching.
Emigration 361
of competition, interference, or aggression always come into being.
However, a moderate amount of crowding may be beneficial to
members of a group, and this result may have favored the evolution-
ary development of integrated social organization in animals. The

fact that an increase in a population progressively curtails its own
growth makes possible a mathematical formulation of population de-

velopment and provides a basis for deducing the optimal yield that
can be obtained by exploiting a population. Numbers of a species
always fluctuate to a greater or a lesser extent. Every species has
definite space requirements, determined either mechanically or by
avoiding reactions related to home ranges, and overpopulation is re-
lieved by the destruction of the extra individuals or by emigration.
The presence of others of the same species thus forms a critical aspect
of the environment, and each organism is involved in many reciprocal
relations with individuals of its own kind.
/o
Relations
between Species
In natural situations the presence of other organisms of different

species is an unavoidable and also a necessary part of the environment.
The existence of other species may be crucially important in the pro-
vision of food, shelter, or some other necessity. Contrarywise, vari-
ous kinds of animals and plants are undesirable neighbors; but the
presence of these species must nevertheless be dealt with as an in-

fluence received from the surroundings. Certain interactions among
species in an area are prominent and clearly discerned, but others
are of a subtle nature not easily studied. Some of the relationships
form an integral part of the operation of the ecological complex as a
whole, whereas others may be of only minor consequence. In this
chapter we shall continue our analytical approach to ecological prob-

lems by surveying the various types of relationships between species
and considering the operation of each separately. In subsequent
chapters the combined operation of these interactions will be con-
sidered in relation to the composition and adjustment of the commun-
ity and to the functioning
of the ecosystem as a whole.
In attempting to delineate the various types of interrelations be-
tween species one realizes that a great complexity exists. Animals
have relations with other animals, plants with other plants, many
animals are dependent upon plants in their environment, and some
plants are dependent upon animals. All gradations exist from rela-
tionships that are vital and lifelong to those that are casual and
tempo-
rary. Interdependency may exist between species of widely different
kinds and sizes as between mighty redwood trees and microscopic
bacteria, or in the animal kingdom between elephants and fleas. In
some instances one species has an exclusive relation with another
sometimes with one short life stage of the other species; but in other
instances species are quite flexible in their dependencies upon their
neighbors.
362
Relations between Species 363
Interrelations between species may be beneficial to both parties,
harmful to both parties, or beneficial or harmful to one and neutral
in respect to the other. Every gradation may be found between these
conditions. The beneficial effect of the presence of another species
is sometimes a vital necessity;but in other instances in which only a
trivial advantage is
provided, decision is often difficult as to whether
the relationship is actually beneficial or merely neutral. Positively
harmful relations grade off in similar fashion to those that produce
only a minor inconvenience or are essentially neutral. The nature
of the relationship may change during the life cycle of one or both
of the species concerned. Furthermore, as with intraspecific relations,
the classification of an interaction between two species as beneficial
or harmful depends upon whether consideration is given to the imme-
diate effect on the individual or to the long-range effect on the species
as a whole. In our present discussion we shall consider first the na-
ture of interspecific relations as they act within the life
span of the
individual, and subsequently their more remote consequences to the
success of the population and
to the evolution of the species.
j With the foregoing qualifications in mind, we may divide the in-

terrelations between species into two main categories: (1) symbiosis,
in which one or both species are benefited and neither species is
harmed; and (2) antagonism, in which at least one of the species is
harmed. Some authors extend the meaning of symbiosis to embrace
all
types of interrelations including harmful ones; other authors, taking
the other extreme, limit the term to relations between a plant and an
animal, or to relations that are mutually beneficial. The present defi-
nition of symbiosis, which literally means "living together," seems
more logical and more in keeping with established usage. Symbiotic
associations are divided into those of mutualism (both species bene-
fited) and commensalism (only one species benefited). Relation-
ships of antagonism between species embrace antibiosis, exploitation

(including parasitism and predation), and competition.^
Various ecological aspects of the foreging subdivisions will be dis-
cussed in the following sections. E. F. Haskell's more elaborate clas-
sification of "coactions" between species is discussed and illustrated
by Burkholder (1952). A simplified arrangement of these inter-

specific relations may be represented as tabulated on page 364.
^j
In this scheme ( -[- ) indicates an increase in a beneficial life process
as the result of the association, ( )
indicates decrease or harm, and
(0) indicates no significant effect. Since these relationships grade
into one another and sometimes change, and since it is often difficult
to determine whether the effect on one of the species is
essentially
364 Relations between Species
neutral, attempts to classify many actual associations in these sub-

divisions may not be profitable. The scheme is chiefly useful in clari-
to be kept in mind while interspecific
fying the types of relationship
reactions are being examined.
Species A Species B Relation
+ + Mutualism
Isymbiosis
+ Commensahsm J
Neutrality, toleration .
Antibiosis
|
+ Exploitation (inc. parasitism and pro- > Antagonism

(iation)
J
Competition
SYMBIOSIS
When members of two species are living together in a symbiotic
relationship, the benefit received by one or both of them most fre-
quently involves the provision of food, but it may also involve shelter,
substratum, or transport. The association may be continuous or
transitory, obligate or
facultative. The two symbionts may be in close
contact, with their tissues actually intermingled, or one partner may
livewithin a cavity of the other or attached to its surface. In some
instances contact between the individuals is transitory, and in some
the two species may influence each other without actual contact.
Associations in which both species derive benefit are termed mutual-
ism; those in which only one species is benefited and neither is harmed
are termed commensalism.
Mutualism
Mutua\\stn \vvt\v Continuous Contact T\\e most intimate type oi

n\utY\a\\sm is seen in those associations in w\\ic\\ contact between the
symbionts is close and is often
permanent as well as obligatory. A
classic example of this sort of relationship is furnished by the lichens,
which are composed of a matrix formed
by a fungus within which
cells of an
alga are embedded. The fungus holds moisture and makes
minerals available for both
partners, in return for which the chloro-
phyll-bearing alga manufactures carbohydrates for itself and also for
the colorless fungus. The
fungus of a lichen can never grow in nature
without the associated
algae, and the algae, although
generally similar
to independent
species, are probably dependent
upon the fungus
under natural conditions. The dual of the lichen
personality be may
demonstrated in the laboratory by suitable techniques that permit the
algal and fungal components to be cultured separately, as is sug-
gested in Fig. 10,1. Many lichens grow abundantly on bare rock
surfaces where the lack of moisture and of organic matter would make
life
impossible for most independent algae and fungi.
Other equally fascinating instances of inutualistic symbiosis in-
volving two plant species are to be found in nature. A well-known
example, and one already mentioned as of great ecological importance
in relation is nutrients, is furnished by the bacteria of the
genus
Mature lichen
Ascospores
Pure cultures
Clasping hyphae^-
Association
Lichen is formed in nature but

rarely in laboratory
the
FIG. 10.1. Diagram the symbiotic partnership represented by
illustrating
lichen Cladonia cristatella and its in the laboratory into algal and fungal
separation
in the laboratory, the typical
components. When the components are recombined
not because of the lack of certain
morphology of the lichen is developed, evidently
Baitsell's
environmental conditions found in nature. ( Burkholder, 1952, Copyright
Science in Progress, Yale University Press.)

Rhizobium, which form nodules on the roots of leguminous plants and

live
symbiotically with their hosts (Fig. 10.2). The bacteria are
somewhat specific as to host, and obtain carbohydrate and other sub-
stances from its In return the rhizobia fix gaseous nitrogen
juices.
and pass it on to their plant host. If available fixed nitrogen is absent
from the soil, the plant is
completely dependent upon its
symbiotic
nitrogen-fixing bacteria.
FIG. 10,2. Vertical section through the root of a soybean plant, showing the root
nodules within which live symbiotic bacteria (Rhizobium japonicum).
A somewhat similar mutualism is seen in the fungi that form mycor-

rhizal structures either inside the roots of certain plants or on their
outside surfaces (Fig. 10.3). Ectotrophic mycorrhizae are found on
various kinds of trees such as pines, oaks, hickories, and beech; endo-
trophic mycorrhizae occur in the red maple and are particularly com-
mon in roots and other tissues of many orchids and heaths. Fungi of
this type are nourished by organic material that they absorb from
their hosts. The ectotrophic mycorrhizae commonly take the place of
root hairs and function in the
absorption of water and nutrient salts
from the soil. The degree of
dependency of the host plant upon
mycorrhizae very variable, but pine seedlings, at least, appear to be
is
quite unable to grow in soils normally deficient in one essential nutri-

ent without the aid of these symbiotic structures. Since the bene-
ficial
mycorrhizae of the blueberry, rhododendron, and heather
Photo by Somerville Hastings, from McDougall, 1949, Copyright, Lea and Febiger
FIG. 10.3. Ectotrophic mycorrhizae seen as whitish sheaths over the branching
rootlets of the hornbeam (Carpinus betulus)
growing in leaf mold.
flourish only in an acid medium, the growth of these plants is im-

proved by a soil of low pH (McDougall, 1949). It is reported that
settlers moving west in the United States were at first unsuccessful in
establishing certain kinds of trees around their homesteads because of

the absence of suitable fungi. The seeds that they
brought with
them sprouted, but often the young trees would not grow. When
fungal spores accidentally reached the plantations in soil samples car-
ried from the east, many of the necessary mycorrhizae were
supplied
and better tree growth became possible. Subsequently deliberate
was practiced,
inoculation especially for the growth of pine trees.
Even more remarkable associations involving the
intermingling
of tissues are those in which one partner is an animal and one a plant.
Unicellular plants live symbiotically in the outer tissues of certain
sponges, coelenterates, mollusks, and worms. Some of these uni-

cellular forms are green algae known as zoochlorellae; others are
brown or yellow cells, believed to be flagellates, and are termed zoo-
xanthellae. Partnerships called plant-animals by Keeble (1910) are
formed by a turbellarian worm (Convoluta roscoffensis) and large
numbers of zoochlorellae. The algal cells, often growing in the tissues
of the worm in such profusion as to give it a greenish appearance,
release oxygen during photosynthesis and produce nitrogen com-
pounds that are nutritionally beneficial to the host. In exchange the
algae obtain a suitable matrix for their growth and receive a supply of
nutrient material resulting from the animal's metabolic
processes.
This balanced symbiotic relationship is thus able to
persist indefi-
nitely, and the nourishment furnished by the algae enables the worm
to live and grow for
long periods without taking in solid food.
Zooxanthellae are found abundantly in the body wall of coral
polyps. These unicellular organisms serve the useful function of
removing nitrogenous wastes and carbon dioxide from the coral and
providing it with oxygen produced as a by-product of photosynthesis.
The zooxanthellae in turn benefit by the absorption of the metabolites
containing nitrogen and phosphorus, which are scarce in tropical
waters, as well as by the absorption of carbon dioxide resulting from
the catabolic processes of the coral animal. The fact that coral
polyps
placed in the sea in sealed glass containers survived for 2 weeks is
evidence that the symbionts in this instance had
approached a state of
balance, in regard to their respiratory exchanges at least, and
perhaps
also to some extent in regard to other needs.
The giant clam Tridacna (Fig. 7.11) grows on shallow coral reefs
with the opening between its shells directed
upwards and the broad
edges of its mantle, containing vast numbers of zooxanthellae, spread
out horizontally where they receive intense radiation from the sun.
The mantle also contains great numbers of small lens-like
organs
which probably serve to focus light into the tissue,
making photo-
synthesis possible for the deeper-lying /ooxanthellae. Since phagocy-
tic blood cells
regularly engulf and digest large numbers of the zoo-
xanthellae, the giant clam may be thought of as
"farming" these sym-
biotic algae and deriving a considerable
portion of its nutrition from
them (Yonge, 1944).
Some symbionts reside in cavities of their hosts rather than in their
tissues. Ruminants and other animals living on a diet high in cellu-
lose are unable to digest this material without the
enzymatic action
of cellulase produced by microorganisms in their intestines.
Sym-
biotic bacteria fulfill this function in cattle and other
grazing animals.
Certain cockroaches and termites can digest wood only with the aid
of a special type of flagellate that is harbored within their guts.
Symbiosis in these instances is mutually beneficial and obligatory for

both parties. Some of the bacteria living in the intestines of animals
also produce various B vitamins and other special materials.
Another manifestation of mutualism in which the symbionts are in
permanent contact, but one in which the contact is entirely external

is the attachment of certain marine
sponges and coelenterates to the
shells of crabs. The attached animal benefits by being carried about
to fresh feeding areas and by avoiding being stranded in the tidal
zone or in stagnant water, as well as by obtaining fragments of food
from the meal of its host. The crab, for its part, is camouflaged to
some extent by the presence of the attached animals on its back and
is often protected by them from attacks by its enemies.
FIG. 10.4. Three sea anemones attached to the shell of a hermit crab, illustrating
mutually beneficial symbiosis. Note the growth of the foot of the anemone over
the surface of the shell. ( Modified from Borradaile, 1923. )
The classic example of this type of partnership under the sea is

furnished by the sea anemone Adamsia palliata which grows on the
shell of the hermit crab Eupagurus prideauxi (Fig. 10.4). Further
relations result in this instance from the fact that the shell
interspecific
inhabited by the hermit crab is the abandoned house of a snail and
from the fact that the same shell may also furnish the abode of the
annelid worm Nereis. The hermit crab starts the
enterprise by ob-
370 Rclatiotis between Species
taining a sea anemone from a rock and placing it on the back of its
shell. When the growth of the crab causes it to move to a larger

shell, the crab loosens the base of the anemone with its claws and
transfers it to the new shell. In some species the base of the anemone
grows over the entire shell and extends beyond it; sometimes the
that the crab's house con-
original shell largely dissolved away so
is
sists almost entirely of the anemone's base. The stinging nematocysts

of the sea anemone are a powerful deterrent to predaceous fish and
protect the members of the partnership
from being eaten. When
Nereis also present as a junior partner, it is said to help keep the
is
inside of the shell clean and, as payment, to snatch fragments of food

from the pincers of the crab.
Mutualism without Continuous Contact. A great many instances
exist of mutualism in which the partners are not attached to each other
or in which they are in contact intermittently or for only a short pe-
riod. In most mutual benefit associations of this sort the fulfillment
of a nutritional need plays a prominent role for at least one of the
species; but the other species, and sometimes both, may derive an
of advantage.
entirely different type
A commonly cited example of mutualism of two animal species in-
volves birds that alight on the backs of large grazing animals and
pick off the ticks or other external parasites. The cowbird in North
America, the oxpecker, the white heron in Africa, and certain
little
other birds obtain a ready supply of food in this way. The host ani-
mals are rid of their pests and are frequently warned of approaching
danger by the activity of the birds as watchmen. An amazing kind
of pest-control service is rendered by the crocodile bird in
removing
leeches from around the teeth of the crocodile, which allows the bird
to enter its mouth for the search. Mixed groups of ostriches and
zebras are said to derive mutual benefit in guarding against attack
by the keener sense of sight of the ostrich and the greater powers of
scent possessed by the zebras.
In the subterranean world we find various
extremely complex re-
ciprocal relations between species. Various ants maintain a popula-
tion of aphids in their nests. The ants obtain a nutritive exudation
from the hind end of the alimentary tract by stroking the aphid's
Abdomen with their antennae. This furnishes the basis for popular
accounts that ants "keep cows" and "milk" them. The
aphids feed
on the roots of plants, or they are carried by the ants out of the nest
and allowed to feed on leaf stalks. The aphid eggs are laid on the
plants above ground, and, although of no immediate use to the ants,
they are carried down into the nest where they are sheltered during
the winter until they hatch and the young aphids repopulate the dairy
farm (Fig, 10.5).
An equally extraordinary agricultural technique practiced by in-
sects isthe cultivation of fungi by certain beetles, ants, and termites.
Various kinds of fungi are grown for food by these animals, and some
species are known only in insect gardens of this sort. In the tropical
forestone may see a band of green moving across the ground; this is
formed by a line of leaf-cutting ants (Atta) carrying pieces of leaves
FIG. 10.5. Diagram of an ant nest containing a "dairy farm" of aphids. ( Burk-
holder, 1952, Copyright, Baitsell's Science in Progress, Yale Univ. Press.)
over their backs like so many umbrellas. The leaves have been cut
from a shrub and are being transported to the ant's underground nest
where they will be chewed into a pulp and spread out to form a bed
in which a particular kind of fungus (Rozites gongylophora) is
planted. The ants cultivate the garden with great care; they weed
out unwanted species of fungi and prevent the fruiting of their
fungus, but encourage it to develop special mycelial outgrowths on
which they feed ( Brues, 1946 )The fungi that live symbiotically in
.
termite nests are nourished by the insects' excreta rather than

by leaf
pulp. The cultivation of fungi by ambrosia beetles (Scolytidae) is
carried on within the tunnels drilled in wood by these insects ( Cham-
berlin, 1939). Sometimes the development of the fungi is sufficiently

luxuriant to clog up the tunnels completely, and the beetles are killed
unless they can eat their way out faster than the fungi grow!
The perpetuation of this mutualistic relation between insects and
fungi is assured elaborate structures and reactions by which the
by
fungi are transferred to the habitats of new generations.
The fungus-
beetles,for example, possess or internal
certain external
growing
structures by means of which they carry spores or fragments of the
to the site
fungi from the old burrow, where the larvae were hatched,
of a newly founded colony. Among the leaf-cutting ants the virgin
queen carries a pellet of fungus in a pocket below the mouth and
deposits the inoculum in her new bridal chamber.
These arrange-
ments by the host for transmission are strong evidence that the guest
species are beneficial, and hence are symbionts. With parasitism,
the
of transmission is for the in one
problem always arranged by parasite
way or another and is resisted by the host. A more detailed consid-
eration of these relationships among insects will be found in the sum-
mary by Steinhaus ( 1946 ) .
The pollination of flowers by bees, moths, and butterflies, and oc-

casionally by hummingbirds, is another manifestation of mutually
beneficial symbiosis, but one in which the species concerned may be
in contact for only a few seconds. The insect derives food from the
nectar, or other product of the plant, and in return carries pollen from
the anthers of one flower to the stigma of another, thus ensuring cross
pollination.
v The coordination of the elaborate behavioristic and
anatomical adaptations that have been evolved in the insect and in
the flower in relation to this cooperative activity is truly remarkable.
The flowers open, producing odors and displaying colors that attract
suitable insects, only at the time when they are
sufficiently mature for
fertilization. In various ingenious ways the flower is so
shaped that
the insect cannot get its food without dusting the
stigma with pollen
carried from another flower and then picking up fresh
pollen to be
carried to the next flower. The reactions and the structure of the
pollinating insects are correspondingly adapted. Bees, for example,
continue to visit the same species of flower as
long as a supply of
mature blossoms is available and thus avoid mixing pollen from dif-
ferent species.
The mutual dependence of insect and flower is
frequently highly
specificand sometimes involves the reproductive cycle of the animal
as well as that of the plant. The
peculiarly enclosed flowers of the
commercial are pollinated only by wasps of the genus
fig Blastophaga,
and special floral structures called caprifigs provide the only place in
which these wasps can lay their eggs. A similar dually obligate and
specific symbiosis is found in the relation between the yucca plant
and the yucca moth (Fig. 10.6). The female moth visits the yucca
flower in the evening and collects a ball of pollen from the anthers.
Then, holding the pollen ball in specially adapted mouth parts, she
flies to another
plant and pierces the ovary of the flower with her
Fie. 10.6. The Yucca moth (Pronuba ijnccasella) approaching flower of the
yucca plant (left). Flower cut open (right), showing the moth placing pollen on
the stigma. ( Modified from Borradaile, 1923. )
After depositing eggs within one of the ovules, the moth

ovipositor.
creeps down
the style and stuffs the ball of pollen into the stigma.
When the moth eggs hatch, the larvae feed on the tissue of the ovule,
and eventually mature to repeat the process for the next generation.
It is difficult to
imagine in such complex intcrdependencies how the
behavior pattern and anatomical structures vitally necessary to both
species have evolved.
Other two-way benefits in the relations between species involve
transport and dispersal of seed. The fruit eaten
by birds, mammals,
and other animals provides them with a source of nourishment, and
the contained seeds are subsequently dropped in their excrement at
varying distances from the original site. The activity of squirrels in

carrying and burying acorns, hickory nuts, and the like may play a
significant role in the establishment of trees in new areas. Many

other illustrations of mutualism, both of an intimate and of a casual
nature, will occur to the reader.
Commensalism
When members of different species are associated in such a way
that only one of the organisms is benefited but neither is harmed, the
^ Such associations no doubt
relationship constitutes commensalism.
began by the mere toleration of "guests" near the usually larger host
species, or on or in its
body. If the guest derived some benefit with-
out interfering with the host, the relationship would tend to persist.
Casual association may have led to a partial or a complete dependency
on the part of the guest. Obligate commensalism established in this
way may have evolved further in some instances to give rise to mutual-
ism, on the one hand, or parasitism, on the other. If the host
species
became adapted to take some advantage of the close proximity of its
guest, a mutualistic symbiosis would result. However, if during the
course of evolution the guest species imposed more and more upon
its host,
finally overstepping the bounds of hospitality and inflicting
harm upon the host, the relationship would change to exploitation and
perhaps to parasitism.
The advantage derived by the commensal involves the provision
of substratum, shelter, or transport, and very frequently of food.
Commensalism means "eating off the same table" as guest messmates.
"The messmate does not live at the expense of his host; all that he
desires is a home
or his friend's superfluities" (Pearse, 1939). As
would be expected, the circumstances of commensalism are more
variable than those of mutualism. Although in some instances the
commensal is in continuous contact with its host attached to a sur-
face or retained within a cavity more frequently the guest is free to
come and go at irregular intervals. Sometimes the commensal species
can associate with only a single host species, but often considerable
species flexibility is observed.

Commensalism with Continuous Contact. Commensals in more or
less permanent contact with their hosts are represented by a great
variety of epiphytes and epizoans. Many tropical orchids, brome-
liads, and other "air plants" grow perched on horizontal branches or
in forks of trees or hanging in streaming festoons (Fig. 10.7). Fa-
miliar in the northern coniferous forest are the hanging "mosses"
Usnea and Alectoriattliat gave rise to Longfellow's well-known lines:
\j
Commensalism with Continuous Contact 375
This is the forest
primeval. The murmuring pines and the hemlocks,
Bearded with moss, and in garments green, indistinct in the twilight,
Stand like Druids of eld, with voices sad and
prophetic,
Stand like harpers hoar, with beards that rest on their bosoms.
Each two species forming a lichen, and

of these mosses consists of
thisgrows an
as epiphyte upon the conifers, thus creating a three-way
partnership. Epiphytes sometimes show distinct preferences for one
host species, as may be seen, for
example, in southeastern United
Photo by H. B. Moore
FIG. 10.7. Bromeliads, including the pendant Spanish moss ( Tillandsia ) , growing
as epiphytes on the branches of the live oak in Florida.
States where the Spanish moss grows more abundantly on the live
oaks than on the pines (Fig. 10.8).
All these epiphytes use the trees only as a point of attachment amid
suitable light and other conditions, and manufacture their own food
by photosynthesis. Since they do not obtain nourishment from the

tissues of the tree,

they are not classed as parasites. For the most
part the epiphytes do no harm to the host plant but occasionally they
become so numerous as to break it down or to. stifle its growth ( Fig.
10.8). Orange growers in Florida are forced to spend large sums
every year for the removal of Spanish moss from their trees.
FIG. 10.8. Spanish moss (Tillandsia) growing in harmful abundance on a live

oak tree in South Carolina but practically absent from a neighboring pine.
Some plants live as attached epiphytes on the surfaces of animals.

One extraordinary example is the green alga that grows on the long,
grooved Since this alga often becomes sufficiently
hairs of the sloth.
abundant to give the animal a greenish appearance, the sloth pre-
sumably derives some advantage in concealment as it sleeps in the tree
tops. Equally remarkable in its habit is the green alga Basicladia, a
genus of the Cladophoraceae, which grows only on the backs of fresh-
water turtles (Leake, 1939). The specific relation here appears to
be due to a dependence on keratin, since the alga can be cultured in
Commensalism without Continuous Contact 377
the laboratory only if this substance is

provided. To the extent that
the host receives benefit from such
epiphytic algae, the relationship
approaches mutualism.
Many kinds of microorganisms take up residence within tissues or
cavities of larger plants and animals without causing any trouble for
the owner but without paying any rent. Since no light is available
for photosynthesis thesecommensals are represented by saprophytic
fungi and bacteria and by Protozoa. Many such organisms are
found in the lower intestines of animals where they consume un-
digested food and secretions and complete their life

cycles unnoticed
by their hosts. The bacteria in the human colon, notably Escherichia
coli, are a familiar example. As already mentioned, these commen-
sals may represent a mode of life that is transitional between that of
the parasites and the beneficial symbionts also found in such habitats.
Permanently fixed commensals in the animal kingdom are repre-

sented by sessile invertebrates that grow attached to plants or to other
animals. Sometimes a highly spedific relation exists between the
epizoan and its host, although the reason for this is often hard to dis-
cern. The
oyster-like bivalve Ostrea frons grows almost exclusively
on the roots of the red mangrove in the shallow waters off the coast
of Florida; special hooks develop on the lower shell of this animal by
means of which it clings tenaciously to its host and is not displaced
by surf. Certain barnacles are found only on the backs of whales

where they benefit by a free ride; since they do not feed upon the
whale's flesh, they are not parasites in the strict sense of the term.
The marine environment furnishes many other instances of attached
commensals (Wilson, 1951, Ch. 11). Anyone who has an opportunity
to catch an elderly horseshoe crab (Limulus polyphemus) in the
shallow water off the New England coast is likely to find several
species of mollusks, barnacles, and tube worms attached to the shell
and a number of more motile commensals living in the "book gills"
or other anatomical nooks of this strange animal.
Commensalism without Continuous Contact. The category of
commensals that are in
temporary contact with their hosts or that are
associated without being in actual contact is also a large one. Inter-
mittent contact between animal commensals and plant hosts is dis-
played on land by squirrels, monkeys, tree frogs, and snakes, and by

a great many birds, insects, and other animals that use trees or other
plants for substrata, for shelter, or for breeding sites without harming
the host plant significantly. yMany examples of partial or complete
specificity inCommensalism of this type will occur to the reader. An

intriguing three-cornered relationship is that of the elf owl that nests
only in abandoned holes made by the Gila woodpecker in the stems

of the large Sahuaro cactus found in Arizona and neighboring re-
gions.
Temporary or intermittent contact between two animal commensals
is seen in such special associations as that of the remora fishes and the
sharks, whales, or sea turtles to which they attach. The dorsal fin of
this "shark sucker" has become modified in the course of evolution into
a most effective suction disc by means of which the fish attaches itself
to the under side of a shark or other large animal. The sucker can
release its
grip at will, swim about gathering fragments of food re-
sulting from the shark's meal, and return to hook another ride on the
body of the host. Several suckers may be found attached to the

same shark, but they do not seem to hinder the powerful fish ap-
preciably.
Commensalism involving between two species but
close association
without the attachment of the guest to the host occurs in a wide
variety of animal groups. The guest may live in a burrow or other
retreat of the host species, and sometimes the relation is highly specific.
Certain beetles, for example, are known exclusively from the nests
of meadow mice. The burrowing owl which often nests in the bur-
row of a prairie dog is an accepted member of the prairie dog "village"
while the village is actively populated by these rodents. Many such
uninvited guests take up residence in the burrows or tubes of aquatic
animals. The decapod crustacean Polyonx lives in the "back entry" of
the U-shaped tube of the marine annelid Chaetopterus, where it is
well hidden from enemies and where it can obtain particles of food as
well as a supply of oxygen from the water forced through the tube by
the pumping action of the worm's parapodia (Fig. 10.9).
Other marine commensals live within a water cavity of their host.
In certain instances in which the relationship is highly specific the
commensal has been shown to be guided to the proper host by chem-
ical emanations or to be kept within the host by definite tactic re-
actions (
The oyster crab is a familiar example of
Davenport, 1950 ) .
thistype of commensalism and one among several of a similar nature

in which the commensal eventually becomes a prisoner. The oyster
crab is
originally carried as a planktonic larva into the mantle cavity
of the oyster by incoming water currents produced by the feeding and
respiratory activity of the host. The tiny larva metamorphoses, and

eventually grows into an adult about a centimeter in length and too
large to escape through the narrow opening between the valves of
the oyster. The crab leads a completely sheltered life, stealing par-
ticles of food from the oyster, to be sure, but
apparently doing its
Commcnsalism without Continuous Contact 379
host no significant amount of harm. The commensal is still within

the cavity of the shell when the oyster is opened in the kitchen, and
1
the crab sometimes startles an uninformed diner by appearing in his
oyster stew. (Oyster crabs are perfectly edible and. are regarded
as a delicacy by some, including many who are not ecologists.)
Perhaps the most extraordinary instance of a commensal living
within its host is that of the small tropical fish Fierasfer which finds
^.k4ffcL'|iti* -
FIG. 10.9. Diagramiuatic vertical section of a Chaetopterus tube in the mud

bottom of the littoral zone, showing the position of the worm and two commensal
crabs (Polyonyx). (By permission from Animal Ecology by A. S. Pearse, Copy-
right 1939,
McGraw-Hill Book Co.)
shelter within the cloacal cavity of a sea cucumber ( Holothuroidea )

.
The fish occasionally emerges to feed in the neighborhood. When it
wishes to reenter its

strange retreat, it
pokes its nose against the open-
ing of the cucumber's cloaca, then quickly reverses its

position, and
allows itself to be drawn tail first inside its host.
Some species appear to derive protection by acquiring
a big brother,
that themselves with other in the
is, by associating species recognized
community as being voracious or poisonous. Pilot fish (Naucrates
ductor) follow along beneath sharks as closely as shadows, but can-

not attach to the surface of their hosts since they have no suction
discs as do the remoras. For some reason they are never eaten by
the sharks. In similar fashion particular kinds of fish find shelter
under the umbrella of poisonous jellyfish;
one species somehow man-
ages to live unharmed among the tentacles of the Portuguese man-
of-war, whose stinging cells instantly paralyze other kinds of fish,
which are used for food. Another commensal fish has acquired a
type of swimming motion that allows it to move among the tentacles
of large sea anemones without causing the
discharge of the poisonous
nematocysts.
Other instances of the protection gained by the weak through as-
sociation with the strong are furnished by scavengers and mimics, al-
though their modes of life should perhaps not be termed commensal-

ism because of the absence of toleration or even awareness. In the
tropics the hyena lives as a scavenger on leavings from the lion's meal,
which consists of animals that the hyena could not possibly have
killed for itself. Similarly, in the far north during the winter the
arctic fox lives largely upon the remains of seals killed by polar bears.
Another type of protection is attained by weaker species through
mimicking in appearance and behavior certain species recognized as
voracious or poisonous by other members of the community, as de-
scribed in Chapter 6. The mimics, as well as the truly dangerous
models, are presumably avoided by the predators.
The foregoing account of symbiosis is sufficient to indicate the
widely separated types of animals and plants that may be associated
in relations of advantage or of toleration. We have also reviewed
the great range that exists in the degree of dependence and of specific-
ity in these relationships. However, many fascinating questions

await future study as to the exact nature of the behavior patterns that
bring the symbionts together and of the physiological interdepend*
encies between them. The examples described show how these asso-
arisen from casual contacts between
ciations may have species, which
then led to a recognizable commensalism. Subsequently the relation
Antagonism 381
may have evolved in some instances into mutualism and in others into
parasitism. The possibility of evolutionary transformation in either

direction may be imagined from such flexible associations as those of
the various mycorrhizae.
The interspecific relationships described in this section are flexible,
and their nature may
change even within the lifetime of the individ-
uals concerned. At one moment the tissue of the giant clam is pro-
viding a favorable environment for the growth of zooxanthellae; at
another moment it is digesting these unicellular organisms. Fungi
associated with the roots of the tomato plant are ordinarily harmless,
or even beneficial; but under certain conditions these fungi become
definitely parasitic on the same plant. Another turn-about relation-

ship, which will be more fully described
in Chapter 12, occurs in
stands of "old field" pine. The grass in abandoned fields in central
New England forms a favorable bed for the development of seeds
that blow from white pine trees. When the pines have grown into
in
a dense grove, however, the resulting deep shade and thick carpet of
fallen needles will kill the grass beneath. These examples will suf-
fice to indicate how easily the line between symbiosis and antagonism
may be crossed.
ANTAGONISM
Relations between members
of different species in which one or
both are harmed during the life span of the individuals concerned
are included under the general heading of antagonism. These antag-
onistic relations are not necessarily harmful for the population or for
the species as a whole. From the broader viewpoint interspecific
antagonism may have beneficial effects through controlling abun-

dance or influencing the course of evolution V
Exploitation and com-
for the very existence of the community, and,
petition are necessary
when kept in suitable balance, make possible its perpetuation.^ Our
scrutiny of the relations between
individuals of different species there-
fore will lead to a consideration of the relations between interacting
populations. The involved in functional adjustments of the

latter are
as a will be discussed in subsequent chap-

whole which
community
ters. As with other ecological factors, when the interaction between
species extended from individuals to populations and the entire
is
community, additional relations with new significance emerge.

Antagonistic relations of
one species toward another involve harm
that may be one or more of several ways: (1) Species
inflicted in
A may produce a poisonous substance or a change in environmental

condition that is inimical to species B without species A deriving any
benefit (0, ). Antibiosis; the production of harmful secretions, is

the outstanding example of this type of relationship. (2) Species A
may inflict harm by direct use of species B for its own benefit in a
Species A may exploit
relationship of exploitation^ -f-, ).
B by
gaining support or shelter for itself, or it
may obtain nourishment di-
rectly from B as a parasite or as a predator.* (3) Both species may be

harmed in a reciprocally unfavorable relationship ( ). Thijj ,
situation is
commonly found in the indirect rivalry, or competition, of
two species for some feature of the environment that they both need
and that exists in short supply. The space, food, light, oxygen, or
other necessity taken by one species reduces the amount available for
the other. In some situations considerable harm is suffered by both
competing species; in other instances the injury may be serious for one
species and relatively
minor for the other. A reciprocally harmful
effect will also occur if
species A produces an antibiotic harmful to
FIG. 10,10. Roots of strangling fig enveloping the trunk of its host in a Florida
hammock.
Antibiosis 383
species B and at the same time an antibiotic secreted by B is harmful

to A. v
In the foregoing
paragraph the term competition was used in a re-
stricted sense
referring to the indirect rivalry of two species striving
for some The
necessity from a limited supply. aggression between
species that prey on one another or exhibit some other form of direct
antagonism may be considered competition in the broad sense.
also
The ecological effects of competition as both direct and indirect
antagonism will be considered both from the point of view of the in-
dividual and of the population and the species.
All gradations exist between relations that are harmful to one
species and neutral to the other, those that are harmful to one and
beneficial to the other, and those that are harmful to both parties.
Associations between species that are at first neutral or even bene-
ficial
may change during the life of the individuals to become harmful
to one species, as already mentioned* and this relationship may change
further to bring disadvantage to both parties. For example, the
strangling fig starts life as a harmless epiphyte on a palm or other

tree, but after a time its
dangling roots reach the ground. Deriving
nutrients from the soil and energy from sunlight, the fig rapidly en-
its host with a network of
velops the trunk of anastomosing roots
(Fig. 10.10). The luxuriant growth of the upper part of the fig grad-
ually shades out the palm, and, after the death of the host tree, the fig
eventually falls to the ground from lack of support. In similar fashion
a parasite or a predator may bring about its own destruction under
certain circumstances by killing its host or consuming all of its prey.
Antibiosis
Manysubstances produced by organisms, or conditions resulting

from their metabolism, are generally harmful to other sr Thus the
generation of carbon dioxide or of organic acids may harm more sen-
sitive species to such an extent that they are unable to continue to
live in the areaând excessive shading by one kind of vegetation will
kill off species intolerant of low illumination.*' The term antibiosis
applies more particularly to the production of materials that are

specifically antagonistic to other speciesXSuch antibiotic substances
are known be generated by many kinds of fungi and bacteria.^ The
to
action of penicillin, streptomycin, aureomycin, and other antibiotics
in destroying various pathogenic bacteria is fa-
produced by fungi
miliar to the reader through the practical use of these substances
either natural or syntheticin medicine. *
Bacteria, molds, and actinomycetes that produce antimicrobial sub-

stances have now been shown to be widespread in nature. Burk-
holder (
1952 ) reports that about half of the species of actinomycetes
and half of the lichens as well as large numbers of higher plants pro-
duce substances that inhibit certain molds and bacteria. Under
some natural conditions antibiotics are believed to protect the species
producing them from bacterial marauders, but this ecological aspect
of the subject is
largely unexplored.
Antagonistic substances formed by algae have been shown to af-
fect other algae adversely in the laboratory and undoubtedly also do
so in nature under some circumstances. Some substance accumulat-
ing in cultures of the green alga Chlorella vulgaris was found to in-
hibit the growth of the diatom Nitzschia frustrulum to a greater ex-
tent than its own growth; similarly, an antibiotic produced by dense
cultures of Nitzschia retarded the division rate of Chlorella grown in
the same culture or in conditioned water filtered off from the diatom
culture. Interspecies antagonism of this sort probably exerts control
on the abundance of different kinds of phytoplankton in ponds or
other aquatic areas, and in some instances it may influence the sea-
sonal sequence of species commonly observed in nature ( Rice, 1949 ) .
Similar products of plant growth have been found to be harmful

to certain animals. Substances produced by senescent cultures of
Chlorella and of the diatoms Navicula and Scenedesmus inhibit the
feeding of Daphnia in laboratory tests. A reaction of this sort

filter
may account in part for the curtailment of the growth of

zooplankton
sometimes observed in natural waters containing senescent popula-
tions of phytoplankton (Ryther, 1954&). Pond "blooms" of blue-green
algae, especially of the genus Microcystis, are known to produce toxic

substances, such as hydroxylamine, which cause the death of fish and
even of cattle that drink the water (Prescott, 1948). In the marine
environment toxins produced by huge populations of certain micro-
organisms, popularly known as "red tide," cause the catastrophic de-
struction of fish and other animals. An outbreak of the dinoflagellate
Gymnodinium brevis off the west coast of Florida in 1946-1947 re-
sulted in the wholesale death of fish throughout an area of several
thousand square miles (Gunter et al., 1948).
Certain land plants are similarly harmful to animals, but they usu-
ally exert their poisonous effects only

when eaten. For example, a
semidesert bush (Halogeton glomeratus) found in Nevada and neigh-
boring states kills the sheep that eat it. The spongy leaves of this
weed are filled with oxalic acid, which combines with the calcium
in the blood and causes death within a few hours.
Exploitation 385
poisoning are known, some of them acting

Many other instances of
in a highly specific manner. The harm done in this way by one
species to another may be completely fortuitous in some instances, but
it
may have evolved as a protective adaptation in others. The ecolog-
ical consequences of this
type of antagonism, involving the produc-
tion of poisons or of harmful metabolites, have been discussed in fur-
ther detail by Lucas (1949).
Exploitation
We shall now turn to relations of exploitation in which the mem-

bers of one species benefit by the utilization of another speciesX The
most common manifestation of interspecific exploitation is the use of
a neighboring species as a source of food^nd this will be discussed
tinder the headings of parasitism and predation. However, an or-
ganism may employ another species of plant or animal in the environ-
ment for attachment, support, or transport, as already mentioned, and
such activity often results in harmful consequences to the host.
Certain highly specialized kinds of exploitation occur, and one of
the most astonishing of these is the enslavement of one species of ant
by another^Talbot and Kennedy, 1940). Polyergus is an obligate

slave-making ant that unable to maintain its colony without the
is
presence of members of certain species of Formica. Polijergm work-

ers raid neighboring nests of Formica and carry home larvae and
pupae. After maturing into the worker stage, these captive ants
undertake feeding and nest building for their masters. Pohjergus
ants will starve even in the presence of abundant food if this slave
labor is not available. ^
Another special type of exploitation is that practiced by so-called
"parasitic birds/'
The cuckoo in Europe and the cowbird in the
United States never build nests of their own; each female lays an
in the nest established by birds of another,
usually smaller,
egg
species. The cowbirds'
habit perhaps arose as a result of their
nomadic wandering after grazing animals, which left them insufficient
time in one place for such domestic matters as raising a family. The
is sometimes able to throw out the
pair of birds thus imposed upon
foisted egg orit
by building a new nest bottom over it, but
to cover
more frequently the unwanted egg hatches out along with the host's
own eggs. The young cuckoo soon pushes the other fledglings out
of the nest, and the young cowbird is able to starve out its nest mates
all the food. After the young imposter has left the nest,
by grabbing
its still calls forth in its foster parents the behavior pattern
squawking
of feedingand caring for it even in the absence of the rightful

young.
The spectacle of a small sparrow or warbler frantically trying to satisfy
the hunger of an adolescent cowbird four times its size is an
perhaps
amazing sight.
The "parasitic" activities of other kinds of birds are really in the
nature of The
reader may have witnessed the spectac-
"hold-ups."
ular sight of an
eagle attacking an osprey high in the air, forcing it
to
drop a fish from its talons, and then catching the fish before it
reaches the ground.
Similarly, a skua or a jaeger will chase a gull or
a fulmar until the
pursued bird drops its fish, or disgorges a meal al-
ready swallowed, and then the bully secures the food for itself.
Various other kinds of animals are professional
highwaymen, includ-
ing some that compound the felony by robbing thieves. Such hi-
carried on by certain flies
is
jacking (Bengalia) that waylay ants re-
turning from a raid on a termite nest. The
fly attacks an ant carry-
ing away a termite, causing it to drop its booty, whereupon the fly
quickly consumes the stolen goods.
Parasitism. Strictly
speaking, a parasite is an organism that
resides on or in the
body of a larger living and derives
organism
nourishment from
Âccordingly, the foregoing instances
its tissues.
of exploitation are not
regarded as parasitism in the more precise
meaning of the term, either because nourishment is not involved or
because one organism does not live in contact with the other. */ Thus
barnacles growing on the back of a whale, as well as other kinds of
epizoans and epiphytes, are not true parasites since they do not eat
the host's tissue. However, the limits of
parasitism even in the strict
sense are not sharp. Just as
gradations exist between parasitism,
commensalism, and symbiosis, so also many borderline situations ex-
ist between
parasitism and predation. Some derive
organisms only
a part of their nourishment from their hosts and some are in contact
with their hosts for only a part of their lives. The
typical parasite
lives in its host without
causing its death, and the typical predator
kills the
prey upon which it feeds. Yet some kill
parasites regularly
their hosts, and some organisms classified as predators eat
only a part
of their prey sometimes without causing significant harm.Generally
speaking in parasitism the weak benefit at the expense of the strong,
whereas in predation the relations are reversed and the
strong exploit
the weak.
Examples of partial parasitism are found among various kinds of

plantsand animals. The mistletoes (Loranthaceae) grow like little
shrubs on the branches of trees. Their specialized roots
penetrate
the vascular tissues of the host whence they obtain water with the
Parasitism 387
contained dissolved minerals. Mistletoe
plants may be definitely in-
jurious to the tree on which they grow, but, since they are abundantly
supplied with chlorophyll, they photosynthesize their own carbo-
hydrates. Blood-sucking bugs, flies, and leeches and vampire bats
may be in contact with their hosts for only short and, al-
period?,
though such animals are commonly regarded as parasites^heir mode
clearly on the borderline between parasitism and predation.
of life is
Little basis exists for
considering mosquitoes, which live on the blood
of animals, to be parasites and not so
considering aphids, which suck
the juices of leaves. But if
aphids are regarded as parasites, their use
of the material of a
living plant is no different in principle from that
of caterpillars which eat the leaf substance from the outside as herba-
ceous predatorsor from that of deer which browse on the same leaves
from the ground. Many partial
parasites can derive nourishment
from a wide variety of host species. These considerations us bring
to the conclusion that
every gradation exists between obligate para-
sitism at one extreme, in which the is
parasite completely dependent
upon a living host with which
permanently associated, and preda-
it is
tion at the other extreme, in which a

free-living predator catches,
kills and devours its
prey.
Many kinds of plants and animals have taken up a completely para-
sitic mode of existence.
Although the largest representation comes
from the lower organisms, certain members of more advanced
groups
have also resorted to parasitism. Plant parasites, mostly fungi and
bacteria, may' attack animals or other plants. Animals that
parasitize
other animals are found in the Protozoa, in various other invertebrate
groups, and rarely in vertebrates. Animals that parasitize plants are

represented by gall wasps and gnats. The eggs of these insects are
commonly laid on stems and leaves, and the activities of the young
cause the formation of galls on the plant host. No
organism is known
that is not susceptible to attack by parasites of some sort.
Some parasites are restricted to one host species, or to one type
of host, whereas other parasites can attack
widely different plants and
animals. Parasitism may even occur within a For example,
species.
in the deep-sea angler Photocorynus spiniceps the male lives as
fish,
a tiny permanent parasite upon the head or side of the female and ob-
tains his entire nourishment from her blood
supply. Thus does this
species solve the problem of the location of one sex by the other in
the inky blackness of the deep sea. This type of
intraspecific relation
occurs frequently in the plant kingdom, as seen, for
example, in the
growth of the pollen tube representing the male plant as a "parasite"
on the tissues of the stigma and style of the flower of an angiosperm, or
again in the growth of the sporophyte of a moss upon the tissues of

chapter, however, we are concerned
the leafy gametophyte. In this
primarily with the parasitism of one species on another species.

Great variety is displayed among parasites in regard to their loca-
tion on their hosts, the duration of their dependency, their adapta-
tions, and their methods of transmission. Only brief mention of the
more ecological aspects of this large subject can be made here. For
a more thorough discussion of parasitism the reader should consult
such authors as Pearse (1942), Chandler (1944), or Baer (1951).
Parasitesmay occur on the outside of the hosts (ectoparasites) or
within their cavities or tissues (endoparasites). Certain worms and
many bacteria and protozoans are obligate parasites practically
throughout their lives; other organisms may be parasitic for only a
small part of their existence. Facultative parasitism occurs widely
and found, for example, in mosquitoes that can live their whole
is
lives on nectar and other plant exudates but will readily parasitize
any available mammal. However, some species of mosquito must

obtain a blood meal before they can reproduce.
Many part-time parasites are parasitic as adults and free living as

larvae, but in other species the reverse is true. A
rather special con-
dition within the latter category is presented by the parasitic wasps
that lay their eggs in the bodies of other insects. When the eggs
hatch, the larvae eat the tissue of the host, which at first remains
active. Since the host is always killed eventually by the growing
is sometimes
stowaway, the term parasitoid applied to insects follow-
ing this life pattern. Certain of the true wasps complicate the pro-
cedure further by stinging their victim into permanent paralysis be-
fore depositing the egg on it. In this way, the parent wasp provides
a living but helpless insect of another species, or a spider, for the
nutrition of the larvawhose mode of life is thus transitional between
parasitism and predation. Perhaps the most extraordinary example
of this behavior is the attack of digger wasps, the largest of which
have a wing span of about 10 cm, upon the even larger tarantulas.
When the female of the giant wasp Pepsis rnarginata is ready for egg
laying, she somehow locates a tarantula Crijtopholis portoricae, and

explores it with her antennae to make sure that it is the correct species.
The larvae of each species of wasp can be nourished by only one
species of tarantula. Although the tarantula could easily kill the

wasp, it does not do so, and makes little attempt to escape. After the
wasp has dug a grave for its intended victim, she stings it (Fig. 10.11 ),
drags it into the grave, and lays a single egg, which she attaches to
the abdomen of the paralyzed monster. At hatching the wasp larva
Parasitism 389
is
only a tiny fraction of the bulk of the tarantula, but, by the time
it is
ready for metamorphosis and independent life, it has consumed
all the soft tissue of the giant spider ( Petrunkevitch, 1952).
FIG. 10.11. The giant wasp (Pepsis marginata) stinging the tarantula (Cnjto-
pholis portoricae) preparatory to attaching an egg to its abdomen. (Petrunke-
vitch, 1952, Scientific American, drawing by R. Freund. )
and animals are susceptible to invasion by several to many

Plants
species of parasites at the same time, and conversely many parasites

can or must have more than one type of host during their lives. The
fact that parasites infest other parasites has been immortalized by the
jingle:
Fleas have lesser fleas

Upon their backs to bite 'em,
And lesser fleas still lesser fleas
And so ad infinitum.
In a chickadee nest two cowbirds were found both of which were in-
fested with hippoboscid flies. Attached to the abdomen of one of
the flies were two mallophagan bird lice that thus obtain transporta-
tion from one bird to another, and within the bodies of the lice bac-
teria were undoubtedly present (Herman, 1937). As many as five
links in chains of such hyperparasites and symbionts have been re-
ported.
During the perpetual war between parasites and their hosts many
special adaptations
have evolved on both sides. The anti-invasion
tactics of the host include external anatomical features and internal
defenses of antibodies and phagocytes. The parasite is

usually pro-
vided with a protective covering and with special ways for gaining
entrance, maintaining position, and avoiding digestion by the host.
On the other hand, certain of the parasite's unused organs have de-
generated. The efficient
parasite gains all the advantage possible
without seriously curtailing the life of the host. If the host is killed,
or when it eventually dies, the parasite must have the means of reach-
ing a new host. Both host and parasite may become so profoundly
modified as to be almost unrecognizable. In its young free-swimming
stage the barnacle Sacculina has the appearance of a typical crustacean
larva, but, as an adult parasitizing the shore crab, it becomes little
more than a sac suitable for the absorption of food and for reproduc-
tion. The crab host is in turn reciprocally affected by the presence of
the parasite for its reproductive glands are caused to atrophy and cer-
tain metabolic changes are brought about.
In many instances the transmission of the parasite to new hosts is
left to chance aided
by a very high natality; in others the parasite
oriented to the host (Thorpe and
Jones, 1937) or the
is
specifically
host attracted to the parasite. A remarkable instance of the latter
is
procedure is displayed by the fresh-water mussel Lampsilis ventricosa

in which the mantle edge is modified to appear like a small fish.
When a real fish, attracted by this mimic, swims over the mussel, cast-
ing a shadow, the mussel discharges its glochidial larvae. Some of

these larvae reach the gills or fins of the fish to which they attach and
live as parasites until
they are ready to metamorphose into adults.
Certain fishes thus parasitized wander upstream where the young
mussels drop off and begin a new life as independent bottom ani-
mals. In this way these sessile forms are distributed against the
current to the upper reaches of the stream.
Many parasites require more than one host for the completion of their
life
cycles; a large number
of instances could be cited for both plants and
animals. One familiar example is the white pine rust that is de-
pendent upon members of the genus Ribes as a secondary host. The

bass tapeworm that causes stunting and sterility in the small-mouthed
bass also illustrates the complexities of multiple parasitism. In the
spring when the bass swims into the. shoal water of a lake for spawn-
ing, segments of the tapeworm (1 in Fig. 10.12) living in the fish's
gut are discharged into the water where they produce eggs ( 2 ) The .
eggs are eaten by copepods (3), a secondary host, and hatch out
within the alimentary tract of these primitive crustaceans. The re-
sulting larvae pass through the wall

of the intestines into the
body
cavity of this host. Meanwhile young bass and other fish (4)
Parasitism 391
we;*
FIG. 10.12. Life cycle of the bass tapeworm (Proteocephalus ambloplitis, in-
the copepod Cyclops, and
volving adult small-mouthed bass Microptcrus dolomieu,
young bass or other small fish. (Hunter and Hunter, 1929, New York State
Conservation Dcpt. )
spawned same shore area have grown to the feeding stage and
in the
eat the copepods harboring the larvae of the tapeworm, A close cor-
relation is thus required between the time for the copepod popula-
tion to develop, for the larvae of the parasite to hatch, and for the
young fish to
appear. The
larvae taken in with the copepods can
resist being digested, as can the other stages of the parasite, and make
their way into the body cavity, liver, or other organs of the small
fish which is a tertiary host. When the infected young fishes are
eaten by the voracious larger bass (5), the tapeworm larvae are
passed on, and this transfer from smaller fish to larger fish may take
place several times. The larvae do not develop into adults while in
the body cavity or internal glands of the small fish. Only if the in-
fected fish is eaten by a large bass at the moment when the larvae
are ready to metamorphose, do they develop into adult tapeworms.
This final stage establishes itself in the intestine of the primary host
and thus completes the life
cycle. In another species of fish tape-
worm the adult stage is found in man and in other fish-eating mam-
mals. The cycles of flukes that parasitize birds, snails, and fish
life
successively are described by Hunter (1942). These multiple rela-

tions involve many special adaptations of the parasites and of the
species in the environment on which they depend.

Predation. The typical predator is free living and catches, kills,
and devours individuals of another species for food, in contrast to the
typical parasite that lives on or in its host and derives nourishment
from it without killing it.y Thus the predator is said to live on "capi-
tal" whereas the parasite liveson "interest." ^/Îthough this concept
is
generally true as far as the individual aggressors and victims are
concerned, it does not necessarily apply to the populations as a whole
Parasites may kill a sufficient number of their hosts so that their
"capi-
tal stock" is
very greatly reduced. For example, in the grouse dis-
ease, which game bird periodically, the host population is

strikes this
so depleted by the ravages of the infectious parasite that the parasite
population in turn sinks to a low ebb. Similarly, in an area in which

the predator and prey populations have struck more or less of a bal-
ance we may find that the predators are limiting themselves to "in-
terest" in the sense that they are
devouring only the increment to the
prey population each year. In such a situation the predator
popula-
tion may continue indefinitely to take a limited number of the
prey
without endangering the breeding stock of the species on which it
depends.
Predatory organisms are almost entirely animals but a few kinds
Predation 393
of plants are also included.

Predatory animals that eat other animals
are carnivores, and those that feed on
plants are herbivores. Some
herbivorous animals kill the
plants on which they feed by consuming
all most of each individual.Âquatic filter feeders necessarily de-
or
stroy all the diatoms, flagellates, and other algae that they digest,
and herbivorous animals on land that feed on small plants may do the
same. However, many insects and many ruminants browse lightly
over the vegetation in such a way as to allow the plants to continue
life
indefinitely. We have seen that the grazing of sheep, for ex-
ample may trim the grass harmlessly and prevent the invasion of
other plants, thus aiding in the development of a healthy permanent
turf.
Usually plants serve as food for animals, but in a very few excep-
tional instances the tables are turned, and animals fall
prey to car-
nivorous plants. ^ln the terrestrial flora these are also known as in-
sectivorous plants since insects are the usual prey on land. Plants
with this food habit are adapted in various remarkable and intriguing
ways to attract, catch, and digest their victims. The sundew
'
Drosera ) for example, has round, reddish leaves provided with hairs
that are progressively longer toward the periphery and tipped with
glistening drops of a sticky, sweet secretion. When an insect, at-

tracted by the color or odor of the plant, alights on the leaf, it sticks
fast to this little bit of natural fly paper. The presence of the insect
causes the hairs to bend over it; a digestive secretion produced, and
is
the body of the insect absorbed>/The

is vase-like leaves of the
pitcher plant are filled with water and serve

as death traps for other
insects (Fig. 10.13). Animals that fall into the pitchers are unable
to climb out because of the steep sides and downward directed hairs;
they eventually drown and are digested by special tissues at the base
of the leaf. Further interspecific relations result from the fact that
several protozoans and aquatic
insects, including certain mosquito
larvae, normally the

livewater
in of the pitchers. In the larger
plants of the tropics frogs station themselves

at the
openings of the
pitchers and spiders spin webs across them to secure their take of
the insects that stray past.

Some plants are hunters of even smaller game. Specially adapted
bladders of the aquatic bladderwort Utricularia open suddenly wher
any plankton organism comes in contact with the trigger bristles

the unlucky plankter is sucked inside and digested. Equally remark-
able are adaptations that permit certain predaceous fungi to capture
and to consume prey even larger than themselves. One species pro-
duces a chain of cells in the form of a lasso in which a portion of a
nematode, or other soil microorganism, becomes ensnared and which
then tightens up sufficiently to kill the victim ( Bessey, 1950 ) .
Some predators are restricted to one prey species or are dependent

upon a small group of food species, whereas others are highly catholic
in their tastes, as has already been discussed in Chapter 8. Con-
versely, certain prey species are
attacked by only one predator, but
others may satisfy the appetites of many kinds of diners. The special
anatomical and physiological of predator species for
adaptations
securing, devouring, and digesting specific prey species,
and the
equally elaborate specializations of the prey to
avoid detection or to
resist capture, have been considered in earlier chapters.
I'twto by H. W . Ail red, U. S. Soil (.Conservation Service
FIG. 10.13. A group of pitcher plants (Sarracenia flava) in

swampy land of
Louisiana.
In some animals only the adults are predatory, whereas the young
are parasitic or live wholly upon the yolk supplied in the eggs. In
other animals, such as many insects, the larvae do most
of the eating,
and in certain species the adults do not feed at all. Some predators
feed upon the adult stage of their prey, some on the larvae, some on
the eggs, and some on more than one stage. If members of the prey
species are killed before they have had a chance to reproduce, the
Predation 395
inroads of the predators on the
population will be particularly serious.
The degree to which food "preferences" influence the feeding of
predators of considerable importance, both theoretically and prac-
is
tically in relation to pest control,

game management, and fishery
biology. The forage ratio of a prey species (A) in respect to a
predator species (B) is defined as the ratio of the percentage of B's
food made up by A to the percentage of the potential food organisms
in the environment represented by A:
Horace ratio =
.
%A in /Ts
7^7 /-. r
food
%A in environment
A forage ratio of 1 indicates that the species is eaten in the same pro-
portion as its abundance in the habitat. An investigation of the

feeding of the black-nose dace (Rhinichthys atratulus atratulus], for
example, revealed a forage ratio of 2.7 for Diptera and of 0.47 for
Trichoptera although the latter were about one quarter more abun-
dant in the stream. In this instance the Diptera were found to be of
size and in positions that rendered them more accessible to the fish
than were the Trichoptera (Hess and Swartz, 1940).

Generally speaking, predators tend to eat the organisms that are
most available either because of their abundance or because of their
accessibility. predator has a specific food preference or becomes

If a
conditioned to continue feeding on the same species, serious conse-
quences may ensue both for the predator and the prey; but usually
when one type of food becomes scarce, the predator changes its diet.
Top minnows (Gambusia) are often stocked in reservoirs because
fishes of this type are known to eat mosquito larvae voraciously. Un-
fortunately for the mosquito-pestered human residents, when the fish
have reduced the larvae to low numbers, they turn to other food and
thus allow the mosquito population to recover.
Studies by N. Tinbergen showed that during one winter, within a
certain locality, when voles (Microtus) were abundant, they consti-
tuted 86% of the food of long-eared owls (Asio otus) and wood mice
(Apodemus) furnished 1%\ whereas in the following winter when voles

were scarce, they formed only 30% of the owls' food, wood mice fur-
nished 153S, and other animals made up the difference. A more elabo-
rate investigation of the forage ratios of the sparrow hawk by L.
Tinbergen has been reviewed by Hartley (1947). These quantita-
tive aspects of prey-predator relations will be considered further in the
section on competition and in subsequent chapters dealing with the
interdependences of the community as a whole.

Competition
In the foregoing sections we have
discussed certain categories of
antagonism in which one species harms another by parasitizing it,

by preying upon it, by poisoning it, or by taking some other direct
advantage of it. In this section we shall consider first the aspect of
competition that involves the "cold war" between species that are
contending for "lebensraurn" or for "consumer goods," and then we
shall discuss certain generalities in regard to competition involving
both direct aggression and indirect rivalry.
An organism competes
with members of its own species as well as with representatives of
other species for space, light, food, or other necessity, but the nature
of the competition between species differs because of the variation
in
precise needs and adaptations of different species.,/
Lichens compete with each other for space on a dry ledge and also
with members of other species; on a submerged rock barnacles simi-
compete for space with one another and also with oysters, mus-
larly
and other sessile animals. Tree seedlings vie with small shrubs
sels,
and herbs for light in the developing vegetation. The roots of a
forest tree engage in a continuing but unseen struggle with the roots
of other trees for water and for nutrients. Various species of para-
sites contend for the choicest tissues of their host. Grasshoppers not
only compete closely with other insects for grass but also contend to
some extent with mice and rabbits as well as with sheep and antelope
for the same food. Various carnivorous species similarly are often
rivals for the same prey.
The moreclosely similarone organism is to another the more nearly
alike will be their needs and hence the more intense will be their
rivalry in obtaining their requirements
from a common environment.
This fact means not only that competition between individuals of the
same species is particularly keen but also that the intensity of com-
petition between species is directly related to
the ecological similarity
between them. The rivalry between species of the same genus is
therefore usually more severe than that between species belonging to
different genera as pointed out long ago by Darwin.
Various aspects of interspecific competition have been investigated
with carefully controlled laboratory populations of flour beetles.
When Tribolium confusum and Tribolium castaneum, for example,
are grown in the same "universe" of flour, one species always becomes
extinct, leaving the other in sole possession, although plenty of food
is available for both. The two beetles are very closely similar in life
Competition 397
history, behavior, and requirements, but subtle differences exist in the

responses to the conditioning of the medium, in the effects of crowd-
ing on natality, mortality, and rate of development, or in the rates at
which the beetles eat each other's eggs. Under certain conditions
T. confustwi was found to inhibit the net
fecundity of T. castaneum
more than the latter species inhibits itself, with the result that T.
confusum drove out T. castancum in competition. Under slightly
different circumstances castaneum gained the upper hand, but
T.
never would the two species coexist permanently in the same unit of
flour (Birch, Park, and Frank, 1951).
In similar tests involving mixed cultures of two species of clado-
cerans, Daphnia pulicaria always caused the extinction of Sirno-
cephalus vetulus. Since these species do not directly attack one an-
other at any stage, the displacement of one species must be due to
difference in toleration for shortage of food, chemical conditioning,
oxygen lack, or physical effects of crowding, but precisely what is the

crucial aspect of the competition awaits further investigation (Frank,
1952).
From both laboratory and field studies we learn that two species
having essentially the same requirements from their immediate en-
vironment do not usually form mixed steady-state populations. The
most closely related species of a genus generally have different geo-
graphical ranges. If they live in the same region, they inhabit dif-
ferent types of habitat or they obtain their food and other necessities
in a in other words, they occupy different
slightly different way
habitat niches, as discussed more fully in Chapter 13.
When two closely competing species are unable to continue living
in the same
habitat, the determination of the species destined to
survive depends upon which species is favored by the existing en-
vironmental conditions. If conditions change, as they do regularly
with the seasons in temperate regions, for example, a different species
may come to be favored. When conditions are not optimal in re-
one ecological factor, the is often reduced
spect to range of tolerance
with respect to other factors. These general relations help to explain
the fact that different species succeed one another during the season,
as is seen in phytoplankton populations, with no one species holding
numerical superiority throughout the year.
The type of competition between species that vie with one another
by direct aggression has been given mathematical formulation by
Volterra (
1931 ) and Lotka (
1934 ) for various cases in which one or
more species feeds upon another species. When the population of
one species grows at the expense of another, as in the predator-prey
relation, the populations of the two species have certain reciprocal

relations. These may be illustrated by the well-known investigations
of Cause (1935) which the interactions were demonstrated in
in
laboratory populations of protozoans (Fig. 10.14). Cause prepared

cultures in which the ciliate Paramecium caudatwn, as prey, was
eaten by another ciliate Didinium nasutum, as predator. An ample
Predator
Predator
III
Time
FIG. 10.14. History of prey (Paramecium caudatum) and predator (Didinium
nasutum) populations in (I) homogeneous microcosm with initial seeding, (II)
heterogenous microcosm providing refuge for prey with initial seeding, and (III)
homogeneous microcosm with repeated seedings. (From Alice et al., 1949, after
Cause. )
bacterial population was present at all times as a source of food for

the paramecia. In the first
type of experiment Paramecium and
Didinium were introduced into a medium
equally available to both;
in the second type the microcosm was heterogeneous so that the
Paramecium could escape into one part of the medium inaccessible to
its
aggressor. A homogeneous microcosm was
again used in the third
type of experiment, but additional seedings of both species were in-

Competition 399
troduced at regular intervals. In the first ?et of tests the prey popula-
tion at first
rapidly, but, as the number of predators increased,
grew
the prey gradually became extinct and thus caused the starvation of
the predators soon afterward (Fig. 10.14, Case I). In the second ex-
periments ( Case II ) since the prey could escape, their population con-
tinued to grow, but the predators died off. In the third set of tests
(Case III), in which the additional seedings simulated immigration,
oscillations of both populations were induced with the maxima and
minima of the predator population following those of the prey
population.
Many illustrations of the interdependencies of prey and predator
shown in the foregoing tests are known in natural populations. The
removal of predators has allowed certain useful fish populations to in-
crease several fold (Huntsman, 1938; Foerster and Ricker, 1941).
The protection of deer from predation in the Kaibab forest resulted
first in their great increase and then in wholesale death from starva-
tion, as discussed in an earlier chapter (Leopold, 1943). In most

natural situations the predators are unable to find and kill all the
prey so that, after the reduction of the predator population, the prf^
population can increase once more. Furthermore, since in nature
not all the prey are successful in finding refuge from their enemies, a
small part of the predator population usually manages to survive
periods of prey scarcity.

The action of these reciprocating influences may cause oscillations
in natural populations of prey and predator even in the absence of
periodic immigration. For example, in the Hudson Bay watershed

the populations of lynx and of its prey, the varying hare, fluctuate
widely and with about the same period of approximately 9% years

(MacLulich, 1937). From the fact that the maxima and minima in
the lynx population follow closely those of the hare population, as

well as from other observations, the periodicity of lynx abundance
appears definitely to be controlled by the availability of its food, and

the oscillations of the hare population are influenced to some extent,
at least, by the extent of lynx predation. Host and parasite popula-
tions similarly affect one another in a generally reciprocal manner
(Debach and Smith, 1947, and Utida, 1950). The repercussions of

prey-predator and host-parasite fluctuations on the whole community
will be considered further in Chapter 12.
To summarize, when two antagonistic species occur in the same
environment, these general types of result are possible: (1) If the
two species are ecological equivalents making common demand on
the environment, and if one inhibits the growth of the other more
than its own

growth, the first species will cause the elimination of
the second from the area. (2) If the two species have somewhat
demands on the environment so that they inhabit different
different
niches, thetwo species may continue to coexist in the area. (3) If
the two species are dependent one upon the other, as parasite and
host or predator and prey, the aggressor may eliminate the other
species and then turn to other types of food. (4) If the attacking
species is unable to
destroy all the other
species and is able to survive
periods of low abundance of the other species, the two species may
continue to live in the area in either a steady or a fluctuating equi-
librium. For a further discussion of these intricate relations of com-
petition the reader may consult Hutchinson and Deevey (1939),
Solomon (1949), or Allee et al. (1949, Ch. 22).
The survey been presented in this chapter
of interrelations that has
has revealed the extraordinarily varied and complex nature of the
dealings of one species with another. We have delineated relations
of mutual assistance, of toleration, and of antagonism; we have re-
viewed the special circumstances of parasitism and of predation and
have considered various ecological effects of competition between
specific types of organisms.
In the natural community many indi-
viduals of many species of both plants and animals are typically
It is our next task to consider the
present. operation, organization,
and alteration of the community as a whole in which each individual
is
responding not only to influences from all neighboring species but
also to concomitant influences from others of its own species and from
the chemical and physical features of the environment.
Community Dominance 409
example inan alpine region, or is largely unmodifiable as in the

open
ocean. This type of condition is also seen in communities of marine-
bottom organisms in which the chief controlling influences are the
mud of the bottom and the water currents. These factors act on the
species presentmore or less
directly, as indicated by the accompany-
ing scheme (case 1).
Mud-Bottom Biocenose
Water currents
/ / / /
Case (1) A B C B
I t t t
- Substratum
Case (2) A -**B *-(>* B
Case (3) A +B *-C-<- B

(Q (C) (C)
In addition to the major effect of the physical features of the en-

vironment a certain amount of interdependency may also exist among
the members of biocenoses of this sort. In communities living on
isolated rocks the interrelations between the species may be quite
minor or entirely absent. When interdependencies do occur, the
relations may be specific or non-specific. In case (2) species C is
primarily controlled by the physical features of the environment, but

it is also influenced
by the presence of A and B specifically. An illus-
tration of this situation is furnished by the fresh-water mussels that
make use of fish past as transport agents for their larvae.
swimming
During most of their lives the primary dependencies of the mussels
are concerned with the bottom, the water, and the plankton of their
biotope, but at the time of reproduction this particular relation with

the fish of their community becomes significant.
In a third type of situation the presence of the other species exerts a
definite effect but not in a specific way. In other words, as far as C
is concerned, A and B may merely be occupying space, and individ-
uals of C
could be substituted for them without significant conse-
quence to C as is suggested in case ( 3 ) above. Illustrations are found
in the competition for space among attached animals on submerged
rocks. Similarly, in the competition
for light
among plants the
proximity of other individuals often has its chief effect in
cutting off
410 The Community
the light with little difference whether the interfering neighbors are
of the same or of different species,
In many communities one species is particularly conspicuous
because the largest or the most numerous. This species is often
it is
called the dominant, and its name is usually given to the community,
as in a spruce forest community. Occasionally two or more species
share the honors in respect to dominance, as in an oak-hickory forest.
Very frequently the species that is dominant in conspicuousness also
appears to exert a controlling influence over the other members of the
community, but sometimes a less conspicuous species may have a pre-

ponderant influence on the inhabitants of the biotope. In a sphag-
num-heath community, for example, the heath plants are the most
conspicuous, but the presence of the sphagnum controls the nature of

the community as a whole. In some biocenoses microorganisms may
play the critical part although they are among the least conspicuous
of the species present. The seedlings of pine and some other trees
do not grow properly unless suitable fungi are present to form mycor-
rhizae on their roots. Thus, although the species designated as the
dominant frequently does exert the major controlling influence on
the biocenose, this is not necessarily true even in those communities
in which a high degree of interspecies dependence exists.
In the second general type of situation, in which the members of

the community owe their presence primarily to a more or less direct
and independent control by physical features of the environment,
a species designated as dominant because of prominence or abun-
dance clearly does not exert a critical control over the other inhabit-
ants. For this reason some ecologists refer to such a species as "pri-
mary" rather than dominant. Dominance based on controlling in-
fluence must be clearly distinguished from dominance based on mere
conspicuousness. In communities of low interspecies dependence the

relative numbers and prominence of the various species are likely to
vary from place to place. Great care must be taken in applying

names to such communities to avoid ambiguity and the implication
that conspicuous species are necessarily controlling.
Ecotone
The transition zone of tension between communities presents a

situation of special ecological interest and is known as an ecotone.
The border between forest and grassland, the bank of a stream run-
ning through a meadow, or the boundaries between any other com-
munities on land or in the water furnish illustrations of ecotones
Ecotone 411
(Fig. 11.4). In
this transition zone of tension the
outposts of each
community are maintaining themselves in environments that are in-
creasingly unfavorable. The tension may result chiefly from a strug-

gle with physical conditions or from a direct competition between
certain species in each community. At the border between a shrub
community and a marsh, for instance, the shrubs may compete
directly with marsh reeds for light, nutrients, or other necessities of
FIG. 11.4. A distinct ecotone dominated by smooth sumac (Rhus glabra) and
goldenrod (Solidago) between forest and grass communities in eastern Massachu-
setts.
life in such a way that one type of plant gives way completely to the
other. In such a situation and in areas where controlling physical
factorschange rapidly the transition between communities is abrupt
and the ecotone is correspondingly narrow. In other circumstances
the two communities may interdigitate to a considerable extent. At
the edge of a forest individual trees may pioneer into a scrub com-
munity, and the scrub species will invade the margin of the forest as
far as they are able to survive.
When one community gives way only gradually to the other com-
munity, wide ecotone results/ The transition zones sometimes
a
more than 100 km in width between major continental communities
are regarded by some as ecotones (Pitelka, 1941), but others restrict
the term to areas of smaller scale. Strictly speaking a transition
412 The Community
area is tension exists between the bordering com-
an ecotone only if
munities, and this often

difficult to demonstrate, especially for
is
large areas. Accordingly, the decision as to what is to be considered

an ecotone depends upon the scope of the biocenose as recognized by
the individual ecologist.
In the ecotone area the conditions of temperature, moisture, light,
wind, and other physical influences are different from, and usually
intermediate between, those existing well within either of the border-
ing communities. These or other conditions, such as food or shelter,

may be superior in the region of the ecotone for certain species.
These special influences of the ecotone area prevail below ground and
high in the vegetation as well as on the surface of the soil. As a
consequence various kinds of plants and animals not occurring, or rela-
tively rare, in the bordering communities may
become abundant in
the ecotone. Shrubs typically grow in profusion at the forest edge
and harbor a distinct fauna. At the margin of a pond willows and
cattails thrive in the transition between land and water, and here
are found such animals as turtles, frogs, herons, red-winged black-

and a host of invertebrates that are entirely absent
birds, muskrats,
or much less abundant in the center of the pond or in the terrain far
removed from the water.
These species favored by the special conditions of the ecotone join
with the outpost representatives of the principal inhabitants of the
bordering communities in populating the ecotone. The species oc-
curring in the ecotone may thus form a distinct functional community
of their own. It must be remembered, however, that the ecotone
inhabitants owe their existence to the presence of the particular con-
ditionson each side and that the ecotone assemblage would disappear,
or be considerably modified, if the bordering communities or condi-
tions were removed or seriously changed.
As a rule the ecotone contains more species and often a denser
population than either of the neighboring communities, and this gen-
erality is known
as the principle of edges. The greater variety of
plants in the ecotone provides more cover and food, and thus a greater
number of animals can be supported. In measurements of bird
populations in a variety of areas in the central part of the United
States Kendeigh (1944) and Johnson (1947) distinguished between
"forest-edge" species and species which confine their territory, nest-
ing, feeding, and roosting to the interior of the forest.

As conservationists and wildlife managers have become aware of
the principle of edges, they have bent their efforts toward increasing
the amount of available ecotone area in each region since this usually
Ecotone 413
A: Poor interspersiond covey)
Cultivation
B: Good interspersion (6 coveys)
FIG. 11.5. Diagram indicating the increase in number of coveys of quail sup-
ported by farmland after increase in the amount of interspersion of the vegetation
without change in the total area of each. (Reproduced from Game Management
by Aldo Leopold, Copyright 1933 by Charles Scribner's Sons, used by permission
of the publishers.)
414 The Community
favors the increase in game birds and mammals. Regions that are
broken up into many units of different vegetation types are said to
have "good interspersion" because the total length of community edge
is
great and hence a large amount of ecotone habitat is provided.
Since coveys of quail tend to establish themselves where several
types of vegetation meet, the number of coveys supported by farmland

can be increased through a change from a few big tracts of woods,
brush, pasture, and fields to a greater number of smaller interspersed
tracts without altering the total area of any one type of vegetation
(Fig. 11.5).
COMMUNITY COMPOSITION
Communities inhabiting distinctive and commonly occurring bio-
topes are the most easily recognized, and their composition and
interrelations have been more extensively studied than others. A
biocenose of this kind is made up of a characteristic, but flexible,
assemblage of species without necessarily containing any species that

is exclusive to it. Certain types of species invariably occur in a
biocenose of a certain kind, but the individual species may vary
widely from place to' place. In a prairie community grass-like plants
are always present and serve as the fundamental plant producers,
but in different regions quite different species of grass fill this niche.
Among the animals of this community fleet browsing forms, jumping
types, and burrowers are commonly present but the species, genera,
or even families of these typical members of the biocenose may differ
greatly, depending upon the region in which they are located. The
ecological relationships are entirely separate from the taxonomic af-
finities ofthe species since the latter depend upon the evolutionary
history of the area. Species having a common evolutionary origin are

placed in "faunal regions" or "floral regions," and these subdivisions
may or may not correspond with ecological subdivisions.
Communities may be large or small. Some may cover thousands
of square kilometers, such as a spruce forest in Canada, a prairie
community in central North America, a pine-land community in

southeastern United States, or an oceanic community. Other biocen-
oses, such as those occupying relatively uniform swamp, desert, or
lake biotopes, may have dimensions in hundreds of kilometers. The
biocenoses of ponds, tide flats, rivers, balds, alluvial sands, chaparral,
mountain meadows, and rocky plateaux typically occupy a more re-
stricted area. Smaller communities inhabit canyons, Tiammocks" in
the everglades, mountain springs, tidal inlets, and clearings in the
forest. The plants and animals living on an isolated boulder, or in a
rotten log, or in the debris washed
up on the shore of a lake constitute
smaller biocenoses. Characteristic communities
still
occupy even
more minute inicrohubituts in the area such as a crevice in a ledge,
or the water in a pitcher plant. We thus have communities within
communities. Theepiphytes, cpizoans, and parasites of an animal
or a plant may also be thought of as
comprising a biocenose. An
apple tree and the plants and animals that live on it and in it illustrate
a special community of this sort, A horseshoe crab with the dozen
or so species of mollusks, worms, and barnacles attached to its shell
exemplify a microcommunity, as do the fauna and flora of a cow's

stomach.
The number of species and the population abundance in communi-
ties also
vary greatly. While maintaining the necessary quantitative
relationships among the consumers and the producers in the eco-
system, the inhabitants of an area often include a wide range of densi-
ties. Furthermore, each component in the ecosystem may be repre-
sented by one, a few, or many species. Biotopes with extreme con-
ditions and little food, such as a rock desert or an ocean deep, gen-
erally support few species and relatively few individuals of each

species. The relatively small number of both species and individuals
in the badlands sections of southwestern United States is familiar to
those who have crossed these areas. In other severe habitats suffi-
cient energy and nutritive materials are sometimes available to allow
huge populations to develop although only a few species are able to

survive. In the relatively simple biocenose on Bear Island in the
Arctic Zone some of the smaller plants and animals are very abun-
dant (Elton, 1939).
Under more equable climates larger numbers of species generally
become interrelated in community groups; each species is usually
rather meagerly represented, but sometimes the abundance of certain
species is great. The food relations of

the North Sea herring are
known to involve a great number of species (Fig. 11.6) some rep-
resented by very large populations and these food species form
of which the herring is a
only a portion of the whole community
member. A bed of kelp (Nereocystis luetkeana) in Carmel Bay,
California, was found by Andrews (1945) to form the matrix of a

community that included 40 species of invertebrates, several kinds
of fishes, and numerous attached algae, protozoans, and bacteria, as
well as associated species of phytoplankton and zooplankton.
Few complete enumerations have been made of both the plants and
animals in even the simplest situations and none in complicated
in those with elaborate populations of micro-
biotopes especially
416
organisms. The complex ecosystems of temperate and tropical ter-

restrial areas are
particularly difficult to analyze, but some idea of the
number and diversity of the living components may be obtained from
existing studies of portions of the biota. A comparison of the num-
bers of species that may be involved in communities of different kinds
has been made by Elton (1946); a few
examples follow.
The biocenoses of certain British rivers include representatives
of as many 131 species of invertebrates in addition to fishes,
as
amphibians, rooted aquatic plants, and vast populations of algae,

flagellates, and bacteria that were not enumerated. A meadow on clay
near Oxford, England, served as the biotope for 93 species of inverte-
brate animals in the soil and surface vegetation, in addition to the
plants and the microorganisms. At the other numerical extreme, the

biocenose on the sandy shore of Amerdloq Fjord, west Greenland, con-
sisted of only 5 species of invertebrates. In the Bettila odorata forest
in Finmark, Norway, only 29 species were reported, whereas
of plants
in an ash forest in Yorkshire, England, 72 species were found, and in
the red fir forest of the Sierra Nevada, California, 93 species of plants
were recorded. The animal components of these forest communities
were not recorded, but they would include birds, mammals, am-
phibians, and a great many species of insects and other invertebrates;
the total number of species would be further swelled by a wide
variety of protozoans and cryptogamic plants. Even within a single

microhabitat a surprising number of animals may be present as evi-
denced by counts of 111 species of invertebrates found in pine logs
and 136 species found in oak logs during investigations in the Duke
Forest, North Carolina.
Another observation in regard to community composition is that
the species present for the most part belong to different genera. In
many biocenoses each genus is represented by only one species; in

others certain genera are represented by several species, as is often
true, for example, of the sedges, willows, and oaks in communities
of temperate and boreal North America. Rarely are more than a few
of a found in the same biocenose. Quite exceptional
species genus
is the occurrence of flocks," such as those in Lake Baikal,
"species
where, for example, 300 species of gammarid crustaceans are found
and these belong to only 30 genera (Brooks, 1950). The typical
biocenose is
species that are separated by at least generic
composed of
differences, whereas in the biota of a country or a region many species
may be recorded for each genus. Comprehensive surveys of the

animals in 55 communities and of the plants in 27 communities from
showed that 86 per cent of the animal genera
a wide range of habitats
418 The Community
and 84 per cent of the plant genera were represented by only one
species (Elton, 1946). The average number of species per genus
in these
widely spread surveys was 1.38 and 1.22 for the animals
and plants, respectively. In contrast the average number of species
per genus for all British insects, for

example, is 4.23.
As a general rule the species of a genus are sufficiently similar so
that their demands on the environment are in serious conflict. During
the course of evolution this conflict has usually resulted in eliminating
all but one
species from each genus represented in the area. In
contrast to a geographical region, a community is a functional unit
and each niche in the community structure is occupied by that species
which has been most successful in the competition for it. Adaptations
for ecological needsand accessibility determine which species will
survive and populate each part of the biocenose. The species that
succeed in establishing themselves are almost always sufficiently dif-
ferent to be members of separate genera. Competition within each
level of nourishment in the food chain thus produces a certain or-
ganization among the species of a community in addition to the

organization imposed by the relationships from one link in the food
chain to another and by other dependencies. The species composi-
tion of each biocenose is therefore determined
by relations within
the same functional level as well as in successive levels from among
the species that are distributed to the area.
To make a census of the inhabitants of a diversified natural com-
munity and to ascertain the role of the various species in any compli-
cated ecosystem exceedingly difficult. Such tasks require the
is
combined efforts of at least several taxonomists and ecologists com-

petent to deal with the various segments of the biota. General ac-
counts of the "plant communities" and the "animal communities" in-
habiting the principal types of environment in the world are available

in such works as Costing (1948), Hesse, Allee and Schmidt (1951),
and Dice (1952). Detailed studies of the inhabitants of specific
biotopes have for the most part been carried out by investigators con-
fining themselves to restricted categories of
animal or plant life. To
obtain an idea of the total assemblage of organisms making up a
biocenose and their integrated activities, it is usually necessary to
piece together the results of several investigations carried out in the

same region. The reader may obtain rather complete information
on the composition of the entire biota of certain specific areas by
reference to the collaborative reports of members of expeditions, of
government surveys, and of field research laboratories. In many

instances the work of areal surveys has been expanded to include
MATURE POPUP COMMUNITY
Baltimore oriole Canker Hairy and Downy
Chickadee Pomes woodpeckers
Least flycatcher
Rose-bstd grosbeak
Dicera
Willow thrush
Saperda
Ruffed grouse
Cooper and
t
Groshawk
\N
Sharpshinned
.Rabbit
hawks Crow.
V\ Great horned owl
FOREST EDGE
IMMATURE ASPEN COMMUNITY
Yellow warbler
Redwinged blackbird
Bronze grackle
ispen
Show-
berry
Hazelnut
Choke-
Red-eyed
vireo cherry
Yellow
warbler
Gold finch
Catbird
Brown
thrasher
Townee
robin
Cutworms
Grasshoppers
Click beetles
Pocket gophers
Ground squirrels
FIG. 11.7.Simplified representation

communities
of four in the parkland region
of Manitoba, showing some of the food relations within and between communities.
and Schmidt, Ecological Animal
(Reprinted with permission from Hesse, Allee,
Geography, 1951, John Wiley & Sons.)
419
420 The Community
studies of the dynamic ecological relationships within the community.
The extensive series of investigations on Wisconsin lakes will serve as
an example (Juday, 1943).
The foregoing discussion has shown that the biocenose is
composed
of species characteristic of the area, but these species may not be
limited to the area nor is the biocenose limited to a fixed set of species.
This concept of the flexibility of the biocenose is well illustrated by
studies made by Bird (1930) of the interrelationships of members of
communities occurring in the parkland region of Manitoba. The
chief food relations of certain of the species within and between com-
munities are indicated in Fig. 11.7. A great many other, smaller
species are present in each community represented in the diagram,
and many other dependencies exist besides the food relationships.
This oversimplified picture nevertheless shows that certain kinds of
plants and animals form a characteristic nucleus in each community
but many of the individual species occur in more than one biocenose.
The principal members of each biocenose have most of their require-
fulfilled within their own
ments biotope but interrelations between the
biocenoses are brought about by animals that move from one biotope
to another for foraging or other purposes.
STRATIFICATION OF THE COMMUNITY
Many communities exhibit a structure, or recognizable pattern, in

the spatial arrangement of their members. A community may be
divisible horizontally into "subcommunities" that is, units of homo-
geneous life-form and ecological relation. The zonations described

in earlier pages of this chapter are horizontal structural units of this
sort. Of more general occurrence is the
aspect of structure that in-
volves vertical changes, or stratification, within the community. In
some communities a complex stratification is present, but in others the
vertical dimension is so much compressed that the entire biocenose
consists essentially of only one stratum. Lichens pioneering on a
rock ledge represent an extreme example of a one-layered community,
but, at a later stage when higher-growing mosses and herbs have be-
come established and bacteria are present in a layer of humus beneath
the holophytic plants, the beginnings of stratification are apparent,
albeit on a very small scale.
In a grassland community subterranean, floor, and herbaceous sub-
divisions canbe recognized. The subterranean stratum contains the
roots of the principal vegetation and forms the permanent residence
of the soil bacteria, fungi, and protozoans, as well as a host of insects,
spiders, worms, and other invertebrates.In addition, it provides a

part-time abode for many other animals including other species of
insects, rodents, reptiles, mammals, and a few burrowing birds. The
main activity in the soil for both plants and animals occurs in the
upper layers, but the longer roots of prairie
grasses extend to depths
of about 2 meters and the
roots of other prairie plants occasionally
reach levels of 5 or 6 meters below the surface. In an extensive
plant parts Weaver and

discussion of the relation of underground
Clements (1938) report that 65 per cent of a group of true prairie
species studied have root systems that penetrate to depths between
1.5 and 6 m. Prairie ants are active to 3 m, and prairie dogs are
known to burrow to more than 4 m.
The floor subdivision of the grassland community contains basal
portions of the vegetation, including particularly the rhizomes of the
grass plants partially covered by litter and debris of both animal and
plant origin. A
characteristic group of animals in which insects,
spiders, reptiles,and rodents are prominent join with the plants to

form this subdivision of the community. Most of the animals inhabit-
ing the grassland floor also usually invade one or both of the other
subdivisions. The herbaceous stratum of this biocenose consists of
the upper parts of the grasses and herbs and a characteristic assem-
blage of animals. The vertical dimension of this subdivision is vari-

able, according to the plant species and the local conditions. The
area may be covered by sparse, resetted grass only a few centimeters
in height or by coarse tough species growing 2 or even 3 high. m
The animals of the herbaceous subdivision include a wide variety of
insects, birds, and ruminants.
Stratification of land communities reaches its
greatest complexity in
the forest. Five vertical subdivisions of the forest biocenose are
typically present, namely: the
(1) subterranean, (2) forest-floor,
(3) herbaceous, (4) shrub, and (5) tree strata (Fig. 11.8). Each
of these may exhibit certain further subdivisions in particular situa-
tions, and the air above the forest canopy is sometimes considered to
comprise a recognizable division of the community. Allee (1926)
distinguished 8 strata in the tropical rain forest of Barro
Colorado
Island. The subterranean layer of the forest is typically damp and
contains a large amount of humus that in extreme cases may be
prominent to a depth of 2 or 3 m. The effective depth of the sub-
terranean division of the forest community is difficult to determine
but in wet forests it
appears to be definitely shallower than in dry
prairies. The adequate supply of water near the surface, the avail-
ability of nutrients,
and the inability of roots to penetrate
into poorly
422 The Community
aerated influence the depth of root growth.
soil In the temperate
forest the tree roots may extend to depths of 3 m, but the bulk of
root development occurs in the upper meter. Animal activity is most
intense in the upper half meter but the larger burrowing forms pene-
trate to depths of several meters.
FIG. 11.8. Stratification of vegetation in a deciduous forest of the temperate
region. The vertical subdivisions of the biocenose are: (1) subterranean, (2)
forest-floor, (3) herbaceous, (4) shrub, and (5) tree strata.
The forest floor is a complex subdivision of the biocenose in which

great biological activity goes forward involving the decay of plant
material and a web of predator-prey and parasite-host relationships.
Huge numbers of invertebrates inhabit fallen logs and other material

on the forest floor, and they are preyed upon by carnivorous beetles
and hunting spiders as well as by higher animal types. Saprophytes,
including the familiar mushrooms and bracket fungi, are particularly
characteristic here but mosses and other low growing green plants
may be abundant. In general the density of the population of the

floor stratum in the forest is higher than it is in
grassland and higher in
warmer climates than in colder climates,
The subdivisions of the forest biocenose above the ground are
variable and are determined
by the arrangement of the vegetation,
but in most instances herbaceous, shrub, and tree strata are distin-
guishable. The herbaceous stratum varies in height up to a meter or
so and frequently overlaps with the shrubs that extend to heights of
perhaps 1 to 5 m. The tree stratum occurs between heights of 5 and
15 min the
typical oak forest, but it extends to 25 or 30 in the m
coniferous forest and to about 40 m
in the rain forests, with individual
large-crowned trees towering to 50 m or more. The upper limit of

the forest canopy in groves of giant redwoods may surpass 100 m.
The herbaceous stratum is poorly represented or absent in some in-
stances as, for example, under a thick stand of pines or spruce, and
the shrub stratum may in
vary greatlyprominence. On the other
hand, the tree stratum may be elaborately developed and divisible
into sublayers. The trees of the rain forest on the Gold Coast of
Africa display a stratification that extends from a height of 2 m or so
to about 40 m (Foggie, 1947).
The herbaceous, shrub, and tree strata are each inhabited by a
characteristic assemblage of epiphytes and epizoans. Herbivorous
insects and web spiders are particularly abundant in the herbaceous
stratum. In the higher strata numerical superiority is held by the
insect group in enormous variety but snails, lizards, snakes, frogs,
arboreal mammals, and many kinds of birds are also present in greater
or lesser abundance. Most of the species are primarily associated
with one of the principal strataor perhaps with one of the subdivi-
sions of the tree stratum but individuals of many of the species range
above or below their usual abode.
In certain aquatic habitats vertical gradients in environmental fac-
tors cause a recognizable stratification among the members of the
community. Such layering may have dimensions of less than a meter

in shallow ponds, or it may involve strata many meters thick in the
open ocean. Mention has been made in earlier chapters of the sharp
limits commonly found for the vertical ranges of attached plants and
animals in the zone, and particularly in the tidal zone, as a
littoral
result of the rapid change in water and light conditions. Where these
sessileorganisms exert a controlling influence on a variety of de-
pendent species, they cause subdivisions of the community to be
formed that may be directly observed (Stephenson
and Stephenson,
1949).
The benthic communities at greater depths is not so
stratification of
easily recognized,
but it is
being investigated by means of dredges
and underwater cameras, and by divers with aqualungs. The situa-
424 The Community
tion for communities in the free water contrasts with that for com-
munities on the bottom or on land in the absence of members that are
fixed in position. Since the pelagic plants and animals are free to
drift or to swim vertically as well as horizontally, subdivisions of the
pelagic community are less well defined and are more flexible than
those determined by sessile algae or rooted vegetation. Nevertheless,
critical changes in the
physical environment, as at the lower limit of
the photic zone, at the thermocline, at a density discontinuity, or at
the margin of an oxygenless stratum bring about a definite stratifica-
tion within many pelagic communities.
investigations have been made in marine and fresh-water

Many
environments of the vertical distribution of various taxonomic groups
among the plankton, fishes, and benthic organisms. Summaries may

be found in such reference works as Welch (1952) and Sverdrup
et (1942). However, relatively little study has been undertaken
al.
of the vertical subdivisions of aquatic communities as functional

units. Certain zones of functional dependence are known to exist
at various levels along the bottom and in the free water. One such
unit of interdependence is formed by the plankton and the fishes of
the well-illuminated surface waters, another is composed of the fishes
and invertebrates inhabiting the bottom in deep water, and several
others may be distinguished at intermediate levels. More intensive
study will be required of the interdependencies among all the in-
habitants of each stratum before specific functional subdivisions of
pelagic communities can be clearly delineated.

Although recognizable stratification does not exist in all communi-
ties, its presence is
sufficiently
common in aquatic situations as well as
in terrestrial areas for be considered a general characteristic of

it to
community structure. Spatial organization may thus be added to

the other attributes of the community that have been reviewed in this
chapter. Each community has been shown to have a composition

its members that
and an integration among separate it from neighbor-
ing communities. The degree of dominance within the community
and the nature of the ecotones between these units of plant and animal
life
vary greatly from one situation to another. All these charac-
teristicsallow the community to maintain itself as a recognizable unit
in a specific area for a period of time. Circumstances that sooner or
of one community by another
later may cause the replacement or
within the community will be con-
that may bring about fluctuations
sidered in the next chapter.
12
Succession
and Fluctuation
In the previous chapter the nature of the community was discussed

as a characteristic
group of plants and animals inhabiting an area.
The typical community maintains itself more or less in equilibrium,
but the members of the community are never in complete balance
with each other or with the physical environment. Changes in the
environment over a period of time are produced by variations in
climatic and physiographic influences and also by the activities of the
plant and animal inhabitants themselves. These modifications of the
habitat may cause sufficient changes in the dominant species so that
the existing community is replaced by a new community, or they may
cause marked fluctuations in the abundance of certain species within
the same community. In this chapter we shall consider the progres-
sive changes in communities leading to a relatively stable type of
community, the classification of communities, and the oscillations
within the community.
ECOLOGICAL SUCCESSION
Progressive changes in communities take place from one geological
epoch to another and also within much shorter periods of time. De-
tailed consideration of large-scale changes in the fauna and flora, such
as those caused by the passage of an ice age or the uplifting of a
mountain range, or those resulting from the evolution of new species,
is
beyond the scope of our present task; such alterations in the biota
have great long-term significance, and they are discussed in treat-
ments on paleontology, climatology, biogeography, and evolution
(Shapley, 1953). Here we shall concern ourselves primarily with
the replacement of one community by another in particular areas and
within the same general climatic conditions.
Observation has revealed the fact that in given biotopes certain
425
426 Succession and Fluctuation
communities tend to succeed one another. The occurrence of a rela-
tively definite sequence of communities in an area is known as eco-

logical succession. The change in the communities may be due in
part to independent physiographic such as alteration of drain-
changes
age, erosion, or deposition, but more especially it is caused by modi-
fications produced by the action of each
community on its own en-
vironment. The two types of causes are frequently operating to-
gether, as isseen, for example, in the replacement of a pond com-

munity by a marsh community. The filling of the pond
is
brought
about by the deposition not only of a certain amount of inorganic
but also of a large amount of the organic remains of successive
silt,
communities, and the accumulation of both types of deposit is en-

hanced by the presence of the roots and stems of the living com-
munity members.
The extent to which ecological succession is self-induced as dis-
tinct from being caused by changes imposed from without varies
greatly in different situations. Similarly, the predictability of the
course and speed of succession is variable. In many instances the
presumed course of succession is based on inference derived from

studies of surrounding areas so that "space is substituted for time";
but in other instances, some of which are described below, the nature
of the succession is substantiated by actual records. Self-induced
ecological succession is another outstanding example of the organism

and the environment acting as a reciprocating system.
Living things modify their own habitat so as to cause one com-
munity to give way to another in a variety of ways. All species of
animals and plants tend to increase in numbers and /or in size. The
conditions of the community consequently change because of the
growth of the inhabitants even without any change in species com-

position. Consider a forest, for example. As the trees increase in
size, they provide more shade, higher humidity, and different condi-
tions of foodand cover. New types of animals find suitable conditions
here; old forms may be eliminated. Wildlife managers have come to
realize that the carrying capacity of a forest area for game changes
with time because the availability of food and shelter in a stand of
saplings is
entirely different from that mature
in a stand of trees.
When populations grow in

respect to numbers or size of individuals,
or both, the total weight of living material in the area tends to become
larger. As predators and parasites increase in numbers, they tend to

reduce the abundance of their prey, but as food becomes scarcer the
consumers in turn are curtailed. At the same time the community
Dispersal and Invasion 427
causes changes in the Plants with-
physical nature of its
biotope.
draw material from the they grow and return it when they die,
soil as
but the material returned to the soil is not in the same form. Humus
accumulates, pH changes, moisture content is modified, and other
alterations of the environment discussed in
previous chapters are
brought about.
The changed conditions caused by the varied activities of the in-
habitants of the area may favor the growth of species other than those
that have been dominating the scene. When this occurs, different
species will soon get the upper hand. These may either be species
already present in a subordinate capacity, or invaders from the out-
side. As one or more species take over the dominant position, a new
community will be formed; its establishment constitutes a step in the
ecological succession of the area.
Dispersal and Invasion
The establishment of the pioneer community on a bare area and

the replacement of this community and of subsequent communities
as ecological succession goes forward are dependent in the first in-
stance upon the existence of means by which new species can reach
the area. The ways in which animals and plants can invade new
areas are extremely varied. Certain non-motile forms may be carried
by wind or water currents for great distances; other species ride as

hitchhikers on or in the bodies of various larger animals. The action
of the air and water mediain providing dispersal and the many special
adaptations of eggs, seeds, and adults for transportation have been

described in earlier chapters.
The organism's own locomotion is responsible for the dispersal of
many animals and of some motile plants. The success with which the
starling extended its

permanent breeding range westward in the
United States from point of introduction is indicated in Fig. 12.1.
its
The great distances traveled by ducks after the breeding season and
before undertaking the southward migration to winter grounds have
already been mentioned (Fig. 9.19), Many other species probably

wander radially from the breeding grounds in similar fashion before
going to their winter quarters. Animals with such ample powers
of dispersal can push rapidly into new areas suitable for their
permanent invasion. Although other animals cannot travel at such
speeds as those represented by the flight of birds, their incessant

pressing against their boundaries nevertheless accomplishes wide
dispersal over a period of time. Even non-motile animals and plants

are able to extend their ranges with surprising rapidity when condi-
tions are suitable.
Barriers.
Against this insistant pressure for dispersal, barriers exist
that retard or prevent the movement of certain species. Ecological
SUMMER 1918 SUMMER 1921

WINTER 1916-18 WINTER 1919-21
SUMMER 1926 SUMMER 1932

WINTER 1924-26 WINTER 1930-32
FIG. 12.1. Westward dispersal of the starling as shown by advancing limit of

breeding range (solid line) and winter records (black dots) outside the breeding
range. Star indicates unusually advanced breeding record in 1934. (Kcssel,
1953.)
barriers may be
either physical or biological. The action of many
physical barriers is
easily understood since they may be too wide to
cross all at once and unsuitable as a way station because of local con-
ditions. A great expanse of salt water or of dry land acts as an effec-
tive barrier against the dispersal of fresh-water forms. A rugged
Ecesis 429
mountain range is a barrier preventing the dispersal of plants that
require a warm, moist soil for their growth.
Biological barriers are sometimes more obscure in their mode of
operation, but they may be equally effective in preventing the spread
of certain species. The
action of biological barriers is perhaps most
easily seen whenlarger plants are involved. If an extensive area is
completely occupied by trees, for example, other species of trees with

closely similar ecological requirements cannot enter the area. If the
forested area is
large, the dispersal of the other species will be stopped
unless their seeds can be carried over or around the barrier. In the
same way a grass sod acts as a biological barrier to the dispersal of
other herbs from adjacent areas. Undisturbed prairie grass regularly
prevents the invasion of trees even though in some situations the

climate can "support" trees once the prairie sod is broken. Com-
munities in which available space for certain types of organisms
all
has been occupied are termed closed communities as far as these kinds
of organisms are concerned. Although these communities thus pre-
vent the invasion and the dispersal of certain species, they may allow
species of very different requirements to enter.

Ecesis. In order for new species to invade an area they not only
must have some means of reaching the new locality but also must be
able to grow and reproduce under the conditions found there. Or-
dinarily the first invaders of a bare area are plants, and for them
the
physical features of the soil and the climate are of primary importance
in determining whether or not ecesis, or successful establishment, can
take place. The pioneer assemblage usually of plants first estab-
itself in a new area is sometimes referred to as a colony but
lishing
care should be taken to avoid confusion with the use of the same
term for groups of social animals. After the ecesis of the pioneers
other species will arrive at the area and will gain a foothold if they
can. The presence of certain plants is usually a primary factor in
determining whether or not invading animal species will be able to
establish themselves.Accordingly, the first animals to arrive will be
primarily concerned with whether the vegetation which they
find is
suitable. The general dependence of the distribution of animals

upon that of plants is not limited to the early stages of colonization,
but applies in more advanced communities, especially in the ter-
restrial environment.
Later arrivals at the area being invaded must be able not only to
tolerate the physical conditions but also to compete successfully with
the species already if
presentthey are to establish themselves. The
absence of a species in a given area does not necessarily mean that it
could not live there as far as the climatic and edaphic conditions are
concerned. Finally the newly established species will be forced to
defend themselves against additional invaders that continue to arrive.
The success with which a species can extend into a variety of new
regions depends on its ability to tolerate widely different ecological

influences both physical and biological.
After the invading species becomes established in a new biotope,
it
may undergo genetic changes. This will further complicate or
modify the situation and the effects of such alterations in the inherent
nature of the members of the community must be taken into account,
especially in any study of an area extending over many generations of

the inhabitants. The consequences of mutation and hybridization
and the effects of isolation are topics beyond our present scope and
the reader is referred to specific treatments such as those of Mayr
(1942), Cain (1944), or Anderson (1949).
In summary we observe that from the point of view of the species
a tendency for dispersal is always present, When pioneers reach a
new the struggle
area, a struggle for establishment takes place, and, if
is successful,
genetic modification or evolutionary change may ensue.
Looking at the same process from the point of view of a specific area
we may observe the change of inhabitants as time passes. If we
watched one spot for a long period of time, we would witness a suc-
cession of communities each one formed from new arrivals, allowed
to flourish for a while, and then replaced by a new community.
Succession and Climax
The species that have successfully invaded a biotope dominate the

scene for a period and form a closed community; further arrivals
cannot at first establish themselves. However, in the course of time
conditions become altered with the result that the members of the
existing community no longer compete successfully with

the invaders.
A new dominant type gains a foothold, and a new community suc-
ceeds the old. By the modification of the environment one com-
munity puts itself at a disadvantage and gives way to another; com-

munities appearing at later stages of ecological succession are estab-
lished partly or chiefly because of the modifying action of earlier
communities.
One community continues to follow another until in many situa-
tions a type of community is reached that cannot be displaced under

the prevailing conditions. The community that can maintain itself
Succession and Climax 431
indefinitely in each biotope is known as the climax. In other situa-

tions doubt exists as to whether the communities would ever be un-
disturbed for long enough to enable an
equilibrium condition or
climax to be reached. For example,
waterways in the black spruce
muskeg of Alaska cause melting of the frozen ground with a conse-
quent destruction of the spruce vegetation through the caving in of
the surrounding terrain and a conversion of the area to treeless
bogs.
In time the spruce forest with frozen ground becomes reestablished
and the cycle appears to continue indefinitely (Drury, 1952). In
such situations the whole area might be considered as being in the
climax condition and the alternation of vegetation types merely fluc-
tuations within the climax. Possibly analogous fluctuations or cyclic
changes in the fauna and flora occur in the oceanic community. This
type of fluctuation within communities that are apparently in the

climax condition will be further discussed in the latter part of this
chapter.
When a climax community has become established, it tends to
remain in possession of the area because it does not change the en-
vironment so as to injure itself or to favor the growth of different
dominant species, and because its members can resist all competition
from the outside. The
succession of communities leading to a recog-
nized type of climax is termed a sere. Seres composed of different
sequences of communities typify different situations.

On land the type of climax community in which the sere culminates
is often determined primarily by the climate, and similar climax com-
munities dominated by plants often extend over large regions. In
local areas with special edaphic or physiographic conditions, or sub-
ject to recurring
fire or disease, a
type of community different from
the surrounding region may be displayed. Clements (1936) and his
followers originally believed that given sufficient time every local area
would eventually develop the same type of climax community the
as a whole under the prevailing
type characteristic of the region
climate, and that the continuing existence of a different type of com-
munity in was due to "arrested succession." Subse-

a local area
the strict application of the mono-
quently ecologists have abandoned
climax theory based on climate alone, and many have adopted a
polyclimax theory in which the type

of community that maintains
itself in each area

is
regarded as the climax for that area. Since
communities do not form a sharply delineated mosaic but grade into
one another with many variations of species composition according to
local conditions, Whittaker (1953) has proposed a climax pattern
hypothesis in which "vegetation is conceived as a

pattern of popula-
tions, variously related to one another, corresponding to the pattern
of environmental gradients."
The climax community remains in
possession of the area until some
unusual change causes its
displacement. The biotope and its biota
may be completely destroyed by a cataclysm such as a volcanic erup-

tion or extensive erosion; serious but incomplete destruction may
result from a forest fire, flood, or hurricane. On the other hand, the
climax community may be
displaced less violently by removal of the
dominant species through lumbering or the attacks of parasites.
Types of Succession
Primary Succession. Ecological succession that begins on a bare

area where no life has existed, or where the previous fauna and flora
have been completely destroyed, is known as primary succession.
Habitats that become available for initial colonization include: new
islands, sand bars, deltas, or glacial moraines; recently formed ponds;
fresh alluvial, shore, or volcanic deposits; and various types of sub-
strata exposed by erosion. These diverse areas may be classified as
xeric, mesic, or hydric according to whether the initial moisture con-
ditions are dry, intermediate, or wet. Seres starting from these types
of situations represent xerarch, mesarch, and hydrarch succession,
respectively.
A striking example of primary succession, and a classical one, is
the hydrarch succession in which a pond and its community is con-
verted to dry land with an entirely different community. The vege-
tation rooted along the margins of the pond is able to push out from
shallow water into deeper water in a variety of ways ( Fig. 12.2 ) As .
the vegetation invades the open water, the margin of the pond is
reduced. At the same time the growth of the plankton and of other
aquatic organisms adds organic matter, and much of this is deposited
on the bottom. Beavers, muskrats, and other animals carry material
into the pond, deciduous vegetation blows in from the shore, and silt
is carried in from surrounding land. Rafts of vegetation from the
pond margin drift offshore, strand, and take root, thus establishing
islets that grow in size until they meet and also join the shore. At
the same time the area available for completely aquatic plants, such
as the water lilies, becomes reduced. As the free water is changed
to swampy land, the water lilies and similar species give way to
sedges and rushes, and these are subsequently replaced by heaths and
shrubs. As succession continues, the soil is further built up, so that
it becomes drier and is also changed
chemically. In time certain smaller spe-
cies of trees invade the area, taking the
place of the shrubs, and eventually full-

sized forest trees will dominate the scene.
Various stages in such a succession are
shown in Fig. 12.3.
The existence of zonation in a com-
munity does not necessarily mean that
succession is going on since distinct hori-
zontal subdivisions may occur in a static
community, as described in the previous
chapter. Furthermore, the change in

the vegetation is not always self -induced
but may be caused by outside influences.
Sometimes the conversion of a swamp to
dry land is brought about primarily by a
lowering of the water table caused by a
physiographic change. But in other in-
stances, as in the situation described in
the preceding paragraph, the vegetation
responsible for building

itself is chiefly
up the land as ecological succession goes

forward.
At the same time that the vegetation
is
undergoing these profound changes in d
the hydrarch succession, the animal life
of the community is correspondingly
altered. Fish, beavers, and muskrats
will gradually be excluded and terrestrial
vertebrates will enter. Less conspicuous
but just as significant will be the mani-
festations of succession among the in-
vertebrates and the microorganisms.
These trends in the animal members of
the community are indicated schemati-
cally in Fig. 12.4 for a hydrosere in Illi-

nois. The changes in bird species asso-
ciated with a hydrarch succession are
shown in Table 19. The changes in the
FIG. Five methods by which vegetation invades deeper water from the
12.2.
pond margin: (a) spike rush, (b) tussock sedge, (c) loosestrife, (d) cat-tail flag,
and (e) sphagnum and heath. (Needham & Lloyd, 1937, Copyright, Cornell
Univ. Press.)
433
FIG. 12.3. Hydrarch succession in Gifford Bog, Falmouth, Massachusetts. Pond

lilies(Nymphaea)> bulrushes (Scirpus), and masses of filamentous algae fill in
the open water. Cassandra or leather leaf (Chamaedaphne) forms islets and also
invades the bog at its margin, where it is followed by sweet pepperbush (Clethra
alnifolia) and later by forest vegetation.
Primary Succession 435
Bare
bottom
(pioneer) stage
Temporary
and prairie
Beech and
maple forest
(climax) stage
POND SUCCESSION
FIG. 12.4. Schematic representation of some of the changes in animal life during
successional stages from pond to forest in Illinois. ( Buchsbaum, 1937. )
insect members of serai communities leading to a red oak-maple climax

are discussedby Smith (1928).
Secondary Succession. When a natural area is disturbed so as to
destroy the community inhabiting it and to set back the course of
succession, the new series of communities tending again toward the
climax constitutes secondary succession. This situation arises when
the principal species of the community have been destroyed by fire,
disease, tornado, flood, or by human activities such as farming or
a community
lumbering. In some instances of secondary succession
is established that is essentially the same as a stage in the previous
primary succession, but

in other instances a quite different
community
is
brought by the
into being special conditions resulting from the dis-
turbance. However, later stages tend toward the type of community
found in the primary succession, and ordinarily the same climax is
eventually reached.
An admirable illustration of secondary succession may be observed
today in old fields of central
New England, and the phenomenon oc-
curred on a large scale during the last century when wholesale farm
abandonment took place. The early settlers of this region cut down
the forests to build farms and homesteads. Where the land was in-
tensively cultivated,
the stumps and roots of the forest trees were
TABLE 19
REPRESENTATIVE SPECIES OF BIRDS PRESENT IN VARIOUS STAGES or A

TYPICAL HYDROSERE IN NORTHEASTERN OHIO TO SHOW THE CHANGE IN
BIRD LIFE THAT ACCOMPANIES THE CHANGE IN VEGETATION
(Aldrich, 1945)
X = present at times; P * permanent resident; 5 seasonal.

Secondary Succession 437
completely removed from the and the cleared areas were planted
soil,
to crops or used as pastures.

Beginning shortly after 1830 the open-
ing of rich farmlands in the west, the
building of railroads, and the
growth of industrial centers, brought about the exodus of the farmer
from the rocky hillsides of New England that were so difficult to work.
The abandoned fields were covered
by a thick sod of grass. In this
turf the seeds of white pine trees blown from neighboring forested
areas found a favorable environment for germination, but other forest
treeswere not able to establish themselves effectively in the aban-
doned fields. The result was that the white pines grew into a dense,
uniform stand, and eventually produced a forest in which all the
dominant trees were of this species. A few hardwood saplings later
became established as an understory throughout the pine forest, but
their further growth was suppressed by the pines.
In the course of fifty years
or so, the pines had grown to a size that
made a valuable timber harvest possible. Great areas were lumbered
off; in regions where lumbering was not carried out the even-aged
trees grew old and finally

fell. In both situations the trees which
then grew into dominant position were not white pines but were
hardwood trees that had existed in subordinate position almost un-
noticed in the pine forest. Pines will not sprout from stumps, and,
being an intolerant species, the pine seedlings are unable to develop

in the shade of the mature trees (Fig. 12.5). The root sprouts and
FIG. 12.5. View mature white pines in southern

of the forest floor in a stand of
New the absence of pine seedlings but the presence of beech
Hampshire, showing
and maple saplings.
seedlings of beeeh, maple, and other tolerant species that existed as

minor members of the community grew into mature trees when they
were "'released" by the removal of the pines. White pine stands
formed abandoned fields of New England were therefore not self-
in
perpetuating. Once the mature pines were gone, the species com-
position of the forests changed completely, and gave way to a com-
munity of mixed hardwoods. Thus, secondary succession originating
in old fields of this region produces a transient community of pines
but eventually results in the restoration of the typical climax forest.
Convergence. Since the bare areas in which primary succession
can be initiated are extremely diverse, it is not surprising to find a
correspondingly great variety in pioneer communities. As succession

proceeds, however, the different biotopes tend to be modified toward
a more nearly uniform condition that will support similar communi-
ties. This convergence is particularly clear in relation to the moisture
factor serai communities cause hydric habitats to become drier, and
xeric habitats to become moister, so that a trend toward a more nearly
mesic condition is followed from either extreme.
As a general prin-
cipal we
recognize a convergence in succession such that seres orig-
inating in diverse habitats within a region of similar climate tend to
progress toward similar climax communities.

An examination of succession in the deciduous forest region of
Indiana revealed a tendency to converge that could be traced in seres
originating from five kinds of biotopes as shown in the scheme on page
439 from Clements and Shelford (1939). Pioneer communities
established on a sand ridge, clay bank, flood plain, shallow pond, or
deep pond initiated the successions indicated by the serial numbers

with the dominant plants named. The exclusive presence, or relative
abundance, of a dominant plant served as a serai index to the stage
of succession in each sere. An animal species characteristic of each
stage could also be recognized, and the names of these animal serai
indices appear in the diagram. Succession in each of the widely
different original areas tended to bring about the eventual establish-
ment of the same type of climax community in which beech and sugar
maple trees and the salamander, Plethodon cinereus, are the indices.
Succession in Special Habitats. The instances of succession dis-
cussed above are typical of broad areas in temperate regions. Other
manifestations of succession occur in special habitatseither large or
small. A classical example of succession in a microhabitat is the
sequence of bacteria, protozoans, and other microorganisms that follow

one another in a hay infusion formed by allowing hay to rot in a
quantity of water.
The seasonal succession of different kinds of
M
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phytoplankton and of zooplankton in natural waters is in part caused

by temporary modification of the medium by the organisms them-
selves and in part by seasonal changes in the physical environment,
as discussed in Chapters 8 and 13. The special circumstances of suc-
cession in arctic areas in which the substratum is repeatedly dis-
turbed by frost action are described by Hopkins and Sigafoos (1951).
Succession on a small scale, but of a complex nature, takes place
on and in the trunks of fallen trees. The sound wood, cambium, and
bark are attacked by a group of boring insects, saprophytic fungi,
first
and various kinds of microorganisms. This pioneer assemblage is

followed by a series of more elaborate communities that cause and
accompany the further disintegration and decomposition of the tree

trunk. The presence of these organisms attracts predators and
scavengers until a very large number of species may be represented
in the dead-tree-trunk biotope. Mosses, lichens, ferns, and, later,
higher plants find the rotting log a favorable substratum for growth.
With the establishment of autotrophic plants a new cycle of con-
structive growth begins, in which the new pioneers take root literally
as well as figuratively from the remains of the previous cycle.
The building up of coast lines is another activity in which the course
of succession and the physiographic changes are mutually interde-
pendent. The
ecological steps involved in and following the forma-
tion of sand dunes along the southern shore of Lake Michigan as one
community succeeds another have been studied by a number of ecol-

ogists. In simplest terms the course of succession was found to in-
volve the capture of moving sand by grass, and the development of
communities dominated successively by cottonwoods, pines, and oaks,
leading to the beech-maple forest. Subsequent investigation has
shown that the succession is far more elaborate and
also more flexible
than was at first supposed and that several interlocking channels for
advancement or recession may be followed by the various series of
communities involved, as indicated in Fig. 12.6.
Along marine coast lines in tropical regions a well-known land-
building succession takes place (Fig. 12.7). Here red mangroves
( Rhizophora mangle )
work out from the shore by dropping viviparous
seedlings that will root only in water more than about 25 cm deep.
By means of spreading, stilt-like roots this species of mangrove can
maintain itself in and slightly below the tidal zone in spite of wave
action. Mudthat collects around the dense jumble of roots builds up
the bottom, causing the gradual elimination of the red mangrove, and
prepares the way for the establishment of the black mangrove

(Avicennia nitida), the seedlings of which will grow only in water
_2 G
rt .2
Q
S ~ >>
.2 M
w s
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O)
:
(^
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Q.
18
shallower than about 25 cm. As the bottom is raised and the soil
becomes drier, buttonwoods replace the mangroves, and these are
subsequently succeeded by the climax palm community.
In the situations described thus far the controlling organisms are
plants. On land, plants are usually the dominant members of the
community, and vegetation development is chiefly responsible for
causing succession, but in some communities animals are found to
exert control. For example, birds nesting in colonies are sometimes
so abundant that their activities and their droppings cause a change
in the vegetation. In this way a rookery of herring gulls on Kent
Island off the coast of New Brunswick caused the elimination of a
grove of spruce trees and its

replacement by grass.
MANGROVE SUCCESSION
..Climax
community Transition Salt-marsh Pioneer community
Red
mangrove
High tide
Seedling
j>
I*/ " '
Lowt.de
Crtll lino
Soil line
Underlying rock
FIG. 12.7. Diagram of succession along the margin of tropical shores as seen in
southern Florida. (Modified from Davis, 1940.)
Control of the environment by animals is more commonly found

in aquatic habitatswhere sessile forms are prominent, The photo-
graphs in Fig. 12.8show two stages in an all-animal warfare in the
tidal zone. Oysters became established on the sea bottom in this
biotope, but mussels soon began attaching to the oysters' shells.

Gradually the growth of the mussels smothered the oysters, and the
latter were replaced by an almost continuous carpet of mussels. Sub-
sequently, barnacles became attached to the shells of the mussels in
sufficient numbers to kill them. After the death of the mussels their
shellsbroke loose from the bottom and the barnacle population was
swept away by wave action so that no enduring change was brought
about in this instance.
Frequently both plants and animals are involved in well-defined
succession in the littoral zone as described by Dexter ( 1947 ) for mud-
bottom communities; similar
relationships occur on hard bottoms and
on submerged surfaces subject to attack
by fouling organisms. When
a jetty or pier is built, or a boat without
anti-fouling paint is left at
her moorings, various plants and animals attach in
recognizable se-
quences. Bacteria are the first to establish themselves, and they com-
monly form a film in which benthic diatoms and various filamentous
algae find a foothold. The subsequent course of succession varies
according to circumstances, but in some situations Bryozoa appear
From Miner, 1934

FIG. 12.8a, Mud flat community dominated by oysters. The shells of these ani-
mals provide a suitable substratum for the subsequent attachment of mussels.
From Miner, 1934

FIG. 12.8k. Mussels have overgrown the oysters and will in turn be smothered by
the barnacles that are seen beginning to attach in abundance to the mussel shells.
next, followed by mussels (Scheer, 1945), whereas in other situations

barnacles, tube worms, or tunicates attach at an early stage (Woods
Hole Oceanographic Institution, 1952). The varied surfaces of the
attached organisms, and particularly the crevices between them, pro-
vide abodes for a host of mobile forms that join the community in
these later stages and that could not inhabit the biotope if it were not
for the presence of the earlier arrivals. Thus, in these small habitats,
as well as in the larger areas considered, the early inhabitants
change
the conditions so as to cause their own displacement and the estab-
lishment of successive new communities.
Modification of Succession
The course of succession may be modified by unusual natural

circumstances, or frequently by the hand of man. Succession may
be changed, for example, by the browsing of an overly abundant
population of deer in a forest. Deer do not browse on all species of
the vegetation indiscriminately but have certain favorite items. If
the deer become numerous, the most appetizing plants will be grazed
down excessively and the unpalatable types will be given an ad-
vantage. In this way the species composition of the vegetation and
the course of succession considerably altered. If man permits
may be
overgrazing by cattle on the range, the usual sequence of succession
may be reversed so that the vegetation is pushed back from perennial
grasses to annual weeds (Graham, 1944,
Ch. 9).
Repeated forest fires sometimes bring succession to a halt at a com-
munity stage that is different from the climax in
neighboring areas.
In the sandy soils near Plymouth, Massachusetts, and on Cape Cod
the prevailing vegetation consists chiefly of pine and scrub oak, and
thisappears to be a climax condition
determined partly at least by
fire.On an undisturbed island in a large pond within this region a
mature stand of beech, maple, oak, hemlock, and yellow birch has
developed. This island has not been burned over during the memory
of present residents, and it may well be that because of the protection
of the water no fire has occurred on the island for generations. The
existence of this island community suggests that other neighboring
the pine and scrub oak stage if the
regions might progress beyond
interference of forest fires was prey0lited for a sufficient time. The
of southeastern United States may similarly
pinelands in some parts
controlled by rather than by climate
represent a climax community
fire
(Fig. 12.9).
Modification of succession by special factors may also be observed
Modification of Succession 445
on a small scale in
many local habitats. One example is furnished
by the activity of certain animals in maintaining open "glades" on the
surface of rocks in the tidal zone The browsing of
(Fig. 12.10).
limpets at the margins of their territories shears the
away enlarging
basal holdfasts of the algae that would normally grow over *he
reel
rock surface in the course of succession. Another animal, Idotea

viridis, bores holes in the fronds of the algae so that
they break away
easily and thus assists in affecting succession in \his microhabitat.
In contrast to the situations mentioned above the activities of man
'
may sometimes add a stage beyond the usual climax. In certain

of southern California the climax vegetation
parts formerly consisted
of grass or chaparral. When man introduced the from eucalyptus
Australia, this tree established extensive stands replacing the former
vegetation and adding a forest stage to the previously recognized limit

to succession in the region.
Ecologists have taken such an interest in pointing out the occur-

rence of succession and its
importance under various circumstances
that emphasis has failed to be laid upon the fact that in some regions
Photo by H. B. Moore
FIG. 12.9. Pineland community in southern Florida, consisting principally of
pines, palmettos, and sawgrass. After such an area is burned over, the same type
of community is reestablished directly.
no succession appears to take place. Where conditions are not

changed by the inhabitants of an area so as to favor other species,
succession does not occur. If the existing community is destroyed in
many regions of the tropics or subtropics, the area will be repopulated

directly by the original species. The communities of the open ocean
and of the deep sea may be regarded as being in a climax condition.
Since the physical conditions of the sea and also of large lakes are
essentially unmodifiable by the plant and animal inhabitants, no eco-

logical succession takes place that is comparable to that found on
land.
In the open ocean seasonal and short-term sequences in the communi-
ties of
temperate regions occur, as they do on land, but growth of
marine organisms in the plankton or on the floor of the deep sea does
not change the nature of the physical environment in such a way as
to cause a permanent or irreversible replacement of one community
by another that regularly extends beyond the seasonal cycle. For a

further discussion of ecological succession the reader is referred to
Costing (1948), Allee et al. (1949, Ch. 29), and for special situa-
tions to Hutchinson (1941), Dansereau (1951), and Niering (1953).
Photo by C. Davidson
FIG. 12.10. Open "glades" maintained by limpets (Patella) on rocks in the tidal
zone of the Isle of Cumbrae, Scotland. The browsing of these gastropods (seen
on the large rock in the foreground ) curtails the invasion of the red alga, Gigartina
stellata. Barnacles also inhabit the "glades."
Community Type 447
COMMUNITY CLASSIFICATION
Since the
community is essentially a functional unit, the size of the
area occupied by the
community, the number of living things included
in it, and its organization are variable As
according to circumstances
we have seen, the groups of plants and animals that constitute a com-
munity may be large or small, and frequently one functional group is
contained within another. Furthermore, one community reâces

another in the same area as
ecological succession proceeds. This
situation presents difficulties when we
attempt to arrange communi-
ties in some sort of order since
any practical classification is based
on descriptive considerations such as the life form of the dominant
species or their position in the terrain. The presence of some types
of communities is more
strongly influenced by climatic factors than
edaphic factors, but for other types the reverse is true.
Studies of the arrangement of communities in hierarchies have
been made chiefly in environments where plants hold the dominant
position in the biocenose. It is not surprising then that common sys-
tems of classifications are based primarily on the vegetation./ We
shall
describe the chief units into which communities are grouped, but
unfortunately complete agreement does not exist as to terminology.
Community Type
Communities of a certain dominant life form, such as deciduous
and the like, obviously compose recogniz-
forests, coniferous forests,
able categories. Some authorities, like Tansley (1939), regard any
mature community of distinctive life form as a "formation," but other
authorities, following Clements (1936), apply this term to climax
communities only. Extreme difficulty is often experienced in ascer-
in a given region has attained its
taining whether the vegetation
climax condition, and for many types of investigation it is not neces-
sary to do so. Also there
seems little logic in using a different term
for, say, grassland that climax and grassland that is on its way to
is
becoming forest. Whenthe state of ecological succession is known,

a modifier may be used for clarity by referring to the community as
a climax formation or a serai formation. Until general agreement on
to specify the particular sense
usage is reached, it will be necessary
in which the term formation is used on each occasion, or to avoid it
by referring to the life form of the dominant organisms or

to the
community type. Thus we may state

that the vegetation in a certain
area is a serai shrub formation, is of the shrub life form, or is of the

shrub community type.
The Biome
Certain communities whose dominant species have a distinctive life

form have become more or less permanently established in certain
climatic regions of the earth and are believed to be in the climax
condition. Associated with these climax communities are communi-
ties in earlier
stages of ecological succession and also communities of
a type controlled by special local conditions different from the general
nature of the region. Such a complex of communities, characterized
by a distinctive type of climax community and maintained under the

climatic conditions of the region is known as a biome. All animal
and plant components of each of the included communities are mem-
bers of the biome. The biomes constitute the great regions of the
world distinguished on an ecological basis, such as the tundra, the
desert, the grasslands, and the various forests, >
The biome consists of a special combination or complex of com-
munities. The essential matrix of the biome is composed of climax
communities with dominants of a certain life form that
give the biome
its
particular character. Communities of different life form are pres-
ent as minor constituents of the biome. The major climax communi-
ties in the biome are of the same type but differ in species composition
in different parts of the biom^ The deciduous forest biome that
extends across the eastern part of the United States, for example, is
characterized by a community type in which deciduous trees are
dominant.- Major communities of equal rank in this deciduous forest
biome are the oak-hickory forest of the central Atlantic states, the
beech-maple forest of the Middle West, the hardwood forest of
northern New England, and others of the same life form. These
major climax communities that form the essential matrix of the biome
have been termed associations by Clements and his followers. The
life form characteristic of the
major climax communities and hence
the general character of the biome is determined primarily by the
nature of the regional climate. Which of several major communities
of the same type will be present in each part of the biome is de-
termined by local variations in both climatic and edaphic conditions.
Within each community Clementsian ecologists recognize certain
subdivisions. Geographical differences in abundance or in relation-
ship of the dominants in the community are called faciations. Within
a region occupied by a grass community, for example, variations in
The Biome 449
climate with latitude and with altitude result in differences in the
selection of dominant species. Each of these various geographical
combinations of dominants constitutes a faciation of the community.
Variations of the dominant species on a more local scale when deline-
able are termed lociations.
The subdominant members of a community
may also form recog-
nizable groups. These subdivisions have been called "societies'" but
this term is unsatisfactory in this connection because of its
specific
use for highly integrated animal groups within the same species as
seen in insect colonies. If we again take a climax deciduous forest
as an and cryptogamic plants
illustration, the shrubs, seasonal herbs,
as well as the various categories of animal life
may be regarded as
constituting definite groups of subdominants. Many aquatic com-
munities including benthic plant and animal groups may be similarly
subdivided, but with the smaller life forms local subdivisions are not
as conspicuous communities dominated by the larger vegetation.
as in
Subdivision of communities involving plankton and nekton are even
less clearly defined because of the mobility of the medium and the
organisms.
Many ecologists feel that communities and their subdivisions cannot
be as clearly distinguished as suggested by the foregoing terms, and
that no system of community classification is really satisfactory.
Some investigators feel that each community is a law unto itself, as
argued (1926), since each is composed of whatever plants
by'Gleason
and animals have reached the area and have found conditions toler-
able. As the inhabitants in each situation have grown and multi-
;
plied, the community and its environment have changed until a stable
condition has been reached. If similar climax communities develop,
it is because similar conditions happened to exist, but an indefinite
amount of individual variation is
possible.
Regardless of the method of classifying communities, each biome
isfound to consist of several major communities (or associations) in
the climax condition and of many minor communities.. Between or
within the major climax communities the developmental stages of
these communities can be recognized and also other community types
produced by local variations in the environment. In the deciduous

forest biome, for example, grass and brush communities are found that
may eventually mature into forest communities. Also present are the
communities of ponds, marshes, rock ledges, and sand hills that will
not be converted into deciduous forest for a long time but are definite
members of the biome.j A certain amount of cohesion within the
biome is
provided by plant species common to two or more communi-
1H TUNDRA AND ICE
TAIGA
TEMPERATE DECIDUOUS FOREST
GRASSLAND
mm DESERT
TROPICAL FOREST
TEMPERATE RAINFOREST
FIG, 12.11, Generalized representation of the major biomes of the continents.

(Modified from Goode, 1943, Copyright, Rand McNally and Co.)
Fluctuation within the Community 451
tiesand by the mammals, birds, and other animals that move back
and forth between various communities of the biome. For example,
beavers are members of a pond
community within a forest biome and
ako are members of an aspen community that borders the pond.
LThe biomes are the result of the equilibria established by living
inhabitants with all
aspects of the climate of the region. Since the
biomes are characterized by major communities of distinctive life
form, they correspond to certain of the life zones that are based on
the same life forms. The chief biomes that occur in a series frp) the
equator toward the poles coincide with the principal continental life
zones described in Chapter 5. (jjjome biomes tend to be continuous in
extent and form a more or less definite unit, but others are discon-
tinuous and the parts may be widely separated geographically.
Communities of the same life form as a biome but too small to con-
stitute a separate biome are referred to as biome types.
The clearly defined major biomes of the land masses of the world
are shown in Fig. 12.11. The sea may be regarded as constituting
an additional biome, but the plants of the open ocean do not exert
the controlling influence on the biotope experienced on landi/ All
parts of the sea are interconnected, and many kinds of plankton and
nekton move readily from one region to another. The biogeographic
subdivisions of the sea are therefore based principally on physical
features of the environment, as has been described in previous chap-
ters and is further elaborated by Ekman ( 1953 ) .
The principal biomes on land are: tundra, taiga, temperate decidu-

ous forest, temperate rain forest, grassland, desert, and tropical forest.
In certain regions, as in central North America, the dividing line be-
tween biomes tends to run longitudinally, but for the most part the
chief biomes of the world are arranged in a general latitudinal se-
quence. For a fuller description of the biomes of the world the

reader is referred to the general account of Cain (1944) and Allee
et al. (1949, Ch. 30) and to suck special "treatments as Richards
(1952) and Beard^(1953)...
FLUCTUATION WITHIN THE COMMUNITY

When the climax community has been established, can we then
take a long breath and say that at last the ecosystem will be entirely
constant? The answer is, No. Even within the relatively permanent
and stable climax community fluctuations occur sometimes of con-
siderable magnitude. It is true that in some situations the community
remains relatively steady. Sometimes, the life cycles of the members
of the population are

staggered so that little
change is
apparent for
the community as a whole. For
example, in certain tropical regions
where individual trees shed their leaves at different times, the vegeta-
tion as a whole remains
green throughout the year. Similarly, in
tropical waters individual species of plankton in small numbers wax

and wane, but the complexion of the community may continue essen-
tiallythe same for long periods. At higher latitudes where climatic
year, the abundance of popu-

conditions change markedly during the
and the composition of the community vary widely with the
lations
season. Yet even here the community may in some instances come
around each season to essentially the same composition as in the
previous year, resulting in a rather faithful repetition of the seasonal
cycle.
On
the other hand, constancy within a community is the exception
rather than the rule, even within the same broad climax condition.
Every species in the community fluctuates to some extent in respect
to rate of vegetative growth or of reproduction, and sometimes the

variations are very large. In earlier chapters we have mentioned the
various types of fluctuation of populations of individual species and
also the reciprocal oscillations of certain populations of species ex-
ploiting one another. We

shall now consider fluctuations in com-
ponent species against the background of the community as a whole.

All but the most minor changes in the growth or reproduction of one
species of a community is bound to have repercussions among other

members of the biocenose. Sometimes the fluctuations are obvious
and spectacular, as in a plague of mice or of locusts; on other occa-
sions changes in the community originate in fluctuations among the
unseen microorganisms of the soil or of the water, or among patho-
genic bacteria or viruses. In either case the effects on the natural
community may be serious,and they often have a drastic direct or

indirect consequence for man.
It is difficult to ascertain the magnitude of fluctuations
exceedingly
in the community in any precise terms. Attempts to take a census of
wild populations have been made for a relatively small number of
biotopes. Even in those instances in which a systematic count has

been undertaken, the reliability of the census methods is hard to
evaluate, and rarely have population measurements been continued
sufficiently long to
reveal significant relationships. The best available
census data for extended are for species
periods that are of concern
to man either because they cause damage or because they provide a
useful product. Records of abundance in wild populations have
been kept for various disease organisms and insect pests, and for
Irruptive Fluctuation 453
certain plants and animals harvested
by man. Among the latter the
most extensive records are those of trading posts for fur-bearing ani-
mals and those of fishery agencies for the landings of commercial fish.
We shall discuss some ecological aspects first of irruptive fluctuations

inwhich a population undergoes wide, irregular swings in abundance,
and then of oscillations that appear to be cyclic in nature.
Irruptive Fluctuation
Data from the commercial marine fisheries will serve admirably to

illustrate the circumstances of irregular fluctuation in a community.
It will be agreed that the general nature of the ocean has not changed
drastically during the last several hundred years or so; yet during that
time certain fish populations are known to have fluctuated in an ex-
treme manner. In the cod fishery off northern Norway, for example,
periods of scarcity severe enough to be recorded have occurred as far
back as the time of Leif Ericson. In certain intervening years, as in
the winters of 1714 and 1715, whole villages along the Norwegian
coast are reported to have starved because of the failure of the cod
fishery, whereas in the years immediately before and afterward codfish

were plentiful. In the same region similar fluctuations have con-
tinued to the present day, and equally extreme variations in the fish-
eries occurred on the American side of the Atlantic. The fluctuation
in the catch of mackeral by the American fishing fleet for a period of
150 years shown in Fig. 12.12 may be taken as representative of the
change in abundance of the mackerel population in the waters fished.
The most spectacular change during this period occurred between
FIG. 12.12. Fluctuations in the landings of mackerel (Scomber scombrus) by the
fishing fleet from the east coast of North America. (From data of the U. S. Fish
& Wildlife Service. )
1884 and 1886 when the catch of mackerel dropped from more than 60
million kg to about 10 million kg.
Subsequently the catch fell to a
still lower
figure but eventually rose again to a value of more than 30
million kg. The
seriousness of the ecological, economic, and socio-
logical repercussions of these tremendous oscillations can easily be

imagined.
Year-Class Analysis. The economic importance of the fluctuations
in the
Norwegian herring fishery led the great marine biologist Johan
Hjort to examine the annual differences in the population of the
herring. By adopting an analytical approach to the problem he was
able to trace the source of the oscillations of the entire catch to the
special success of fish hatched in certain years. Hjort began by con-

sidering a hypothetical, ideal situation in which the same number of
young herring were spawned each year and in which mortality

caused a uniform reduction in the abundance of each "year-class."
Under these circumstances 1-year-old fish would always be the most
abundant, 2-year-old fish the next most abundant, and so on. Since
herring of 4 years of age and older are taken in the commercial catch,
an analysis of the catch under these ideal conditions would show that
the 4-year-old fish were always the most numerous, the 5-year-old
fish were the next most abundant, and so on.
Applying this reasoning to the catch in 1910, for example, the fish
hatched in 1906 should be the most numerous, followed by the 1905
year-class,and then by earlier year-classes, provided always that the
same number were spawned each year and underwent the same mor-
tality. When the age composition of the actual catch in 1910 was
examined, however, a very different picture was presented, as shown
in Fig. 12.13. The 1906 year-class, which was 4 years old in 1910,
was by no means the most abundant, but occurred in extremely small
numbers, and the 1904 year-class was found to represent more than
80 per cent of the catch. Looking back in the diagram we see that the
1904 year-class dominated the catch in 1909 and in 1908; this same
year-class continued to dominate the catch for a good many years after
1910 and remained recognizable until 1921. No other dominating
year-class appeared until 1917 when the 1913 year-class entered the
scene, but in 1922 the 1918 year-class became prominent. are We

confronted then with the surprising discovery that during 20 years
only three year-classes were successful.

A good year-class is obviously
the exception; in spite of the tremendous fecundity of the fish en-
vironmental resistance is overpowering in most years. The fluctua-
tion in the herring fishery, and in many others, is now known to be
Year-Class Analysis 455
Age in years
4 5 6 7 8 9 10111213 14 15 16 17 18 19 20 21
FIG. 12,13. Age composition of catch of Norwegian herring (Clupea harengus) in

each of the indicated years, showing the predominance of a few good year-classes.
( Russell and Yonge, 1928, Copyright, Frederick Warne after Lea. )
due, in part at least, to the occasional production of a year-class in

which many more fish reach commercial size than in other years.
The cause of the varying success of different year-classes is difficult
to ascertain. Usually plenty of eggs are spawned every year; rarely
does unusual mortality occur after commercial size has been reached.
The critical point comes somewhere Since the gen-
in very early life.
eral nature of the ocean does not change, some subtle variation in the
environment must arise during a sensitive stage of development. Per-
haps an abnormal temperature in the spawning area is to be blamed;
possibly a serious change in the abundance of planktonic members of
the community-cither enemies or food organisms for the young fish-
is
responsible. Quantitative measurements of plankton in various
Feb. Mar. Apr. May June July

FIG. 12.14. Annual variation in average volume per standard net haul of a com-
mon species of planktonic copepod in the
waters between Cape Cod and Cape
Hatteras. (Sears and Clarke, 1940, Biological Bull.)
Cyclic Fluctuation 457
parts of the ocean have revealed great variations in abundance from

year to year. As an example, the drastic differences in the seasonal
growth of a population of a planktonic copepod during 1929 to 1932
are shown in Fig. 12.14. Presumably these great annual changes in
the copepods influenced other members of the oceanic community,
but no adequate explanation for these fluctuations in the plankton is
known. Possibly the survival of young fish is dependent upon a
simultaneous reduction in number of predators and an increase in
availability of food.The relation between conditions favoring plank-
ton growth and the abundance of young fish in the English Channel
is discussed
by Harvey (1950). Search for the factor or factors that
control year-class success in specific instances is one of the many fasci-
nating problems in ecology awaiting solution and obviously one of

great practical significance.
Cyclic Fluctuation
The fluctuations of the population in some communities appear to

be regular or cyclic. The phenomenon particularly noticeable in
is
high latitudes of the northern hemisphere among small mammals, cer-

tain predators, gallinaceous birds, and some fish such as the salmon.
The abundance of certain insect species and the rate of tree growth
also exhibit cycles in various parts of the world. In addition, the
occurrence of disease epidemics and other biological events are be-
lieved by some to be definitely cyclic.
Periodicity among northern mammals may
be studied from the
records of the Hudson Bay Company and other fur-trading agencies.
As with commercial fisheries we must realize that the variations in the
take by man may not represent the actual fluctuations of the entire
population. Nevertheless the mass of evidence from fur traders is

corroborated by other observations and leaves no doubt that oscil-
lations of approximately uniform periods take place over long periods
of time (Elton, 1942). Cycles of abundance with 3 to 4 years be-
tween peaks are exhibited by mice, voles, and lemmings, and also by
arctic foxes, martens, and snowy owls. Many of these cycles can be
traced back in the records for a hundred years or more. The snow-
shoe rabbit, the grouse, and certain insects are commonly reported to
fluctuate with a 9- to 10-year cycle. Data for the Canadian lynx are
available from 1735, but difference of opinion exists as to the size of
the that constitute maxima.
peaks
If only the "big" peaks are counted,
the average period of fluctuation is 9.6 years. If every year in which

the population is greater than the preceding and the following year
is considered to be a maximum, then analysis of the same records for

the Canadian lynx yield an average of 5.8 years for the period of the
cycle (Cole, 1951). Lynx populations in Finland oscillate with an

average of 3.5 years for the cycle. The 10-year cycle may be inter-
preted as the average distance between "high peaks" of a basic cycle
with maxima 3, 3, and 4 years apart, of which the maximum after the
4-year interval is larger than the others (Siivanen, 1948).
Possible cycles of other periods have been reported and are sum-
marized by Hutchinson and Deevey ( 1949 ) The long-haired rat in
.
western Queensland, Australia, appears to display an 11-year cycle.

Irruptions of the chinch bug (Blissus leucopterus) between 1823 and
1940 occurred with an average period of 9.6 years for the cycle, but
individual periods varied from 7 to 13 years and the outbreaks are not
considered to be truly cyclic. The population of guano-producing sea
birds off Peru undergo catastrophic reductions about every 7 years
when the warm current "El Nino" moves farther south than usual, as
described in Chapter 5.
Of a different nature is the periodicity of certain other insects in
which the cycles are controlled by the simultaneous emergence of
the adults after long, but often regular, periods of dormancy. Per-
haps the most spectacular is the 17-year cicada (or "locust") Magi-
cicada septendecim which in "locust years" swarms over the country-
side but during the intervening years remains unseen as a nymphal
stage in the ground. During one irruption in an oak-maple forest near

Chicago these cicadas occurred at a density of 500,000 per hectare (50
per sq m), equivalent to 77 kg of dry tissue per hectare (Strandine,
1940).
Fluctuation in growth of certain trees as revealed by the width of
their growth rings often appears to be definitely periodic. The records
studied by Douglas (1936) revealed a recognizable cycle of 9% to 10
years. The growth of hemlock trees in Pennsylvania, traced for over

230 years and of Douglas firs in Utah traced for about 1500 years
oscillated with average cycles of almost exactly 3 years (Schulman,
1948). Nevertheless, during the period 1703 to 1939 when the data
for both tree species can be compared, the fluctuations in the two
series appear to be completely independent.
Causes of Fluctuation
Discovery of the causes of fluctuations in communities is one of the

most desirable objectives of ecological research but also one of the
most difficult. We have seen that changes in the size of population
Causes of Fluctuation 459
are dependent upon the interplay of biotic and environmental
potential
resistance. Ordinarily the full biotic potential of animals and plants
in nature is not reached environmental factors
usually keep a strong
check on the population. In some instances, however, even a
slight
variation in environmental resistance a marked effect on
may produce
abundance. Consider
the accompanying hypothetical
example for
the annual recruitment of a fish population. From the
point of view
Number spawned each year: 1,000,000

of fish
1st year: 999,000 die; mortality 99.9%; 1000 left to catch
2nd year: 998,000 die; mortality 99.8%; 2000 left to catch
= 100% increase in number surviving
of the original number of fish spawned the variation in

mortality ap-
pears to be trivial, but from the point of view of the survival of fish
to form the adult population a very great difference is
produced in
the two sample years.
In addition to variations in physical features of the environment,
changes in the living elements, including both food supply and ene-
mies, will produce modifications of the environmental resistance. The
biotic potential of the prey is usually greater than that of the predator,
but the predator can destroy a much greater supply of its
prey than it
actually assimilates. As a
result predatory animals may sometimes
be the chief factor holding prey species in check. When an unusually
favorable combination of both physical and biological features of the
environment occurs, the species "breaks out" from its controls. If the
species possesses a high biotic potential, only a short period of un-

checked reproduction will cause the population to irrupt. One North
American species of field mouse can breed when it is 3 weeks old and
can produce 13 litters per year. A little arithmetic will indicate the
size to which the population could expand if most of the young sur-
vived for a year or two. Many invertebrates, including notably the
insects, can multiply even more explosively. After a population that
has escaped from usual control has increased for a period, its ex-
its
pansion may eventually be stopped by another set of predators, by

disease, or finally by a lack of food. The maxima of snowshoe rabbits
are brought to an end reputedly by a disease that causes death from
shock, possibly induced by shortage of some mineral element in the
diet.
which influence or combination of influences in the en-

Precisely
of a population and
vironment is
responsible for the augmentation
which brings the increase of a population to a halt or causes it to re-
cede is difficult to determine. In attempting to solve the
extremely
problem as complete a knowledge as possible must be obtained of

the interrelations of the species in question with all aspects of its en-
vironment at every stage in its life cycle. Apparently correlated con-
ditions must be examined over a period of time and in different parts
range. When it is feasible, the factor suspected of causing the
of the
fluctuation should be tested by laboratory or field experiments.
Many growth and reproduction by

illustrations of the control of
individual environmental factors, or combination of factors, have been
given in previous chapters, and others may be found in reports dealing

primarily with this topic, such as those of Shelford (1951, 1951a).
Following extensive investigations of the fluctuations in certain forest
and non-forest animal populations in the Middle West, Shelford
emphasized the fact that environmental influences exert their con-
trol
during sensitive periods in the life histories of the
chiefly
species concerned. Forest invertebrates, for example, were found to
develop larger populations in years when rainfall is great during late

March, April, and May with short dry periods, whereas for certain
birds and mammals the conditions in
February and March are more
significant.
Origin of Cycles. Because of the lack of sufficiently precise data
we cannot at present state that the fluctuation of any population is
strictly cyclic in the sense that accurate prediction of the times of
maxima and minima can be made well into the future. Further study
of the possible existence of truly cyclic behavior is very much desired
since, in addition to the theoretical interest, prediction would permit
precautions to be taken against outbreaks of pests and preparations to
be made for glut or scarcity of fish, game, or furs. Since fluctuation
may be only approximately cyclic, we must include random variations
as a
possible explanation. As possible causes of these biological
cycleswe shall examine self-induced effects within the population,
random effects of many external factors, and single controlling in-
fluences that themselves fluctuate in a random fashion or are truly
cyclic.
Regular oscillation in abundance of a single species might be pro-
duced in a constant environment as the result of delays in the effects
of excessive or favorable numbers. Such a population would alter-

nately overshoot and undershoot a possible equilibrium level, as de-
scribed in Chapter 9. When two or more species affect each other's
abundance reciprocally, as may occur in prey-predator or host-parasite
combinations, oscillations may be similarly set up, as pointed out in
Chapter 10. Such oscillations caused by reactions within populations
of one or more species are termed
intrinsic
cycles.
Origin of Cycles 461
As a species becomes more abundant, its metabolites tend to accu-
mulate and other physical and chemical
changes are brought about
that may curtail the reproductive rate of the
population. With in-
creased density of the population are attracted and disease
predators
parasites can spread more easily. As crowding continues, food and
other necessities become scarcer, and the animals tend to wander
more extensively. This latter fact causes the further distribution of
infectious disease and exposes the animals more frequently to detec-
tion their predatory enemies.
by Furthermore, with progressive re-
duction of the vegetation by herbivores less cover for protection
against their predators is available. Thus increase in abundance may
automatically cause a curtailment of the population in ways that might
tend to bring about regular self-induced oscillations. However, the
inevitable occurrence of other changes in the environment would be
expected to impose irregularities upon any intrinsic cycle.
Approximately regular fluctuations in populations caused by in-
dependent changes in the environment either animate or inanimate-
are known as extrinsic cycles. Obviously intrinsic and extrinsic
types of cycles are not sharply separated since many populations

respond both to self-induced and independent variations in their en-
vironment. Irregularities in the environment might set in motion
regular internal oscillations by a sort of resonance, just as irregular

waves can cause a boat to roll in its natural period.
Scrutiny of the data on populations with apparently cyclic fluctua-
tions has led Cole (1951) to conclude that "the oscillations of any
hypothetical factor determining population size need only be about as
regular as would be expected of a random variable." Since we know
that the abundance of every species is affected by many environmental
factors, it
may be that chance combination of favorable and unfavor-
able influences in any year will produce apparently cyclic fluctuations.
Support for this simple explanation is found in a comparison of certain

fluctuations in nature with those of a series of random sampling num-
bers in which the former are no more convincingly "cyclic" than the
latter (Fig. 12.15). Cole also urges that here, as elsewhere, the so-
called "law of scientific parsimony" be applied, that is, the avoidance
of a more complicated hypothesis than is actually required to explain
the observed facts. However, it is
unlikely that random fluctuation
alone would produce cycles of the same period in widely different
species.
In some instances variation in abundance or growth has been defi-
nitely related to the

variation of some extrinsic factor in the environ-
ment, and no doubt many more such relationships will be found in the
Tippett's numbers, 2-pomt moving sum
1850 1860 1870 1880 1890 1900 1910

FIG. 12.15. Fluctuating series of animal populations and tree growth compared
with series of random numbers. In the 2-point moving sum curve the height of
each point depends upon the values of two successive random numbers, much as
the
population size might be affected not only by conditions this year but also by
size of last year's population. (Cole, 1951.)
Origin of Cycles 463
future. Fluctuation in populations of
guano-producing birds is defi-
nitely determined by periodic influx of warm currents. The growth

of Douglas fir in Utah was found to be correlated with
closely precipi-
tation, but the growth of hemlock trees in Pennsylvania the
during
same years with the same average cycle period exhibited no significant
relation to rainfall. The oscillations of the arctic fox ( Alopex lagopus)
and of the snowy owl (Nyctea nyctea) are evidently timed by those
of the lemmings and mouse-like rodents on which
they feed. As the
rodent population diminishes, the snowy owls disperse in search of
food, and their periodic appearance as far south as New England
attracts much interest among ornithologists (Gross, 1947). Maxima
of the Canadian lynx follow those of the snowshoe rabbit by a year or
so. The fact that the colored fox (Vulpes fulva) and the marten
(Maries americana) exhibit a 4-year cycle in the arctic and a 9- or
10-year cycle farther south is perhaps due to a corresponding differ-
ence in the periodicity of its prey in the two regions. In at least some
of these instances the fluctuations of the prey are not controlled by
the predators as in reciprocating intrinsic cycles. For example, the
snowshoe rabbits on Anticosti Island, where there are no lynx, fluctuate
with the rabbit population on the mainland where lynx abound.
In the foregoing type of situation the apparently cyclic fluctuation
of the species is not due to a random effect of many environmental
influences but is
by the changes in one par-
controlled principally
ticular factor. The biological cycle might then be caused either by
apparently cyclic random fluctuations of this controlling factor, or by
a truly cyclic change in this factor. A long search has been made for
a predictably regular cycle in the climate or in some other feature of
the environment that might serve as a master timer for biological
cycles, but no convincing and generally accepted factor has been

found. Sunspots were at one time thought to be correlated with cer-
tain population cycles since their average period is about 11 years and
is thus closely similar to certain cycles of growth and abundance.
However, the sunspot cycle is more variable than the Canadian
lynx
cycle, for example,
since the period of the former has varied from 7 to
17 years as against 8 to 12 years for the latter. In addition, a com-
parison of the two cycles

from 1750 to 1935 shows that, although they
for a number of years, they subsequently become com-
correspond
pletely out of phase (Fig. 12.16).
A
satisfactory explanation
of these intriguing cycles of growth and
abundance accordingly continues to elude us. These fluctuations are

another manifestation of the many changes going on within the com-
munity that have been reviewed in this chapter. We have seen that
1750 1760 1770 1780 1790 1800 1810 1820 1830 1840
Time in years
1850 1860 1870 1880 1890 1900 1910 1920 1930

Time in years
FIG. 12.16. Population trends of the Canadian lynx (solid line) graphed against
sunspot numbers (dotted line). (Alee et al., 1949 after MacLulich, 1937.)
modifications brought about by the life

processes of the inhabitants in
their reciprocal relations with the environment cause ecological suc-
cession to go forward from the initial colonization of a new biotope
to the climax condition. The biomes of the world represent the per-
manent establishment of the principal climax communities in general
equilibrium with their environment. Within these generally main-

tained conditions, however, many seemingly regular and many ir-
regular fluctuations continually take place.

13
Dynamics of
the Ecosystem
In the two previous chapters we have discussed the composition of

communities, their progressive changes, and the fluctuations that occur
among their members. We shall now consider the dynamics of the
ecosystem as a whole, that is, the operation of the community and its
environment as a functional unit. We
shall also discuss certain quan-
titative aspects of the interdependencies in the ecosystem, including
the concepts of productivity.
FUNDAMENTAL OPERATION
natural situation shall we select for the study and illustration
What
of the fundamental operation of the ecosystem? We might take the
whole world as a unit. We could measure the energy received by
the earth and follow its transformations; we could measure the
amounts and kinds of organic matter that are elaborated. In other
words, we
could evaluate the earth as a mechanism for biological
transformations. A study of this sort is of interest and involves sub-
ject matter treated in the field of biogeochemistry in which the distri-
bution and transformation of biologically important materials are
traced (Hutchinson, 1948). However, we are more often concerned
with the operation of particular types of ecosystems, and, in any event,
an investigation of the variations from place to place and from time to
time would be required before an adequate summation could be made
for the entire world.
In selecting an area for study we might go to the opposite extreme
and investigate a single plant together with the space immediately
surrounding it. Although certain relations of the exchange between
the plant and its environment would be revealed, the unit studied
would probably not be large enough to include all related influences.
Ideally, it would be desirable
to study an area that is large enough
to be representative and to contain all the fundamental factors, and at
the same time that is cut off from outside influences. A completely
465
466 Dynamics of the Ecosystem
self-sufficient ecosystem rarely occurs in nature, but situations ap-
proaching this condition may be found. A balanced aquarium
represents an artificially established, self-contained system in which

the number and type of plants and animals are adjusted so that a glass
cover can be placed over the top and the organisms within will main-
tain themselves
indefinitely. A pond with no inlet or outlet, or with
very water
little
exchange, may approach self-sufficiency. This idea
was first set forth by Forbes ( 1887 ) in his classical essay, "The Lake
as a Microcosm." In the following quotation it is clear that Forbes
was thinking primarily and was not taking
of the animals in the lake
the complete ecosystem into consideration, but he states the essence
of the concept of self-sufficiency, and his essay has become a landmark
in the development of ecological thought:
The animals of such a body of water are, as a whole, remarkably isolated
closely related among themselves in all their interests, but so far independ-
ent of the land about them that if every terrestrial animal were suddenly
annihilated, it would doubtless be long before the general multitude of the
inhabitants of the lake would feel the effects of this event in any important
way. It is an islet of older, lower life in the midst of the higher, more re-
cent life of the surrounding region. It forms a little world within itself
a microcosm within which all the elemental forces are at work and the play of
life
goes on in full on so small a scale as to bring it easily within the
mental grasp.
If an isolated and self-sufficient microcosm cannot be found, a

habitat may be used for study in which exchanges with other areas
are slight and regular, or can be measured. Thus, the living and non-
living materials entering a pond by way of the inlet and the substances
being carried away by the outlet can sometimes be measured, and
suitable allowance can be made. Another possibility is the study of
a unit area, or unit volume, within a uniform region. The operation
of ecological forces may be investigated in the ecosystem represented
by 1 hectare in the middle of a large tract of forest, or in 1 sq km of
grassland inrange country, or in a certain area of the ocean. In these

instances we know very well that the unit we select is not isolated from
neighboring units, but we may assume that transfer out of our unit is
compensated for
by from surrounding areas so that
transfer into it
ecological activity within our area may be treated as if the study

unit
were self-contained.
Principal Steps and Components

The fundamental steps in the operation of the ecosystem
are: (1)
reception of energy; (2) production of organic material by producers;

Principal Steps and Components 467
(3) consumption of this material by consumers and its further elabora-

tion; (4) decomposition to
inorganic compounds; and (5) transforma-
tion to forms suitable for the nutrition of the area
producers. If the
is
occupied by a self-sufficient community, all these steps will go on
within it; otherwise materials must enter from surrounding areas.
These steps in the operation of the
ecosystem not only involve the
production, growth, and death of the living components but also may
influence the non-living aspects of the habitat. For example, con-
siderable energy is represented in the organization of the soil that is
brought about by the growth of the vegetation.

The diagram in Fig. 13.1 illustrates the fundamental steps in a self-
Essential components Non-essential components
Producers
Herbivores Carnivores
[ [ |
Consumers
Transformers
[Nutrients |< |
Non-living Living
components components
FIG. 13.1. Principal steps
and components in a self-sufficient ecosystem.
the nature of the components taking

sufficientecosystem and shows
and material. The non-living components
part in the cycle of energy
and nutrients for the
that must be present are light energy inorganic
growth of photosynthetic plants.

The living components of the eco-
and consumers. The producers are repre-
system consist of producers
bacteria.
sented primarily by green plants but also include synthetic
The activity of the synthetic bacteria is of minor consequence in most
instances, but in specialized situations

it
may be of quantitative im-
portance. The consumers include the
allother types of organisms in
the community. Free-living herbivores feed
directly upon the green
plants. The principal herbivores in terrestrial habitats are insects,
rodents,and ruminants, and in aquatic habitats they are small Crus-
taceans and mollusks. Since most other prominent
types of animal
life in the
community depend upon these herbivores, the latter have
been referred to by Elton (1939) as "key industry animals." The
herbivores serve as food for primary carnivores; these in turn may fall
prey to secondary carnivores, and several more stages of nutritional

dependency may exist. The tissues of the various plants and animals
in the community may also be eaten
by parasites, and after the organ-
isms are dead, by scavengers and saprophytes of many sorts. The
plants and animals that depend successively one on another form the
links of a food chain.
The dead bodies of the producers and of the consumers mentioned
above are attacked by decomposers, consisting of bacteria and other
types of fungi. The decomposers render the organic matter soluble
and break it down chemically. The material is then attacked by
transformers other types of bacteria that change the inorganic com-
pounds into forms suitable to serve as nutrients for photosynthetic

plants once again. Among the living components only the photosyn-
thetic plant producers, the decomposers, and the transformers are
essential. Some communities could theoretically maintain themselves
indefinitely without the presence of any animals whatsoever, and,
although it may injure our ego to state it, we realize that man is not a
necessary part of the ecosystem. However, animal life is ordinarily
present in most natural communities, and sometimes animals hold a
controlling position or take a prominent part in the operation of the
ecosystem.
Niches. Different species of animals and plants fulfill different
functions in the ecological complex. The role of each is
spoken of as
its niche. The term so used stresses the function of each organism
in the community rather than its physical place in the habitat. The
term originated, however, from the characteristic location of different
types of organisms in the area under consideration, and niche is still
used by some ecologists in this sense. The "functional niche" is more
fundamental than the "place niche," but both concepts exist and
should eventually be given different names*.
Niche may be used in a broad sense to refer to the principal func-
tions involved in the operation of the ecosystem, or it
may be employed
to describe the subdivisions of these and the various methods of
"making a living" within each functional category. Among herbivores

that feed on trees, for
example, some fill the niche of eating the leaves
whereas others use the twigs, sap, bark, or roots as a source of food.
In different geographical
regions each type of niche is often filled by
Activity in some niches requires extreme and
quite different species.
often bizarre of or behavior.
specialization anatomy, physiology,
Minute differences in function are exhibited
by the niches of birds that
obtain food organisms from crevices in bark. Small
woodpeckers
characteristically fly to the base of a tree and climb up the trunk
looking for grubs. When they reach the top, they fly diagonally down-
ward to the base of the next tree and again work their way upward.
In contrast, nuthatches common in eastern United States ordinarily
fly to
the top of a tree and work toward the base, clinging upside
down to the trunk. In this way woodpeckers readily remove grubs
and other food from crevices easily reached from below, whereas the
nuthatches obtain their prey from cracks in the bark more accessible
from above. In the Galapagos Islands a finch that has evolved toward
the woodpecker type fills this niche, but having no barbs on its tongue
it
wedges grubs out of crevices by means of a thin twig held in its
beak.
Trophic Levels and Relations. Each successive level of nourish-

ment as represented by the links of the food chain is known as a
trophic level. The plant producers within an ecosystem constitute

the first
trophic level, the herbivores form the second trophic level,
and the primary carnivores represent the third level. Additional links
in the main food chain, and in side chains such as those formed by
parasites, constitute further trophic levels.

Three types of pyramidal
relations may be found among the organisms at different trophic levels
in the ecosystem, resulting in the production pyramid, the pyramid of
biomasses, and the pyramid of numbers.
At each step in the food chain a loss of energy and of material from
the system takes place because the processes of assimilation and
growth are not 100 per cent efficient. This means that the organic
matter produced per average unit of time, and the energy represented
by it, become less at each successive trophic level. The production
rates of the components of a self-sufficient ecosystem may thus be
thought of as forming a pyramid. The base of this production

pyramid is represented by the organic synthesis of the green plant
component, and higher levels are represented by the growth rates of
the herbivore and carnivore components.
Frequently, the size, growth rate, and longevity of

the species
making up a particular ecosystem are such that the living weight, or

biomass, of the members of the food chain present at any one time
also form a second type of pyramid. In a land biotope the biomass of
the vegetation existing at the moment of observation is
commonly the
greatest, and the biomasses of herbivores, carnivores, and further
Jinks in the food chain are progressively smaller. Thus a pyramid of
biomasses results, but this pyramid is an indirect consequence of the
particular kinds of organisms present, in contrast to the production

pyramid which is a dynamic necessity in the operation of the eco-
system. In some aquatic biotopes the biomasses of some of the in-
habitants form a pyramid similar to the production pyramid, as in
Weber Lake, Wisconsin (Fig. 13.2), and on Georges Bank (Fig.
13.12), but in others the biomasses of fish and particularly of the
Bottom flora
Phytoplankton
FIG. 13.2. Pyramid of biomasses as illustrated by the weight relationships of the

various constituents of Weber Lake, Wisconsin. (Juday, 1943.)
bottom fauna are considerably greater than those of shorter-lived

components at lower trophic levels (Harvey, 1950, Table IV). The
operation of the ecosystem thus limits the relative
amounts of the
components that may be produced, but departures from a regular
reduction in the biomasses of components present at any one time
occur because of special circumstances.
Interdependencies within the ecosystem also exert a certain
influ-
ence on the size of the individual organisms involved. The essential
plant producers vary in size from the very largest organisms in some
types of community to the smallest in others. The primary producers
in different biocenoses range from giant trees, such as the redwoods,
to diatoms, often less than 30 in diameter, and
//.
green flagellates or
photosynthetic bacteria of still smaller dimensions.
The possible size of consumers is generally influenced by the size of
their food, but the
possession of specialized food-getting equipment
enables some consumers to be very different in size from the
organism
that provides their nutriment. Herbivores may be either larger or
smaller than the vegetation upon which
they feed. Insects inhabiting
forests are characteristically minute fraction
only a of the size of their
food plants, Some
rodents are larger and some smaller than their
food; crustaceans and mollusks that filter phytoplankton from natural
waters are generally very much larger than the plants upon which
they feed.
In the next step in the food chain, namely the subsistence of the
carnivores on the herbivores, the predatory species must be stronger
in some respect than its prey and is usually larger. Among animals,
therefore, a tendency exists for smaller species to form the early links
in the food chain and for larger species to form the later links. The
maximum size of a prey species that can be attacked successfully de-
pends upon the feeding apparatus of the predator. Thus some or-
ganisms in the community rnay be too large and some may be too
small for the consumer to catch or to obtain in sufficient quantity.
Since total biomass tends to become smaller at successive levels in
the food chain, and since size of individual generally becomes larger,
at leastamong the animals, it follows that in the typical situation a
reduction in the number of individuals comprising successive links
in the food chain takes place. A relation of numbers is another con-
the ani-
sequence of the operation of the, ecological complex. Among
mals of a community the herbivores are typically the most numerous;
the plant producers and
they take in food material synthesized by
consumers. Primary carnivores that prey
pass it on to the subsequent
upon these "key industry animals" are less abundant, and secondary
and tertiary carnivores generally exist in still fewer numbers. This
numerical relationship with the more abundant species near the base
of the food chain and the less abundant species near the top is known
In food chains involving parasites the
as the pyramid of numbers.
size relationships are reversed because the parasite is smaller than its
host, and hyperparasites must be still smaller. For this reason the
for the successive steps of parasite
pyramid of numbers is reversed
of each link are generally more numer-
dependency, and the parasites
ous than their hosts. *^
An illustration of the pyramid of numbers among mammals and
birds inhabiting range land is presented in Table 20. The differences
TABLE 20
ABUNDANCE OF BREEDING ANIMALS ON 1 SQUARE MILE (3 sq km), SANTA RITA

RANGE RESERVE, ARIZONA (After Leopold, 1933)
Species Number
Coyote 1
Horned owl 2
Redtail hawk 2
Blacktail jackrabbit 10
Hognosed and spotted skunk 15
Roadrunner 20
Cattle (over 1 year old) 25
Scaled quail 25
Cottontail 25
Allen's jackrabbit 45
Gambel quail 75
Kangaroo rat (Dipodomys) 1,300
Wood rat (Neotoma) 6,400
Mice and other rodents 18,000
in abundance of the carnivores, such as the coyotes, hawks, or owls,

and of the herbivores, such as the quail or the mice, are striking; an
even greater contrast in numbers would have appeared if the census
had included the insects and other invertebrates. An example of the
pyramid of numbers on a small scale is furnished by a census of the
metazoans inhabiting the floor of a deciduous forest (Fig. 13.3).
In addition to the foregoing relations of biomass, size, and num-
16-20
E !
lM5
6-10
1-5
<1
Number per kilogram

FIG. 13.3. Pyramid of numbers among the metazoans of the forest floor in a
deciduous forest near Evanston, Illinois. (Park, Alice, & Shelford, 1939, Copy-
right, Univ. of Chicago Press.)
Ecological Cycle in the Ocean 473
bers, certain aspects of availability are of crucial concern in the eco-
system. Food organisms must be sufficiently

concentrated in relation
to their size so that the can meet its nutritive requirement
predator
within the time available. In other words, the abundance of the
food per unit area must be adequate in relation to the
possible forag-
ing range of the animal concerned.] If there were only one lion in
all of Africa and zebras lived 100 miles
apart, plenty of food would
obviously exist for the one lion, but he would not be able to obtain
it
enough to satisfy his needs. The same general point is illus-
fast
trated by the facetious remark that in parts of Texas
grass is so scarce
that the cattle must feed on the run. The necessity for adequate con-
centration as well as for adequate total amount of food is not so
immediately apparent for organisms in which feeding processes are

not directly observed. The total amount of planktonic food in an
oceanic area may be more than sufficient for the nutritive needs of
the clams or oysters living on the bottom, but in order for these
bivalves to survive the food must be concentrated to such an extent
that the volume
of water that they can filter in a day will furnish
enough plankton to fill their daily needs.

The necessity for an adequate rate of replacement of nutritive
materials another aspect of availability in the food relations of a
is
community. The growth of terrestrial plants is dependent not only

upon the concentration of nutrient salts in the soil at the moment of
observation but also upon the replenishment of these nutrients after
the existing supply has been absorbed. The same dynamic de-
pendency exists in all natural situations where the nutrition of one
step in thefood chain depends upon the rate of supply by the previous
step. In many situations the amount of nutriment supplied per unit

of time is than the amount of the material present at any one
larger
moment. The same argument applies to the dependency of a car-
nivore on a prey species. Whether or not the food organism is
adequate in amount depends both on its abundance at the moment
and also upon the rate of replacement of the population.
Ecological Cycle in the Ocean
The of the ocean will be used

ecological cycle in temperate regions
to illustrate the principles involved in the general operation of the
ecosystem as well as many of the relationships discussed in earlier

chapters. The open sea is especially suitable for this purpose since
a sampling can be made in the middle of a large uniform area with
fair assurance that no serious modification by interchange of materials
from surrounding areas of a different sort will take place. As has
been pointed out in earlier chapters, the ocean represents a relatively
uncomplicated biotope compared to many others, particularly those
environment. In a volume of sea water it is possible
in the terrestrial
to measure with accuracy the physical and chemical factors with
which we are concerned and to enumerate the living components with
sufficient precision.
Producers. In the open sea the planktonic plants are the essential
producers since the reduction in light with depth prevents benthic
plants from growing in important numbers in deep water. The main
bulk of the phytoplankton is represented by diatoms, but green flagel-
lates and other types of nannoplankton may also play a significant
role because of their great numbers and the rapidity of their repro-
duction. The preponderance of unicellular plants in the oceanic com-
munity is related to the retardation of sinking rate resulting from small

size and to the increased effectiveness with which widely distributed
cells can absorb dilute nutrients. The zones of diatom growth are
sometimes spoken of as the pastures of the sea, and the phrase current
on land, "All flesh is grass/' has been paraphrased for the ocean as
"All fish is diatoms,"
Since diatoms and other types of phytoplankton represent the pro-
ducer component of the oceanic ecosystem, we shall consider briefly
their typical cycle of abundance and the factors controlling it. In the
temperate ocean a sudden outburst of diatom growth generally occurs

in the spring months (Fig. 13.4), and this is often called the "spring
Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.
FIG. 13.4. Generalized diagram of seasonal cycle of diatom abundance and cer-
tain controlling factors in the temperate ocean.
Producers 475
flowering" or"bloom"an amusing misnomer since these algae are far

removed taxonomically from flowering plants. In the Gulf of Maine,
for
example, the number of diatoms per unit volume in the spring
averages 1000 times greater than the winter population and may be
over 60,000 times greater in local
regions. Hardly has the diatom
population reached its maximum size in March or April than numbers
begin to drop off and characteristically continue to fall to a low level.

An intriguing interplay of ecological forces accounts for these
enormous changes in the numbers of
plant producers. The thorough
stirring of the waterthe upper layers of the ocean during the
in
winter brings about a replenishment of the nutrients to the
euphotic
zone from the deeper levels, but the same vertical movement of water
also the diatoms down to levels below the compensation
carries
depth. As the spring comes on, two things happen: the compensation
depth extends to deeper levels because of the greater intensity of sun-
light, and stirring becomes less because of the progressive stratifica-
tion of the water. As a result the diatoms remain for longer and
longer periods above the compensation depth, until after a time con-
structive growth of the population becomes possible (Sverdrup,
1953). Under favorable conditions diatoms can divide at a rate
greater than once in 24 hours. Exponential growth of the population
ensues and the sea may become green with diatoms within a week.
Certain definite ecological reactions bring the spring increase in
phytoplankton to an end. As diatom growth goes forward, nutrients

are progressively used up until their concentration drops to a point
where the plant cells are starved and either die or form cysts.
all
Furthermore, the great abundance of diatoms has tended to reduce

the transparency of the water and at the same time has provided a food
supply for a new generation of zooplankton. Copepods

and other
herbivores "graze" on the diatoms, arid, as the zooplankton increases
in abundance, the diatom population is consumed at an accelerating
rate.
The diatoms of the open sea characteristically remain at a low

ebb throughout the summer months because the majority of dead
cells decompose and release nutrients again at levels below the com-
pensation depth. Because

thermal stratification prevents deep stir-
ring, the euphotic zone remains depleted of

its nutrients
during the
summer. In the autumn when stronger winds and lower surface tem-
to take place once again, nutrients
peratures allow effective stirring
are restored to the upper layers of the ocean. A rather sudden in-
crease in the diatom population may then take place; this is known
as the autumn "flowering" or "bloom." As the diatoms again grow
in abundance, they reduce the nutrient supply concomitantly for a
time. Following the subsequent complete breakdown of thermal
stratification, wind stirring and reduced illumination stop diatom
growth as the area returns to its winter condition once more. How-
ever, the same deep stirring during the winter brings the nutrient-
rich water to the surface. This annual restoration of deep water to
the upper layers gives rise to the statement that "the sea is
plowed
once a year."
Consumers. When we turn to a consideration of the consumer in
the typical food chain of the open sea, we are confronted with a
situation of special interest because of the nature of the producers.
The simple fact that the plants at the base of the food chain are micro-
scopic in size has far-reaching repercussions in the oceanic ecosystem.

(a) Herbivores. Scarcely any of the larger marine animals are
able to feed directly on diatoms because of their small size. The
menhaden is one of the few fish of commercial importance that can
do so. The gill rakers of this species are usually long and provided
with interlocking hooks that form a fine, sieve-like structure by means
of which the plant cells can be filtered from the sea in sufficient
quantities. The production of the menhaden thus represents a two-

link food chainthe shortest possible and a rarity among the large
animals of the sea in complete contrast to the situation on land. In
the terrestrial environment the availability in respect to size and
abundance of trees, grass, and other plant producers is such that deer,
elk,sheep, and other large ruminants can feed directly on them. Thus
short food chains are common on land.
Since most of the larger animals in the sea are unable to use phyto-
plankton as food, they must depend for their nutriment upon "middle
men" or "key industry animals" of intermediate size. On the sea
bottom many mollusks and other sessile invertebrates possess feeding
mechanisms that enable them to filter the smaller plankton ic forms
from the water. In the open sea planktonic crustaceans, especially
copepods and euphausiids, fill this niche. Copepods can filter out the
smallest diatoms by forcing a current of water through a meshwork of
bristles attached to a special set of appendages. We might think of
these small crustaceans acting as miniature harvesting machines,
packaging the microscopic plant cells. The filter-feeding zooplankton

incorporate the substance of the diatoms and represent parcels of
food of sufficient size for larger animals (Fig. 13.5).
When
the importance of copepods as essential middlemen in the
economy of the sea was recognized, special attention was directed

toward the factors controlling the food supply of these animals.
Consumers 477
However, food dependencies are difficult to determine, and several

divergent views have been held as to the critical relations. Since
observations at sea indicated that diatoms were often abundant when
copepods were scarce and vice versa, some investigators argued that
no direct food dependency occurred. The quandary of this apparent
contradiction represents the danger of generalizing from isolated ob-
servations or from statistical tabulations of the occurrence of animals
and plants without due regard to the dynamic aspect of their growth
and life
cycles. If diatoms are scarce at a given point where
copepods are abundant, does it mean that the copepods have just
finished consuming the population, or does it mean that copepods are
primarily dependent upon some other source of food?
Clarke. 1943
FIG, 13,5. Simplified representation of main ecological cycle in the

sea.
As data on this problem were gathered, two theories emerged.

One was the theory of animal exclusion proposed by Hardy (1936)
that high concentrations of phytoplankton are harmful to zooplankton
and, therefore, that water masses containing large numbers of diatoms
are actively avoided by copepods. Exclusion was originally believed
to have taken place chiefly in the vertical direction and thus to curtail
the upward phase of the diurnal migration of the zooplankton. Hori-

zontal currents moving differentially at various levels would secondar-
ily bring about a patchiness in horizontal distribution. Subse-
quently, the scope of the theory has been extended to include other
types of exclusion of one population by another. Dense diatom con-
centrations might be inimical to copepods either mechanically or
chemicallypossibly through antibiotic effects. Such effects have

been demonstrated for oysters and cladocerans in certain instances
as described in Chapter 8.
At about the same time, Harvey and co-workers ( 1935 ) came for-
ward with the theory of grazing, according to which copepods are
believed to reduce the diatom population locally by the intensity of
their feeding activity. At first sight these theories seemed mutually
exclusive, but more extensive investigation has shown that the in-
fluences involved in both may be in operation simultaneously without

conflict. Copepods depend upon suitable abundance of diatoms
may avoid thick concentrations of diatoms and, by

for food, but they
grazing around the margins of diatom populations, may accentuate

the patchiness in the distribution of the latter. The timing of the
supply of diatom food in relation to the appearance of new broods of

copepods also critical (Clarke, 1939).
is Investigation of the pro-
duction of Calanus finmarchicus in the Clyde Sea area gave evidence
that those broods of this copepod which hatched during periods of
diatom scarcity failed to reach maturity, whereas broods appearing
when suitable diatom food was available developed successfully
(
Marshall et al., 1934).
Another possible source of food for copepods as the key herbivores
in the oceanic food cycle is the nannoplankton. Lohmann drew at-
tention long ago to the abundance of such representatives of the
nannoplankton as the green flagellates, peridinians, other protozoans,

and certain bacteria (Sverdrup et al., 1942). These organisms may
occur in concentrations of millions of cells per liter, and, since they
are able to divide several to many times a day, they represent a large
and rapidly replaced potential food source. The fact that certain
copepods at least can sieve these extremely small organisms from the
sea is shown by the successful culturing of Calanus on flagellates 1.5
to 3 p. in diameter (Raymont and Gross, 1942); and Calanus can de-
rive at least some nourishment from bacteria (Clarke and Gellis,
1935). Nannoplankton is undoubtedly of particular importance in

the feeding of young stages.
Dissolved organic matter and fragments of dead organisms as de-
tritushave also been suggested as important food sources for primary
consumers in the sea. Analyses have shown that more than three
Consumers 479
times as much organic matter is
present in dissolved form as is repre-
sented by the phytoplankton. Putter claimed to have evidence that
this material was used
extensively for the nourishment of marine
investigation of the problem now indicates
animals. However,
strongly that, if any dissolved organic matter is available to multi-
cellular organisms, it forms an
insignificantly small fraction of their
requirement (Krogh, 1931). We know that some dis-
total nutritive
solved organic matter in the water can be used by many kinds of
bacteria and other saprophytes, but a
large fraction of it is resistant
even to bacterial attack, and this portion
represents an unrecoverable
loss for the oceanic ecosystem.
Organic detritus appears in the water directly from the disintegra-

tion ofdead plants and animals and is also stirred up from muddy
bottoms. This material occurs in particles of all sizes, and even the
most minute may be removed from the water by animals possessing
mucous filtering apparatus. Echiuroid worms can obtain material
with dimensions as small as 4 m/x (MacGinitie and MacGinitie, 1949),
and Daphnia magna is
capable of taking in detritus of colloidal size
in sufficient quantities for growth (Gellis and Clarke, 1935). De-
tritushas been shown to form a significant portion of the food of at
least certain filter-feeding bivalves (Coe, 1948), and no doubt it
does likewise for many kinds of zooplankton. The nutritional de-

pendencies of copepods and other key primary consumers are there-
fore very complex, but directly or indirectly their food is ultimately
derived from the organic material synthesized by the diatoms as the
chief plant producers of the open sea.
(b) Carnivores. Since the herbivores in the typical community of

the open sea are mostly small filter feeders, the primary carnivores-
representing the third link in the food

chain are confronted with the
problem of obtaining food from prey species that are often less than
a centimeter or two in length. Nevertheless, certain sizable fish,
such as the mackerel, herring, and shad, are able to feed directly upon
the copepods and other primitive crustaceans of the plankton. In-
the animals in the sea including the bask-
terestingly enough, largest
ing shark and the
whalebone whales are also able to live upon the
tiny zooplankton.
The great sheets of frayed whalebone attached to
the upper jaws of these whales serve as gigantic sieves (Fig. 13.6).
When the whales are feeding, they take in huge mouthfuls of the sea
and by means of their massive tongues force the water out through
the whalebone. The copepods, euphausiids, and other types of zoo-
within the mouth cavity are then gulped down.
plankton retained
The fact that the food chain of these whales is composed of only three
FIG. 13.60. North Pacific right whale, showing adaptation of mouth for plankton
feeding. A cluster of barnacles can be seen on the lower lip.
Photo J. T. Ruud
FIG, 13. 6b. Stomach of southern right whale split open, showing mass of euphau-
siids ("krill") that had been eaten.
links,and hence is relatively efficient, no doubt accounts for the suc-

cessful maintenance of such large, warm-blooded mammals in the sea.
Most of the sizable marine carnivores are the result of longer food
chains. Many adult fish, such as cod, haddock, halibut, and flounders,
Productivity of the Ecosystem 481
are bottom feeders and
depend upon worms, mollusks, crabs, and
other benthic animals for food. As mentioned above, certain of the
mollusks are primarily herbivores, but other
types of benthos are
secondary or tertiary carnivores, scavengers, or mud feeders. Many
worms and echinoclerms pass large quantities of mud through their
intestines, extracting nutriment from its components. Other types of
fish feed
primarily upon smaller members of their own class; the
growth of a shark, for example,
may represent a food chain of 5 or
more links (Fig. 13.5).
(c) Decomposers and Transformers. The ecological cycle in the

large group of decomposers and
sea is completed by the activities of a
transformers. Material excreted by living organisms and the bodies
of dead plants and animals from every link in the food chain undergo
decomposition as the result of the activity of certain groups of marine
bacteria. Dead tissue is rendered soluble and decomposed to in-
organic form. The
material is then attacked by the transformers, as
represented by other types of bacteria, and converted to forms suit-

able for the nutrition of green plants once more. Living members of
the oceanic community are thus necessary to carry out each step in
the chemical regeneration of the organic material. However, before
the resulting nutrient salts can be assimilated again by the phyto-
plankton, the water containing them must be restored by currents to

the euphotic zone where light is sufficient for photosynthesis. On
certain submarine banks, such as Georges Bank, and in regions of
permanent upwelling, such as along the western coast of the United

States, physical regeneration may take place almost continuously.
In other regions the deep stirring of winter is required before renewed
development of phytoplankton can initiate constructive growth in the

oceanic ecosystem once more.
PRODUCTIVITY OF THE ECOSYSTEM

In the sections the functional interdependencies be-
previous
tween the various living and non-living components of the ecosystem
have been discussed. We shall now consider relations of productivity
both for the individual links in the food chain and also for the eco-
system as a whole. When reliable methods of measurement
have
been developed, the productivity of one type of organism may be
compared with another, and the productivities of different communi-
ties or regions can be evaluated. From the practical point of view it
isdesirable to ascertain whether the production of a given area is as
great as it could be. In order to

determine whether an area is being
to the number of organisms that can be harvested. In addition to
the yield taken
by man other material may be removed from the area
by active or passive emigration or by the withdrawal of organic matter
from circulation through the formation of The second con-
deposits.
cept of productivity is the material removed from the area per unit
time, and includes the yield to man, organisms removed from the
ecosystem by emigration, and material withdrawn as organic deposits.

When the modern ecologist enters the area, he views the situation
in a different light.
still He should, of course, be aware of the num-
bers and types of organisms present, and of the extent of their removal
from the area by man and in other ways, but his attention is directed
chiefly to the dynamic aspect of the growth and interdependency of
the inhabitants as members of a functioning system. The ecologist
is concerned with natality and mortality in the various components
of the population and with rates of metabolism and growth. He in-
quires not only into the different types of machinery in the factory
but also into the speeds at which the wheels are turning. The third
concept of productivity is the production rate, or the rate at which
growth processes are going forward within the area./ Each of the
fundamental concepts of productivity will be considered separately
in further detail.)
Standing Crop
Measurements of the standing crop reveal the concentration of
individuals in the various populations of the ecosystem. This in-
formation is essential for judging whether the degree of crowding in
each species is exerting a harmful or a beneficial effect.. Further, in
the study of the dependence of one species on another the enumera-
tion gives a measure of the intensity of predation and of the avail-
size of the standing crop is also of
ability of the forage species. ^The
vital concern in relation to the exploitation of natural populations by
man. ,The abundance of the plant or animal influences the efficiency
with which the enterprise is carried forward. However, since a
knowledge of the standing crop gives no information on its replace-
ability, other aspects
of productivity must also be taken into account
for intelligent use of natural resources of this kind and for a main-
tained harvest.
Relation to Population Growth. The relation of the size of the
standing crop to the rate of growth of the population will be con-
sidered for the three types of situations illustrated in Fig, 13.8. In
the first a rapid increase in population size is indicated, leading to a
overexploited or underexploited, abundance and growth must be

measured in accurate terms, but before this can be done the various
aspects of productivity must be clarified.
Concepts of Productivity
The three fundamental concepts of productivity are: (1) standing
crop, (2) material removed, and (3) production rate (Fig. 13.7).
Failure to distinguish these concepts has resulted in much confusion
in attempts to measure and to discuss the production of animals and
plants in different areas.
1. Standing crop
Carnivores |
Herbivores
Green plants
1
Pyramid of numbers
D Yield to man Yield
Emigration
Supply
D Deposits
2. Material removed 3. Rate of production
FIG. 13.7. Concepts of productivity. (Clarke, 1946.)
When the early naturalist entered a new area, his interest was at
first centered
upon the identification and description of the species
present. With the growth of the ecological point of view the observer
added enumeration of each species and soon built up a
to his task the
picture of the differences in abundance of the various members of

the food chain, leading to the concept of the pyramid of numbers.
The standing crop is the abundance of the organisms existing in the
area at the time of observation; it
may be expressed as number of
individuals, as biomass, as energy content, or in some other suitable
terms.
When the hunter, fisherman, or forester enters an area, he sees the
situation from a different viewpoint. The attention of such practical
ecologists is directed not so much to the fauna and flora as a whole as

to the number of organisms that can be harvested. In addition to
the yield taken
by man other material may be removed from the area
by active or passive emigration or by the withdrawal of organic matter
from circulation through the formation of The second con-
deposits.
cept of productivity is the material removed from the area per unit
time, and includes the yield to man, organisms removed from the
ecosystem by emigration, and material withdrawn as organic deposits.

When the modern ecologist enters the area, he views the situation
in a different light.
still He should, of course, be aware of the num-
bers and types of organisms present, and of the extent of their removal
from the area by man and in other ways, but his attention is directed
chiefly to the dynamic aspect of the growth and interdepend ency of
the inhabitants as members of a functioning system. The ecologist
is concerned with and
natality mortality in the various components
of the population and with rates of metabolism and growth. He in-
quires not only into the different types of machinery in the factory
but also into the speeds at which the wheels are turning. The third
concept of productivity is the production rate, or the rate at which
growth processes are going forward within the area./ Each of the
fundamental concepts of productivity will be considered separately
in further details
Standing Crop
Measurements of the standing crop reveal the concentration of
individuals in the various populations of the ecosystem. This in-
formation is essential for judging whether the degree of crowding in
each species is exerting a harmful or a beneficial effect.. Further, in
the study of the dependence of one species on another the enumera-
tion gives a measure of the intensity of predation and of the avail-
size of the standing crop is also of
ability of the forage species. ^The
concern in relation to the exploitation of natural populations by
vital
man. ,The abundance of the plant or animal influences the efficiency
with which the enterprise is carried forward. However, since a
knowledge of the standing crop gives no information on its replace-
ability, other aspects
of productivity must also be taken into account
for intelligent use of natural resources of this kind and for a main-
tained harvest.
Relation to Population Growth. The relation of the size of the
standing crop to the rate of growth of the population will be con-
sidered for the three types of situations illustrated in Fig. 13.8. In
the first a rapid increase in population size is indicated, leading to a
Dynamics of the Ecosystem
high equilibrium level for the standing crop. In the second situation
the growth of the population is much slower, but the same ultimate
size of standing crop is attained. In the third situation the same
.
period of time is required to reach equilibrium as in the second, but

a lower equilibrium level is established. \ If a census of the standing
crop taken before the populations have reached their maximum

is
equilibrium levels, the larger standing crop will clearly occur in the
area in which the rate of production has been the greatest. ,
Usually,
Equilibrium standing crop
Growth
and mortality
Rapid
increase
Short season Long season

Time
FIG. 13.8. Relation between size of standing crop and rate of growth of popula-
tion. (Clarke, 1946.)
however, the observer will enter a natural area long after the initial
growth of the population has been completed and will measure the
standing crop at or near its equilibrium level. At this point the
magnitude of the standing crop is not an index of growth rate. The
size of the population at any moment is the result of the accumulation
of production minus the amount of destruction which has taken place
up to the time of observation.

In the initial growth of the
population the length of life of the indi-
viduals influences the steepness of the population curve because, the
longer each individual lives, the faster will the total numbers increase.
Once equilibrium has been reached, however, individual length of
life has no effect
upon the size of the standing crop. As discussed
in Chapter 9, natality is equal to mortality at equilibrium, or A = M.
If 50 animals are born per unit of time, 50 animals will die in the same
period, and it makes no difference whether those animals have

lived
5 days or 5 years. Similarly, the size of the population is not de-
termined by the magnitudes of A and Af, As indicated in the dia-
gram, a population with a large standing crop may exhibit a low
Determination of Equilibrium Level 485
natality and a
low mortality, whereas another population of smaller
equilibrium size may display high birth and death rates. In anal-
ogous fashion the level of water in a tank (the standing crop) gives
no clue to the rate of inflow (natality) or the rate of outflow (mor-
tality). A
high level of water in the tank may be maintained by a
trickle of water running in and out, or, in another instance, a low
water level may exist with a large inflow and a large outflow.
Determination of Equilibrium Level. What then does determine
the magnitude of the standing crop at equilibrium? The total
biomass of organisms inhabiting an area always tends to increase.
This is due to the numerical increase of the population and to the
growth in size of the individuals, or both. In some kinds of organ-

isms, such as fish or trees, the individuals continue to grow larger
but in others, such as rodents and insects, full size is
all their lives,
attained as soon as adult condition is reached. Although the number

of trees in a forest stand tends to decrease due to competition, the in-
crease in size of the individuals, for a time at least, more than makes
up for the loss in numbers so that here also the total biornass of the
population tends to grow larger. Since the size of the standing crop
is not limited
by factors intrinsic within the population, the increase
in total biomass must be stopped by external factors. The biomass
of the population will
eventually reach the maximum size possible
for the circumstances regardless of the speed of increase. \ The maxi-
mum standing crop that can maintain itself
indefinitely in an area
is
spoken of as the carrying capacity of the area. }
The ultimate limit to the growth of the population is determined

by the supply of nutrient materials and energy,^ If the supply of
these needs is no other factor interferes, the population
large and if
will grow If food and energy continue to be pro-
to a large size.
vided at a rapid rate, the large population can be continuously main-
tained. A clear example of such a situation is found where a continu-
ous upwelling of deep water supplies nutrients for the growth of the
marine organisms in the area. Conditions of this sort account in part
for the large carrying capacity of certain fishing banks.
When a population lives up to the limit of its food supply, it is
living dangerously. Should the food supply he curtailed or should

the population increase still
large-scale destruction of the
further,
inhabitants would ensue. If
populations of animals were checked by
food supply only, critical conditions would be widespread because we
know that animals can destroy their food supply faster than it can
grow. The essence of this idea in relation to man was pointed out by
an early "ecotogist" by the name of Malthus. Because *of the in-
efficiencies of feeding and digestion and because of the great amount
^ energy used for the various life
processes, each animal destroys
a
rrtMch larger mass of food than its
growth increment.
These facts clearly indicate that animals could them-
quickly eat
selves out of house and home.
Temporarily unchecked populations
of insects or rodents sometimes increase to such numbers that every
green thing within the area of their abundance is destroyed. Plagues

of locusts in the Middle East have been known to consume all vegeta-
tion over areas as great as 600 sq km. Even deer, which reproduce
relatively slowly, when protected from other checks, soon multiply to
a point where the population needs more food than can grow within
their home range.
When one realizes the fact that animals are able to destroy their
food supply faster than it can
grow, the question arises why the
herbivores of the world have not destroyed all the vegetation, and
why carnivores have not killed all of their prey and thus brought
about their own destruction. One reason is that climate and other
conditions vary locally, so that the animals do not find conditions
suitable for maximal increase everywhere at once within their geo-
graphical ranges. If a local population does completely destroy itself

by starvation, the area can be repopulated from neighboring regions.
Another reason that extinction is
usually avoided is that other factors
frequently stop the increase in the population before the entire food
supply is exhausted. For example, excessive grazing tends to change
the species composition of the vegetation, and this condition, recog-
nized as range deterioration, curtails the growth of the herbivores.
Increase of some animals is checked by accumulation of metab-
olites, by disease, or by intensified predation, as discussed in previous
chapters, long before a food shortage is threatened. In special in-

stances lack of breeding sites may be the chief factor limiting popula-
tion growth. A striking illustration of the operation of this factor is
provided by studies of the wood duck made at the Great Meadows
Wildlife Refuge in Massachusetts. In an area of 120 hectares, in-
cluding a large swamp, 13 tree holes were located that appeared to

be suitable for nest sites in regard to size, shape, and location, and 6
of thesewere occupied by breeding ducks. After wildlife managers
had put up 65 nesting boxes in the area, ducks built nests in 43 of
these, thus revealing the extent of the previous underpopulation; the
number of wood ducks had been much smaller than could have, been
supported by the supply of food and other necessities in the area.
The tendency for some species to limit themselves to home ranges
or territories and thus avoid excessive concentration of the population
Regional Differences 487
has already been
pointed out. Furthermore it should be noted that
the biotic potential of
prey species of animals is usually much higher
than that of the predatory This relationship helps to retard
species.
the growth of the more remote links of the food chain. As
prey be-
comes scarcer, predators have more and more difficulty in finding
them so that their rate of feeding is automatically reduced. Often
predators turn to other species at this time, relieving the pressure on
the species originally used as a source of food.
This type of check also applies to man as a predator to some extent.
When deer become scarce, sportsmen may voluntarily desist from
huntinglTufficiently to allow the population to recover. In some man-
aged fish
ponds unrestricted fishing by hook and line can be permitted.
After the number of fish in the pond has been greatly reduced by the
anglers, sir much food is available for the remaining fish that they
are too well fed to take bait. When the depletion of an oceanic
fishery reaches the point where fishing-for this species is no longer

profitable, fishermen turn to other species. In such instances, if the
population has not been reduced below the point of recovery, as dis-
cussed in Chapter 9, and if the habitat has not deteriorated in the
meantime, the population may be able to return to its former
abundance.
Regional Differences. The can be sup-
total quantity of life that
ported in an area at its full carrying capacity may be made up of a

few individuals of a large number of species or many individuals of a
few species. Generally speaking tropical communities tend to ex-
hibit the former condition and non-tropical communities the latter, but
there are many exceptions. The fauna and flora of tropical biotopes
are typically very rich in species but an observer often has difficulty
in finding more than one or two specimens of each kind of animal or
plant. The diversified nature of tropical vegetation is illustrated by

the results of a transect 2 m wide and 150 m long run from the edge
to the center of Castellow Hammock near Miami, Florida, in which
111 species of flowering plants were found belonging to 98 genera and
48 families (Phillips, 1940).
In contrast, the vegetation in high latitudes may be very monotonous
from the point of view of species composition. The great spruce
forest extending across the North American continent and Eurasia con-
sists of but a few species of trees with a relatively small number of
subordinate plants. The same general contrast in relation to numbers
of species and individuals occurs among the land animals and also in
the biota of aquatic areas (Table 21). Tropical marine areas, for
example, support hundreds of species of fish with relatively small
TABLE 21
GENERAL DECREASE IN ABUNDANCE OF SPECIES WITH INCREASING LATITUDE

IN NORTH AMERICA As INDICATED BY REPRESENTATIVE DATA FROM
VARIOUS SOURCES
Approximate Number of Species

Florida Massachusetts Labrador Baffin Land
Beetles 4000 2000 169 90
Land snails 250 100 25
Mollusks (tidal zone) 425 175 60
Reptiles 107 21 5
Amphibia 50 21 17
Fresh-water fish 75 20 1
Coastal marine fish 650 225 75

Flowering plants 2500 1650 390 218
Ferns and clubmosses 70 31 11
numbers of each, whereas in northern waters many fewer kinds of fish

are found but some of them exist in huge populations. It is no acci-
dent that most of the large commercial fisheries are located in high
latitudes. Enough cod are found among the Lofoten Islands off Nor-
way during the early spring to keep 4000 fishing vessels busy, and 100
tons of cod may be taken with one cast of the purse seine. Many
other illustrations of this generality may be found in faunal and floral
lists for regions of contrasting latitude, such as those of Russell
(1935).
Although this
typical difference between tropical and non-tropical
populations has been recognized for a long time, opinion differs as to
its cause. One explanation lies in the fact that, since climatic condi-
tions are more favorable at low latitudes, many different kinds of
species can survive; and, because the total quantity of life in an area
is limited, each
species can be represented by only a few individuals.
Furthermore, at high temperatures, with higher rates of reproduction,
mutations probably occur more frequently so that in the course of
evolution a larger variety of species has originated and more mutants
would be expected to survive under the favorable conditions of the
tropics. At high latitudes the reverse situation exists, and in addition
the repeated expansion of the polar climate may have reduced the
number of species as well as the time available for evolution. The
relatively few species that do survive in colder regions find little com-
petition and hence can develop large populations that exploit the en-
vironment to the limit
In addition to this general reciprocal relationship between abun-
dance of species and individuals, the standing crop in marine environ-
ments tends to be
absolutely larger in high latitudes than in regions
nearer the equator. This difference is particularly
prominent if the
standing crop in the tropics, which often remains uniformly small
throughout the year, is
compared with the standing crop in temperate
or polar regions at the height of the growing season following the
winter replenishment of nutrients,
^pr instance, the standing crop of
zooplankton in the antarctic summer was observed to be about 10
times greater than that in the tropical Atlantic. Again many excep-
tions exist in local areas as in the Great Barrier Reef lagoon, where
the standing crop was found to be as great as in temperate coastal
regions. Under tropical conditions organisms grow faster and, after

death, their tissues are decomposed and nutrients regenerated at a
more rapid rate, but as already explained this difference in
growth
rate does not in itself account for the difference in the standing crop.
Part of the explanation of the latitudinal difference in total abun-
dance of marine life is the fact that at low
temperatures the metab-
olism of poikilotherms generally is much reduced. This means that
a larger total number of organisms can be supported on the same
amount of food. However, regional differences in metabolism are by
no means simple and clear cut, as pointed out by Riley, Stommel, and
Bumpus, (
1949 ) . Evidence exists that the rate of supply of nutrients
to the surface in temperate and polar regions is usually greater than
in the tropics owing to the nature of the vertical circulation of the
ocean. Nutrients may thus be supplied more rapidly in relation to
their use in the cooler water, with the result that a larger standing
crop can be supported. However, in tropical regions with exception-

ally great upwelling
as along the Peruvian coastan enormous stand-
ing crop of plankton exists and supports the food chain leading to
the production of guano, as described in Chapter 8. These regional
differences in abundance of marine organisms are discussed in further
detail by Sverdrup et al. (1942, Ch. 19).
Material Removed
Various aspects of the material removed from an area, as the second
concept of productivity, are of interest to both the theoretical and the
practical ecologist. Some may believe that the yield to man is the
only component of the material removed that concerns farmers, fisher-
men, hunters, and foresters, but losses from the area in the form of
emigration and organic deposits must also be considered. For the
purpose of maintaining the yield to best advantage other concepts of
also be understood and applied.
productivity must
In subsistence
490 Dynamics of the Ecosytem
farming or hunting in which products are not taken to the market, the
human agent may be regarded as an integral part of the local eco-
system; but in most farming, fishing, hunting, and forestry operations
a large part of the organisms harvested is carried away, and hence
the material of which the plants or animals are composed is perma-
nently removed from the area.

Permanent emigration of animal or plant life similarly represents a
loss of material from the ecosystem.
^
Ôrganisms may leave the area
by their own locomotion
1
'
or they may be carried away by wind or

water currentsA When the activity of natural predators results in the
one-way transfer of organic material from one ecosystem to another
as when a fish hawk removes fish from a pond the loss of material
from the ecosystem under consideration is also to be included in this
category. (Another type of loss is represented by dead organic matter
that is buried in the soil or in the mud of an aquatic biotope in such
a way that it is
functionally removed from the system,, Also in-
cluded ismaterial that has gone into an inert chemical form and thus
is
essentially unrecoverable during
the period under consideration.^
If a community is to maintain itself, material in some form must be
supplied to it in an amount at least equal to the total amount of

material removed from the area in the various ways discussed. The
supply in the form of nutrients and immigrants usually must be con-

siderably larger than the removal because of the inevitable loss of
energy and of material from the system at each level in the food chain.
Ordinarily, man's harvest, which is only one portion of the total
amount of material removed from the area, is
very much smaller than
the total supply to the area from all sources. The ratio of yield to
supply gives one measure of the efficiency of the use of the area.
Such a measure can show whether a needlessly low utilization is
taking place or overexploitation under way.. is
The yield to man may be improved

in one or more of several
ways.
Although practical limitations must be considered in each specific
situation, a summary of the theoretical possibilities is desirable as a
basis for realistic procedure. A maintained yield can never average
larger than the supply. However, the natural supply of materials
entering the area may be augmented artificially by adding fertilizers
or by stocking; and the amount of materials removed or lost in
ways
other than man's harvest may be curtailed. In addition, the degree
of crowding of the desired species may be adjusted to the optimal
level, and the competitors of this species may be removed.
The foregoing points will be illustrated by a schematic considera-
tion of the interdependencies of a natural pasture from which it is
desired to obtain the maximal yield of
sheep (Fig. 13.9). Increased
growth of the plant producers will result from the addition of fer-
tilizers, but it is essential that this added supply of nutrients reaches
the grass rather than the undesirable weeds. The grass supports a
flock of sheep but it usually also
supports a population of field rodents.
If the rodents are excessively abundant, a reduction in their number
would allow a larger portion of the grass to be consumed by the sheep.
Fertilizer Man
IJ Coyotes
FIG. 13.9. Schematic representation of the relation of man's harvest from an area,
such as a natural pasture, to the supply of materials and to the reduction of the
product in various ways.
Often the best method of controlling the rodents is to allow the pres-
ence of predators such as coyotes, but should the coyotes become too
numerous they would compete with man for the sheep and their num-
bers might have to be reduced. A suitable balance must be main-
tained among all the members of the ecosystem. The population of
the to be harvested should also be adjusted to the level of abun-
species
dance at which
its
growth rate will be as great as possible without
of the ma-
disturbing the equilibrium of the community. Study
terials entering and leaving the area should reveal whether the area
is
beingmanaged to the best advantage from the point of view of its
desired use. The maximum potential yield is the largest maintained
harvest that can be removed from an area under the best population
and environmental conditions.
An of great practical importance is the yield per
aspect of the yield
unit effort. This may be measured in terms of man power, cost, or
other basis, and depends among other things upon the size of the
standing crop. Thus one aspect of productivity affects another
aspect. These relationships may be illustrated by data for the total

yield ofhaddock caught on the Icelandic fishing grounds in compari-
son with the catch per unit effort (as measured by the landings per
day's absence from port) and
the total hours of fishing by British
trawlers (Fig. 13.10). The total yield of haddock declined slowly
1910 1915 1920 1925 1930 1935
1905 1910 1915 1920 1925 1930 1935

FIG. 13.10. (Upper) Total catch of haddock from Icelandic Fishing Grounds by
all countries. (Lower) Catch per day's absence from port (solid line) and total
hours of fishing (broken line) by British First Class Trawlers. (Modified from
Russell, 1942, The Overfishing Problem, Cambridge Univ. Press. )
from 1905 to 1915, and the catch per unit effort declined regularly
after 1911. During the war period 1914-1918 the haddock population
had an opportunity to recover, and, with the resumption of fishing
by the British trawlers in 1919, the catch per unit effort reached an
all-time high. The total yield reached a peak in 1920 and a some-
what higher peak during 1927-1929, owing in part to the increase in
fishing time and in part to the adoption of a new type of trawl.
From
1924 to 1937 the hours of fishing were doubled, but the catch per unit
effort exhibited a downward trend to a value in 1937 less than one
quarter of that obtained in 1919. The explanation is found in the fact

that the standing crop of haddock in Icelandic waters became progres-
sively depleted, as a result of

the intense fishing, and consequently a
Production Rate 493
smaller and smaller catch was obtained from each day of fishing. A
similar study of the
drop in the catch per unit effort following de-
pletion of the stock has been made for the Antarctic whales Ruud by
(1952).
Production Rate
A knowledge of the standing crop of an area or of the yield from

the area does not give a complete picture of
productivity; in addition
the rates at which the different constructive and destructive processes
are going forward within the area must be considered. Sincefthe
plant producers, herbivores, carnivores, and other components of the
ecosystem are all transforming energy and material simultaneously,
the evaluation of production rates is extremely complex. For an un-
derstanding of the interdependencies of the entire ecosystem we

should know the production rate for each of the trophic levels, that is,
the amount of material formed by each link in the food chain per unit
of time per unit area or volume, and we must distinguish between
gross and net production. To disentangle these relations the analyti-
cal approach will again be adopted.
Let us consider first a very much simplified case such as an isolated
pond in which no gain or loss of materialsoccurs during the period
under consideration. The transformations going on within such an
area are indicated schematically in Fig. 13.11. At the left of the dia-
gram the energy represented by the standing crop at the beginning

of the period is indicated. To keep the problem simple the food
chain is considered to have only three links. The size of the rectangles
for the green plants, herbivores, and carnivores are progressively

smaller to indicate the typically reduced energy content of the stand-
ing crop as a pyramid of biomass. In the main part of the diagram

the energy absorbed at each level of the food chain and its transfor-
mations are shown.
The largest rectangle at the bottom represents the total amount of
incident light reaching the area during the period under consideration.
Not all of the radiation is absorbed by the green plants. In the
aquatic environment the lion's share of the light energy is absorbed
by the water, and in the desert most of the sunlight falls on bare
ground. In regions where the vegetation forms a complete cover,
however, a large fraction of the incident radiation is absorbed. Thus
the various rectangles of the diagram are not drawn to scale but
simply illustrate the relations of the energy transformations among

the various components. Of the light energy actually absorbed by
photosynthesis and appears
the plants only a small fraction is used in
as carbohydrate formed since the process is far from 100 per cent ef-
ficient
especially under natural conditions.
The energy of the carbohydrate formed represents the gross plant
production and therefore the gross production of the producer com-
ponent of the ecosystem. Not all the energy present in the photo-
synthetic product takes part in the realized growth of the plants since
Initial standing crop Energy transformations during period

i
D
Herbivores
Growth fP2 ) -
Assimilation (f\)
'^Decom-
Green plants Respired
posed j
Plant growth i
Carbohydrate formed (Pi)~
Energy used in photosynthesis
* Absorbed light *
^ Incident light *.
FIG. 13.11. Energy transformations during unit of time in an idealized ecosystem

with no net increase or decrease (not drawn to scale). Pi gross production;
P, = net production; K respired; D
~ decomposed. Cross hatching indicates
total portion of food organisms killed by consumers; a fraction, I/, of this is un-
assiinilated. (Clarke, 1946.)
the growth process also is not 100 per cent efficient. The accompany-
ing catabolism of the plant is measured by the amount of respiration.

Actual plant growth is
represented by an amount of energy equal to
the total carbohydrate formed minus the respiration and is designated
as the net plant production. Since the formation of additional links
in the food chain depend upon the elaboration of organic compounds
by the green plants, the photosynthesis of plants in the ecosystem is
sometimes referred to as "primary production" or "total production,"
*
but the more specific terms used here are preferable.
Part of the plant material formed in the ecosystem Is consumed by
herbivores, and part dies from other causes and undergoes direct de-
composition. Only a portion of the plants designated as "consumed"
Production Rate 495
is
actually assimilated by the herbivores, for a certain amount of food
that is eaten remains undigested and frequently more food is destroyed
than actually eaten.
is Sometimes this unassimilated portion is the
greater of the two. A colony of beavers, for example, may cut down
large numbers of aspens, but the bark that they consume is obviously
only a very small fraction of the total biomass of the trees. Similarly,
as mentioned earlier, the grazing activities of copepods often result
in the destruction of a quantity of diatoms far greater than that actu-
ally assimilated.
We
have assumed that in our simplified ecosystem the new plant
growth was entirely removed by being consumed or by decomposing.
Under natural conditions the net plant production per unit of time
would rarely be exactly balanced by the loss due to herbivore feeding

and to death of the plants from other causes. Any portion of the net
of the period constitutes a net in-
production remaining at the end
crease to the standing crop. Under other circumstances the amount
of foraging and of other destruction may exceed the amount of new
a net decrease will take
growth during the period, with the result that
place in the standing crop.
We
can the same line of reasoning to an analysis of the energy
apply
transformations at the herbivore level. The plant material
assimilated
This amount, less that
is
equal to the gross herbivore production.
is a measure of the actual growth
lost in respiration, of the herbivores,
or the net herbivore production. At the end of the time unit one
has died and decomposed, and all
portion of the new herbivore growth
of the other portion may have been destroyed by carnivores, as indi-
cated in the diagram-or part of the growth increment may remain as
a net increase in the population.
At the third trophic level the primary carnivores often eat only part
of the prey that they kill, arid, of course, only a fraction of the material
is
digested and assimilated.
The gross carnivore production is equal
to the total assimilation, and the net carnivore production is equal to
the actual growth increment of these animals. Since the primary
carnivores are at the top of the production pyramid in this assumed
three-link food chain, no part of the carnivore production is consumed
and goes back into the system after
by predators, but all decomposes
death. In most natural situations secondary and tertiary carnivores
and other high-level consumers would also be present. The same
typesof relation apply to the gross and net production of these further
links in the food chain.
all trophic levels at
example the standing crop at
In this simplified
the end of the period is the same as it was at the beginning of the
496 Dtjnamics of the Ecosystem
period, and there has been no yield or other material removed.
Nevertheless, productive activity has been going onthe wheels of the
factory have been turning but the products of the factory have been
taken in again as raw materials. An analytical scrutiny of the eco-
system reveals what processes are going on and at what rates, and
how they are interdependent. A basis is also obtained for judging the
productive capacity of the area and for deciding on the best pro-
cedure for the continued or improved use of the area for human needs.
yield to man is
In an unfished lake, for example, the zero. If fishing is
to be initiated, information on the production rates of the various

living components of the lake is desirable in order to understand the
factors that would control the size of the fishery that could be per-
manently supported by the lake.

In areas in which rates of production are higher than rates of de-
struction a net increase in one or more of the components will occur.
The surplus material may be removed by man, thus constituting a
yield, or it may be removed in other ways described in the previous
section, orit
may be permanently added to the system. In the last
case an increase in the organic components of the ecosystem will be
experienced, but such an increase can take place only to the extent to
which an equivalent supply of materials is added to the area. The
amount of organisms and of nutrients in a lake, for example, are fre-
quently seen to increase in this way over a period of time with the
result that the lake becomes more eutrophic; the process is known as
eutrophication. Similar increase of the living and non-living com-
ponents of the ecosystem occurs in other types of habitat.

Turnover. / In our discussion of production rate nothing has been
said thus far about ^the length of the period considered. The time
required for each type of organism to complete its growth, to die and
decompose, and to start the cycle over again is known as its period
of turnover.* The length of the turnover period usually differs widely
for the organisms at different trophic levels in the production pyramid,
and may differ for the same level in different situations and at differ-
ent seasons of the year. Herbaceous plants in a temperate terrestrial
area usually have only one turnover per year, whereas the phyto-
plankton of an aquatic area may turn over within a few days or a few
weeks. In the latter instance the same material may be used again
several times during the year, and it would thus have little meaning
to add up the increments of growth for the whole year in an attempt
to reach a "total" annual value. If the growth of the plants is meas-
ured as energy, it is permissible to sum up the amount of energy

which has been transformed during the year since the energy can be
Efficiencies 497
used by the plants only once. A
comparison may thus be made be-
tween the annual incident radiation and the content of the
energy
plants produced. In the organisms at successively higher trophic
levels, both the materials and the energy are used again one or more
times. A summation of amounts of production at all trophic levels for
a long period, such as a year, therefore,
similarly has little meaning.
The procuring of significant measurements of
production rate is
further complicated by the fact that growth, consumption, and decom-
position may all be going on simultaneously. This difficulty may be
dealt with by reducing the period considered to such a small size that
instantaneous rates for the various processes are obtained; in other
words, by obtaining the differentials of the curves describing these
processes (Clarke, 1946). The productive activity of an ecosystem is
therefore best measured as a set of rates applying to the gross pro-
duction, net production, and net increase for each category of or-
ganism or each trophic level. The term "productivity" has sometimes
been used specifically for the rate at which assimilation (gross pro-
ductivity) or growth (net productivity) is taking place since it is a
measure of the rate at which the wheels of the system are turning. In
view of the fact that this term is used in various broader senses it is
preferable to employ the more explicit term, "production rate," and

to specify the units of time and area considered.
Efficiencies.In ecosystems in which the magnitudes of the rates of
production at the different trophic levels can be measured, calcula-

tions may be made of the quantitative relationships of the various
steps in the production. The ratios, or efficiencies, on page 498

are of interest. In addition ratios including several or all of the steps
within a trophic level, or embracing more than one trophic level, may
be calculated. For example, the efficiency of the growth of a plant
or animal component may be obtained as a percentage of the incident
radiation.
Where the rates of the processes involved in the ecosystem have
been measured, the various ratios, or efficiencies, can be calculated.
Very few accurate measurements are available for organisms under
natural conditions, but the order of magnitude is suggested by the
following generalities. In ponds with dense algal blooms as much as

14 per cent of the light incident on the water surface may reach and
be absorbed by the plant cells; in the open ocean less that 1 per cent
of light energy is absorbed by the phytoplankton. Complete forest
cent of the sunlight, but desert vegeta-
canopies absorb up to 99 per
tion obviously captures a far lower percentage. The efficiency of the
that is, of plant assimilation, for Chlorella
photosynthetic process,
cultureswas found to be 12-20 per cent in moderate light and 2-3 per
cent in the full light of summer, and higher plants are reported to
display efficiencies of the same order (Wassink, Kok, and van

Oorschot, 1953 ) . When the small amount of light reaching the algae
in a series of lakes was combined with their average photosynthetic
efficiency,
values for the rate of utilization of light incident on the
water surface in the manufacture of carbohydrate ranged from 0.043
to 0.38 per cent (Clarke, 1939).
For the animals of the ecosystem, sample values may be given for
herbivore and for carnivore production. It is said that 40 kg of grass
are required to make 1 kg of beef. This is a value involving 2 steps
in our analysis the ratios of herbivore assimilation and of growth and
Efficiencies 499
represents a net production for the herbivore equal to 2% per eent of

the food consumed.
Obviously in the case of a mammal a large part
of the food
energy goes into heat production, and any active animal
burns up considerable fuel in its
locomotory activities, including
foraging for food. Fish culturists have found that about 1 kg of fish
can be produced for
every 5 kg of meat consumed-or an efficiency
of about 20 cent for this Measure-
per highly digestible type of food.
ments of the efficiencies of various
steps in production for pond com-
munities have been made by Lindernan (1942), for oceanic
plankton
by Riley, Stoinmel, and Bumpus (1949), and
for certain farm
crops by
Transeau (1926). A
schematic representation of the chief compo-
nents in the ecosystem of
Georges Bank, a fishing area off the coast of
Massachusetts, shown in Fig. 13.12, summarizes our present knowledge
of the quantitative relations involved. The blanks indicate the gaps
Radiant energy: 3,000,000 g-cal/day
1.6-7.7g-cal/day
Currents Migration
Migration Currents
FIG. 13.12. Principal components and processes involved in production on

Georges Bank, off Massachusetts. Values are averages for the whole bank per
square meter of sea surface. Maximum and minimum values within the cogwheels
are for the standing crop; those over the wheels are for net production rate.
Average yield during 1923 to 1945 is indicated at right. (Clarke, 1946.)
in our information, and the large range in values draws attention to
the variability in the abundance of the organisms, in the yield, and in
the production rates. It will be observed that an average of 3 million
g-cal of radiant energy fall on each square meter of sea surface over
the Bank per day and that the energy content of the fish landed per
day averages about 5 g-cal per sq m, or an overall efficiency of 0.00015

per cent. This very low value is due to the small fraction of the sun's
radiation actually absorbed by the plant producers and the losses at
each step in the long food chains involved (Clarke, 1946).
PRODUCTIVITY OF LAND AND WATER

Let us conclude our consideration of the ecosystem with a general
comparison of the circumstances and efficiencies of its operation on
land and in the water. The terrestrial environment clearly has certain
advantages. More light is usually available, and large plants are
ordinarily much more abundant. For this reason food chains can be
shorter with correspondingly greater efficiency of production. Be-
cause the soil does not have the mobility of the water medium,
nutrients are typically more concentrated, and products of decomposi-
tion tend to be retained as a humus near the surface of the ground
where they can be immediately used again. On the other hand,
the lack of water and the extreme changes in temperature are promi-
nent among the disadvantages of the land as a place to live.
In the littoralzone of the ocean and of lakes, light is

sufficiently
intense for the growth of both planktonic and benthic plants. Since
the water in this zone is continually mixed, nutrients are rapidly re-
distributed and food particles are carried in abundance to sessile ani-
mals. The disadvantages of the littoral zone include excessive molar
action, considerable temperature change, and other special dangers

near the water's edge. Beyond the littoral zone the replenishment of
nutrients to the surface and of oxygen to the depths must await sea-
sonal stirring. In the open sea and in the deepest lakes the euphotic
zone represents only a tiny fraction of the entire depth, and effective
wind stirring never extends below the permanent thermocline. The
result is that nutritive materials sinking into deep water are regen-
erated only very slowly. Although organisms inhabiting large water
bodies enjoy such advantages as relatively stable temperatures, they
are handicapped by a restricted nutrient supply at the surface and by
a lack of light at greater depths.
In relation to the distribution and availability of the critical ma-
terials in the major environments of the world we realize that the run-
Productivity of Land and Water 501
oft from the land, including river and sewage discharge, represents a
transfer of nutrients from the terrestrial environment to inland waters
and thence to the ocean. Much of the nutrient material is utilized
by the marine community, but, as we have seen, a portion of it is being

permanently removed from the system as inert dissolved compounds,
as submarine deposits, and as additions to the reservoir in the ocean
abyss. The land is therefore continuously losing nutrient materials

to fresh water and finally to the sea, and very little return movement
of these materials to the terrestrial environment takes place.
Relatively few channels exist by which the crucial nutritive sub-

stances are being restored to the land. The use of fish, shellfish, and
algae as food by land animals and the harvesting of marine products

by man for fertilizers and for industrial uses, as well as for food,
represent a return movement of materials to the land. A large frac-
tion of the world's photosynthesis takes place in the sea. Estimates
differ as to whether the average annual growth of the phytoplankton
per unit area is as great as that of the land vegetation (Riley, 1944) or
somewhat lower (Steeman Nielsen, 1952). Nevertheless, since the
oceans cover about 70 per cent of the earth and since phytoplankton
exists in all surface waters, perhaps half or more of the primary syn-
thesis of organic matter for our globe is carried out
by marine plant
producers. Yet less than 1 per cent of man's food comes from the
marine environment! From the point of view of conservation what-
ever increase can be made in man's use of marine resources will help
to restore the balance and will represent a gain from the point of view
of the inhabitants of the land (Clarke, 1950).
In view of the widely varying ecological conditions found in the
major environments it is of interest to compare all

aspects of their
productivities in so far as this is possible. Characteristic differences
in the standing crop of various areas have already been discussed.
Far more difficult is a comparison of the production rates in view of
the complexity of the relations between the various trophic levels in
each ecosystem. Very few data exist for comparative purposes in
natural environments. we may say that the efficiency of
In general
plant growth is
usually greater than that of animal growth, and the
efficiency of animal production is

greater in the terrestrial than in the
aquatic environment because of the shorter food chains.
Of particular interest to man is the yield which may be obtained
from different types of environment. As we have seen, the magnitude
of a sustained yield depends upon the supply of energy and of material
and upon the production rates and efficiencies of the various links in
the food chain. Table 22 gives some sample values for man's annual
harvest per unit area. As would be expected from the previous dis-
cussion, the yield is greater for organisms at lower trophic levels than
for those further along in the food chain. The highest yields are
obtained when man can harvest the plant producers directly. The
value given for corn is a maximum crop after a growing period of 100
days (Transeau, 1926). The solar energy reaching the cornfield dur-
ing the rest of the year is lost as far as this crop is concerned. The
figure given for sugar cane in Hawaii, where a growing season of 11
months exists, represents one of the largest annual yields obtainable
from any crop.
TABLE 22
COMPARISON OF ANNUAL YlELD TO MAN FROM VARIOUS ENVIRONMENTS

Values (from various sources) are in pounds per acre (roughly, kilograms per
hectare) for fresh weight of whole organism.
Grass Rich pasture 30,000

Com Ohio 40,000
Sugar cane Hawaii (max.) 200,000
Cattle and sheep Range 150
Rich pasture 700
Fish Georges Bank, Massachusetts 20
Lake Mendota, Wisconsin 19
Fertilized ponds, Alabama 300
Milkfish, Philippines 700
Cultivated ponds, India and China 3,000
Mollusks Clams, Cape Cod (max.) 2,000
Mussels, Europe 4,000
Since cattle, sheep, and other farm animals are products of a two-
link food chain that are raised in a
relatively advantageous environ-
ment, it is not surprising that the yields are considerably higher than
those ordinarily found for animals in uncultivated aquatic environ-
ments. The harvest taken from natural populations of fishes such
as those from Georges Bank and Lake Mendota is thus much below
the yields obtainable from the growth of farm animals, but probably
it does not
compare unfavorably with man's take from wild animal
populations on land.
In the fertilized fish ponds of southeastern United States an average
of 300 kg of fish per hectare per year may be produced. An excellent
effect of the length of the food chain is furnished
example of the by
the sizes of the yields of fishes of differing ecological position in these
ponds. When goldfish, which are herbivores, are raised a yield of

500 kg per hectare is obtained, but the yield for bluegill sunfish, a
primary carnivore, is 250 kg per hectare, and that for bass, a secondary
Conclusion 503
carnivore, is
only kg per hectare per year. The herbivorous
100
milkfish in the Philippines, raised under
very favorable tropical con-
ditions, provides an annual yield of 500 to 1000 kg per hectare without
the addition of fertilizer (Frey, 1947), and even
larger harvests are
reported from ponds in India and China. The very high yields of
shellfish in the littoral zone are
partially due to the fact that these
animals act as food concentrators.
They feed on plankton and detritus
brought tothem by currents from regions far beyond the area in which
they are harvested. The efficiency of their production is thus not
exactly comparable to the other items in Table 22, but the harvesting
of these shellfish is clearly a very efficient use of marine resources
by man.
CONCLUSION
The comparative productivity of land and water is a fitting topic
with which to close our discussion of the elements of ecology. In this
chapter we have shown how the individual physical and biological
influences of the environment interact in the functioning of the eco-
logical complex. We
have considered the availability of solar energy
for green plants in various habitats and the circumstances under which
organic compounds can be synthesized by these primary producers.
We have traced the transfer of materials and energy through succes-
sive links in the food chain and have discussed the qualitative and
quantitative restrictions placed on the various plant and animal com-
ponents of the ecosystem by the specific conditions of each habitat
and each type of dependency. Thus the concept, with which we
began, of the living community and its environment as a reciprocating
system in dynamic balance has been analyzed and illustrated. We
have seen how the intricate relations between the organism and the
environment underlie the striking differences in the abundance and
kind of plants and animals that populate contrasting habitats, in
the harvest of living material that may be removed, and in the rates
of production that are displayed. An understanding of these ecologi-
cal relations enables man to live in adjustment with his natural en-
vironment, to preserve and develop it, and to derive the greatest
benefit from it.
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Index
Abrasive action of medium, 53-58 Austausch coefficient, 248

Absorption of sun's rays by various Autecology, 14
media, 135-136 Autotomy, 58
Abyssobenthic zone, 30 Autotrophic nutrition, 278
Aestivation, 121, 146-148
Age distribution, 344 Bacteria, 298
Aggregation in response to temperature, antibiotics and, 383-384
157 chemotrophic, 278
Aggregations, see Groups commensal, 377
Agriolimax, tactic reactions, 214 effect of pressure, 31
Air, distribution by, 45-48 in soil, 77
Alewife, orientation, 267 luminescent, 204
Algae, antagonistic substances 384
of, symbiotic, 365-366, 368-369
depth in relation to light, 235-237 thermophilous, 130
in hot springs, 130 i see also Decomposers, Decomposi-
on tree trunks, 118 tion, Nitrification, and Trans-
on turtle shells, 376 formers
zonation in tidal zone, 102-105 Bactericidal effect of sunlight, 231
Alkalinity of natural waters, 263 Balanoid zone, 104-105

Allen's rule, 157 Barnacles, setting of larvae, 60
Alpine plants, 160-162 temperature as affecting distribution,
see also Life zones 171
Altitude zonation in, 104
effect on O 2 availability, 243 Barriers, 428-429
effect on distribution of animals, 27 Bass, 7
29 Bass tapeworm, 390-392
effect on temperature, 134 Bathypelagic zone, 30
effect on vegetation type, 175-179 Bats, echo location by, 61
limitation of distribution by, 253 hibernationin, 147
Altitudinal life zones, 175, 179 Beavers, 25, 495
Aluminum in pedalfer soils, 86-87 Bees, control of hive temperature, 322-
Amphibious environments, water prob- 323
lem in, 99-107 orientation in, 215
Anaerobic organisms, 243, 255, 258 Benthic division, 30
Animal exclusion, theory of, 447 Benthos, 36
Antibiosis, 383-385 Bergmann principle, 157
Ants, leaf-cutting, 372 Bicarbonate, 263
slave-making, 385 Biocenose, 402
*
symbiotic relationships in, 370-371 see also Community
territory in, 353-354 Biogeochemistry, 303

Aphids, 325, 370-371, 387 Bioluminescence, 204-207
Aphotic zone, 241 Biomass, 469-470
Aquatic communities, stratification of, Biomej 448-451
423-424 ftiotliermal zones, 180
Association, 448 Biotic potential, 334
523
524 Index
Biptope, 404 Carrying capacity, 485
Siotypes, 325-326 Caterpillars, tent, 211-213
Birds, changes with succession, 435 Cave communities, 198-199, 207
colonial, 5, 324-325 Chemotrophic organisms, 278
homing in, 355 Chernozems, 86
migration, 228-229, 355-357 Chilling, effects of, 168-171
reproduction in, 324-325 Chironornid larvae, 253
territory in,350-352 Chloroplasts, effect of light on, 198
Black Sea, circulation, 251-252 Chromatic adaptation, 235-237
oxygen lack, 258 Chrysanthemum, 226
Blood, effect of carbon dioxide on, CAcada, fluctuations in, 458
266-267 Ciliates, prey-predator cycles, 398-399
Bluegill sunfish, competition in, 321- Cladocerans, competition in, 397
322 effect ofcrowding, 325
Bogs, 433-434 effect of temperature, 156
Boll weevil, 296
Climatographs, 109
effect of humidity and temperature, Climax, 431-432, 447
181-182 see also Succession
Bottom fauna, 29, 6&-69, 253 Closed communities, 429
Breeding sites as population limit, 486 Co-action, 18
Bromeliads, vertical zonation, 235, 375 Cod, distribution in relation to food,
Buffer capacity of natural waters, 263- 294
264 effect of temperature on egg devel-
Burrowing animals, 72, 76-77, 149-150 opment, 154
Burrows as microclimates, 118-119, Colloidal complex of soil, 79-80
149 Colony, 310, 331, 429
Buttresses of ceiba trees, 54 Color, of animals, 199-202
Color changes in animals, 200
Cactus plants, 122 Combined water, 111
Caddis-flies,57-58 Commensalism, with continuous con-
Calciphobes, 285 tact, 374-377
Calciphytes, 285 without continuous contact, 377-381
Calcium, in pedalfer soils, 86 Community, classification of, 405
in pedocal soils, 85-86 closed, 429
in soil, 284-285
composition of, 414-424
Calcium carbonate, in fresh water, fluctuations within, 451-464
271-272 interrelations, 419
in oceans, 271-276 limits of, 405-406
in soils, 270-271
species/genus ratio in, 417418
Capillary water, 111 stratification of, 420-424
Carbon cycle, 301 zonation in, 406-407
Carbon dioxide, effect on aquatic ani- see also Succession
mals, 266-268 Community concept, 401
effect on aquatic plants, 265 Community dominance, 408-410
effect on land plants, 261
Community type, 447-448
in aquatic environment, 262-268
Compensation depth, 238, 475
in atmosphere, 260-262
Compensation intensity, 238
reactions with water, 262-265
Competition, 317-322, 396-400, 409-
Carbonate, 262-263 410
Carnivores, 393, 467-468, 479-181 Composition of natural waters, 92
Carnivorous plants, 279, 393 Conditioning of medium, 319
Index 525
Conservative factors, 21 Deer, competition for food, 320-321,
Consumers, 467-468 399
in the sea, 476-481 Denitrification, 299
Convergence in succession, 438-439 Density of media, 26
Copepods, 476-479 Density-dependent factors, 335
fluctuations in, 456-457
Density-independent factors, 335
orientation to light, 214-215
Deposits, 482-483
survival of, 8 Desert, temperatures, 134, 149
temperature tolerance, 163-164 water conditions, 114, 119-120
zonation in tidal species, 106-107 Desiccation, for
adaptations resisting,
Corals, 58, 69, 274-275, 368 100-103, 120
Corn, production of, 502 Detritus, 479
Corrosion in soil formation, 77
Diapause, 146-147
Cowbird, 296, 385 need for chilling in, 170
Crabs, commensal, 379 Diatom ooze, 71
zonation in, 103 Diatoms, abundance cycles of, 474-476
Cricket, chirping rate and temperature, nutrient requirements, 288-289
153
photosynthesis of, 239
Crowding, beneficial effects, 322-328 see also Phytoplankton
effect on sex ratio, 325 427-430
Dispersal, 42-53,
effect on tree growth, 317
Dispersion, 333
harmful effects, 317-322
D'sphotic zone, 241
Cuckoo, 385 Distribution, in relation to nutrients,
Cultch, 60 294-296
Current demand, 45 in relation to oxygen supply, 253-
Current indicators, 50 259
Currents, in basins, 252 in relation to temperature, 159-175
in lakes, 247 in relation to water factor, 121-128
in ocean, 42-44, 250 Diurnal fluctuation, in oxygen, 245
in streams, 42, 45, 48, 52, 57 in relative humidity, 115
Cycle, carbon, 300 in temperature, 115, 131, 246
nitrogen, 299 Diurnal migrations, 217-219

sunspot, 463-464 Diurnal rhythms of activity, 217-218
Cyclic fluctuations, 457-458 3132
Diving mammals,
Cyclic populations, 342 327-332
Division of labor,
Cyclomorphosis, 41-42
Dominance, community, 408
Cypress trees, 254
social, 328
Cyprids, 60
Dominant in communities, 410
Dace, forage ratio, 395 Drosophila, 319, 320, 323

Drought, adaptations to resist, 100
Daphnia, competition in, 397
effect on geographical distribution,
cycloinorphosis in, 41
123
parthenogenesis in, 100
343-344 Ducks, emigration of, 358
population fluctuations,
temperature relations, 155-156 mortality of, 6
Daylength, 190 Dy.strophic lakes, 287
Decomposers, 467-468
Earthworms, action on 78-79
Decomposition, 298-304, 467 of, soil,
in marine ecosystem, 481 Ecesis,429-430

in ponds, 9, 291 Ecological complex, 16-18, 465
in soils, 77-79 Ecological succession, 426
526 Index
Ecology, development of, 13-18 Fish, production, 347, 492, 499, 502
human, 3, 19 utilization of, 346
Ecosystem, 16, 465 Fisheries, management of, 344-348,

productivity of, 481 453-456, 491-492
Ecotone, 410-414, 419 Fjords, 311
Ecotypes, 325-326 circulation in, 251
Eddy conduction, 247-248 Flounder, color changes in, 200
Eel, migration of, 50-52 thermal migration in, 150
Eel grass, 70-71 Flowering, effect of photoperiod on,
Effective temperature range, 143 224-229
Efficiency of production, 497-500 Flowers, pollination of, 372
El Nino current, 141, 458 Fluctuations, in communities, 451-464
Elasmobranchs, osmotic pressure of, 95 in populations, 342-344
Elk, 329-331 Fly, house, temperature relations of,
Emigration, 357-361 144

English sparrow, photoperiodism in, Food, competition 321-322
for,
229 Food 494-496
chain, 419, 468,
nature 5-13 Food preferences, 293-294, 395
Environment, critical of,
meaning of, 3-5 Forage ratio, 395

Environmental resistance, 335 Forest, ecotone in relation to, 411-412,
Ephippium, 49 419
Epilimnion, 138 effect on radiation, 187-188
Epiphytes, 44, 66, 374-377 effect on water
factor, 115-116
Epizoans, 66, 374-377 humidity in, 114, 460
Estuaries, 48-49, 94, 98 penetration of light, 234-235
Euphotic zone, 241 soil
profile in, 83, 88
Euryhaline organisms, 97-98 species composition, 123, 417
Eutrophic lakes, 287 stratification of, 421-423
Eutrophication, 496 types in eastern North America, 174

Everglades, 406 see also Biome, Life zones, and Suc-
Excretory systems, water retention in, cession
120 Forest floor, 422, 437, 472
Exploitation, 385-396 Formation, plant, 447
Extrinsic cycles, 461
Fouling, prevention of, 64-65
succession in, 443
Faciation, 448
Foxes, population fluctuation, 462-463
Feeding, adaptations for, in animals,
ear development of, 158
293-294
286 Fragmentation of rock, 77
Fertilizers,
use in ponds, 289-291 Freezing, adaptations for resisting, 144-
145
Fiddler crabs, 403
Field capacity, 111 Frogs, temperature requirements of, 168
Fire, effect on seed cones, 145 Frost, killing, 173
effect on succession, 432, 444-445 see also Chilling and Solifluction
Firs, Douglas, 462 |

Frost line, 173
Fish, commensal, 379 Frostless season, 165
farming, 290-291, 502 Fungi, dry rot, 63

mortality, 5, 9, 344, 459 luminescent, 204
pilot, 380 predaceous, 393
ponds, 257-258, 502-503 symbiotic relations of, 364, 366-367,
population, fluctuations in, 453-456 371-372, 381
Index 527
Gambusia, 395 Hermit crab, symbiotic relations of,
Gannets, 5 369-370
Geotaxis, 211-212 Herring, fluctuations in, 454-456
Geotropism, 207 food relations of, 415-416

Gills in tidal-zone crabs, 103 Hibernation, 145, 147
Globigerina, 39 Holophytic organisms, 278
Globigerina ooze, 71 Holozoic organisms, 278
Gloger's rule, 126, 157 Home range, 350-357
Grass, beach, 403, 407 Homing, 215, 353-355
production, 116, 502 Homoiotherms, 130
Grasshopper, Australian, development Homoiothermy, 148
of, 170 Horizons of soil, 80-84
Grassland community, ecotone in rela- Humidity, absolute, 112
tion to, 411 relative, 113-117
fluctuations in conditions, 115 Humus, 79-80, 301
soil profile, 87 see also Soil
stratification of, 420-421 Hurricanes, transport of organisms by,
see also Biome, Life zones, and Suc- 47
cession Hydrarch succession, 432-435
Gravitational water, 111 Hydrophytes, 99
Gravity, orientation to, 208-209 Hygroscopic water, 111
Grazing, on land, 444, 491 Hyperparasites, 389
of zooplankton, theory of, 478 Hypolimnion, 138
Group organization, 327-332
evolution of, 331-332 Indeterminate plants, 224
Indifferent plants, 224
Groups, originating through locomotion,
312 Inflection point in population curve,
originating through passive transport,

338
311 Insectivorous plants, 279, 393
originating through reproduction, Interstitial fauna, 23

309-310 Intolerant plants, 233
Growth rings, 317 Intrinsic cycles, 460-461
Grunion, reproduction of, 221-222 Invasion, 427-430, 433

Ions, abundance, in natural waters, 92-
Guano, 141, 303
95
Gypsum, influence of flora, 285
Iris, ecotypes in, 326
Gypsy moth, 47 Iron in pedalfer 86-87
soils,
development of, 170-171
Irruptive fluctuation,453-457
Habitat ecology, 14 Irruptive populations, 342

Ivy, distribution in relation to temper-
Haddock, yield of, 492-493 162
ature,
Halophytes, 96-97
Hammock communities, 405-406, 487
Jordan's rule, 158
Heat, 129
control and distribution by, 164-168
Kelp community, 415
transfer of, in water, 136-138 208
Kinesis,
Heat sum, 165-166 Knees of cypress trees, 254
Heath hen, 327
Herbivores, 393, 467-468 Lakes, calcium carbonate in, 271-272,
in ocean, 47&-479 circulation in, 138-139, 247
on land, 471, 491 light penetration in, 191-198
528 Index
Lakes, nutrients in, 287 Logistic curve of population growth,
O supply
2 244-249
in, 339-340
pH and carbon dioxide in, 264, 266 Logs, communities in, 417
productivity of, 502 succession on, 440
stratification, 138 Long-day plants, 224
temperature in, 133, 139 Longspurs, Lapland, 9
Laminarian zone, 104-105 Luminescent organs, 204-207
Law of the minimum, 281-283 Lunar periodicity, 219-224
Leaching of soils, 82-89 Lynx population fluctuations, 399, 457-
Leadership, 328-329 458, 462-464
Leafless plants, 120
Leaves, orientation of, 209 Mackerel, 315
Lemmings, 357-359, 457 development of eggs, 153
Lepisma (silverfish), humidity as af- 453-454
fluctuations in,
fecting, 126 mortality of, 5

Lepomis, vision in, 203 Malacosoma, 211-213
Lichens, 78, 364-365, 375, 420 Mangrove, 377, 406
Life form, of plants, effect of environ- succession, 440-442
ment, 182-183 Marl, 271
Life table, 344 Mass sinking, 248
Life zones, 175-181 Maximum effective temperature, 143
distribution in water,
Maximum survival temperature, 143
Light, angular
196-197 Medium, abrasive action of, 53-58
27-32
biological effects of, 198-241 density,
transport by, 42-53
changes in spectral composition in
water, 191-193 viscosity of, 32-42

distribution of, 186-198 Mesarch succession, 432
duration and amount, on land, 188- Mesophytes, 119
191 Metabolic water, 119-120
influence on activity,202-204 Mice, 201, 217, 335, 353, 395, 457
187-189 Microclimates, 117-119
intensity in forests,
Microcosm, lake as, 446
intensity on land, 187-188
orientation to, 208-216 Microorganisms in soil, 77, 298
see also Bacteria and Nannoplankton
penetration in water, 191-192, 238-
241 Midge larvae, 253
periodicitiesdetermined by, 216-230 Migration, 215, 355-357

diurnal, 217-219
spectral composition on land, 186-
187 in birds,150-152, 228-229
of eel,50-52
ultraviolet, 230-233
thermal, 148-152
Limiting factor, 20
446 Mimicry, 200
Limpet glades,
Littoral zone, 101-107
Minimum, law of, 281
Minimum effective temperature, 143
Littorina zone, 104-105
Minimum survival temperature, 143
Lobster, temperature relations of, 166-
Mistletoes, 386
168
Mixotrophic organisms, 279
Lociation, 449 Modifiable factors, 21
Locomotion, speed of, in air and water, Moina, control of sex ratio by temper-
34-35 ature, 159
Locust, migratory, 359-360 Moisture, action with temperature, 181-
Locust trees, 318 184
Index 529
Moisture, in soil, 111 Ocean, circulation in, see Currents
influence on growth and distribution, currents, 42-44, 250
121-128 ecological cycle of, 473-481
Mollusks, nutrition in, 293 light penetration in, 191-198, 230-
production of, 502 233
Monophagous animals, 294 nutrient cycle in, 301-308
Mortality, 5-10, 333-334, 459 nutrients in, 287-289, 305-307
Mosquito larvae, feeding of, 62-63 O supply in, 242-251
2
Mud-bottom community, 409, 443 pH and carbon dioxide in, 263-265

Muskeg, 183, 431 photosynthesis in, 235-241
Muskox, 329 salinity, 93
Mussels, 64, 390, 443 utilization of resources by man, 500-
Mutualism, with continuous contact, 503
364-370 vertical temperature structure of,
without continuous contact, 370-374 136, 139-141
Mycorrhizae, 366-367 zones of, 30
Old-field succession, 436-438
Nannoplankton, 478 Oligotrophic lakes, 287
see also Plankton Optimal yield, 344-348
Natality, 333-334 Organic matter, decomposition of, 300
Nekton, 36 dissolved, 478
Neritic province, 30 Orientation, 207-216
Neuston, 62 in anadromous fish, 267-268
Niche, 46S-469 in bats, 61
Nitrates, in lakes, 288 in bees, 215
in plant growth, 280-283 in birds, 355
in oceans, 288-289, 306-307 Osmotic pressure, of elasmobranchs, 95
in soil, 284 of natural waters, 93-95
Nitrification, 298-299 of plants, 96-97
Nitrogen, availability of, 80, 280 of teleost fishes, 95

release of, 299-301 Overturn of lakes, 13
requirement of plants, 280 Owls, 335, 377, 395
Nitrogen cycle, 299 Oxidation-reduction potential, 251-253
Nitrogen fixation, 299 Oxygen, adaptations of aquatic animals,
Nomadism, 357 255-256
Non-conservative factors, 21 availability in air, 243, 253
Nutrients, availability for plants, 284 availability in aquatic environment,
in fresh water, 287 244-251
in oceans, 287-289, 475-476 availability in soil, 243, 253
in soil, 80, 284-286 consumption, 256-257
influence on animals, 292-297 effect on distribution, 253-259
limitation by, 283-292 partial pressure in aquatic environ-
regeneration of, 304-308 ment, 255
requirements of animals, 292 pulse, 245
requirements of plants, 280-292 requirements of animals, 253, 257
stagnation of, 303-304 requirements of plants, 253
trace elements, 280, 285, 297 saturation values, 244
Nutrition, modes of, 278-280 supersaturation, in sea and lakes,
244-245
Ocean, bottom, 66-69 supply to deep waters, 246-251
calcium carbonate in, 272-276 Oyster, population sizes in, 326
530 Index
Oyster, salinity tolerance of, 98 Photosynthesis, in aquatic plants, 235-

setting of, 60, 443 240
temperature relations of, 164, 166- in land plants, 233-235
168 relation to carbon dioxide, 260-262,
Oyster crab, 37&-379 26,5-266

Phototaxis, 207-211
Palmettos, 445 Phototropism, 207-210
Palms, 173, 406 Phytoplankton, 36-37
Palolo worms, 220 effect on light extinction, 194
Paramecium, 398 in ponds, 291
296
Parasite-host relationships, in sea, 474-476
Parasites, 279, 467-468 photosynthesis in, 237-241
ectoparasites, 388 Pigmentation, influence of light on,
endoparasites, 388 198-202
modifications of, 390-392 Pine, white, 349, 437-438
Parasitic birds, 385-386 Pineland community, 445
Parasitic insects, 387-389 Pioneer assemblage, 429
Parasitism, 386-392 Pit vipers, temperature orientation in,
Parthenogenesis, 100 157

Pea seedling, effect of temperature, Pitcher plant, 393
154 Plankton, 36-42
Peck order, 328 traps, 311
Pedalfer soils, 86-88 see also Nannoplankton, Phytoplank-
Pedocal soils, 85-86 ton, Zooplankton
Pelagic division, 30 Plant lice, see Aphids
Penguins, 354 Podzol, 86-87
Periodicity, diel, 216 Poikilotherms, 130
diurnal, 216-219 Polarization of light, 215
lunar, 219-224 Pollination,by insects, 372-374
seasonal, 224-230 by wind, 46
thermal, 131, 166 Ponds, production in, 289-291, 493-
tidal, 101 496, 502
Permafrost, 174-175 succession in, 432-435
pH, effecton distribution, 269-270 Poplar community, 419
effect on lime formation, 272-273 Population cycles, 457-464
in aquatic environment, 263-265 334-336
Population density, 317-328,
in soils and muds, 269-270
Population ecology, 15
tolerances of organisms, 268-269 Population growth, 336-340
Phosphate, in lakes, 288 in relation to standing crop, 483-485
in ocean waters, 288-289, 302, 306- in United States, 340
307 Populations, development of, 333-348
in plant growth, 280-283 340-344
equilibrium in,
in soils, 284 fluctuations in, 451464
Phosphorescence, see Bioluminescence minimum size, 326-327
Phosphorus, requirement of plants, 280 mixed, 333
Photic zone, see Euphotic zone optimal yield, 344-348
Photokinesis, 202 single-species, 333
Photoperiodism, in animals, 227-229 space requirements, 348-350
in plants, 224-227 Pore space in soils, 111
Photophores, 206 Potassium, in soil, 284
240 requirements of plants, 282
Index 531
Prairie community, 419 Regeneration, of nutrients, 304-308
Prairie dogs, 310 rate, of 305
Prairie soils,85 ratio of materials, 306-307
Prairies, 124 Rernora, 378
see abo Life zones Reproduction, in colonial birds, 324-
Precipitation-evaporation ratio, 125- 325
126, 183-184 effect of photoperiod on, 227-229
Predation, 392-396 effect of population density on, 320-
Predator-prey fluctuations, 397-399 321, 323-328
Pressure, 27-32 production of aggregations by, 309-
in air, 28-29 310
in water, 29-30 Respiratory systems, adaptations of, for
Primary succession, 432 water conservation, 120
Principle, Bergmann, 157 Return migration, 355-357
of common transport, 248 Rheotaxis, 57
of edges, 412-413 River communities, 417
of minimum population, 327 Root secretions, 78
of partial equivalence, 75 Roots, 76
Producers, 467-468 in relation to
oxygen, 253-254
in the sea, 474-476 in relation to water, 121-122
Production pyramid, 469
Production rate, 482-483, 493-500 Sacculina, 390
Productivity, concepts of, 482-500 Sagitta as indicator species, 50
of land and water, 500-503 Saline soil, 97
relation to material removed, 489- 97-99
Salinity, limiting effects of,
493 92-95
of natural waters,
Protective coloration, 60, 199-202 Salmon, spawning of, 313-314
Pteropod ooze, 71 Salt, content of natural waters, 94
Pyramid of biomasses, 470 lakes, 93-94
Pyramid of numbers, 471-472, 482 Salt-marsh community, 403

Pyramiding in tree growth, 234 Sand, organisms in, 24, 417
Sand dunes, succession in, 440-441
g >0 , 152 Saprophytes, 278, 467-468
Quail, 413-414 Sargassum, 50
Saturation deficit, 115-116
Radiation, solar, 186-191 Sawgrass, 406, 445
Radiolarian ooze, 71 Scattering layer in oceans, 218
Ragweed, photoperiodism in, 226 Scavengers, 467-468
109-111
Rainfall, Sea anemone, symbiotic relationships,
in relation to temperature, 113-115 369-370
Rain-forest,234-235 Sea 329-330
lions,
Ranatra, 210 Seals, fur, 310, 329
Range animals, 472, 502 Secondary succession, 435-438
Recognition in group organization, 313- Seeds, distribution by animals, 373
316 distribution by water, 48, 442
Recruitment, 346 distribution by wind, 46
Red 384
tide, resistance to desiccation, 121
Redfish, 295 resistance to temperature extremes,
Redox potential, 251-253 144-145
Reef building, 69, 274-275 Sequoia, 4, 7, 33
see also Corals Serai index, 438
532 Index
Sere, 431 Species flocks, 417
Serpentine, toleration by plants, 285 Species/genus ratio, 417-418
Sessile existence, 44-45, 310 Spectral distribution of radiant energy,
Sex ratio, influence of crowding on, 134-135, 192-194
325 Springs, 132
influence of temperature on, 158 thermal, 130
Sheep, 315, 384, 491, 502 Spruce forest, 431
Shells, formation of, 273-276 community, 408
Short-day plants, 224 Stagnation, in cycles, 303
Sinking rate ef marine organisms, 36- in lakes, 138
42 Standing crop, 482-489, 494-496
Size, of organisms, in relation to equilibrium level of, 485-487
medium, 32-34 in relation to population growth,
in food chain, 470 483-485
Slave-making ants, 385 in, 487-489
regional differences
Snails, 104, 121, 312 428
Starling, dispersal of,
Social dominance, 328 Stenohaline organisms, 97-98
Societies of animals, 331-332
Stereotropism, 61
Sodium, requirements of plants for, Strangling fig, 383
282 Stratification, in communities, 420424
Soil, action on plants, 73-76 in lakes,138
calcium carbonate in, 271 in ocean, 136
chemical composition, 72-73 Streams, 42, 48, 52
community, 420-421 adaptations to current, 57
decomposition in, 301 Substratum, aquatic, 66-71
effects of organisms on, 76 attainment of, 60-61
erosion, 11 breakdown 69of,
formation, 77-79 build-up 69-71
of,
moisture 111-112
in, reactions to, 61-62
nutrients in, 284-286 terrestrial, 71-89
organisms in, 76-77, 472 types of, 62-66

oxygen in, 244 Succession, 425-446
particles, 74 arrested, 431
pH, 269-270 convergence in, 438-439
profile, 80-84 in old fields, 435-438
regeneration of nutrients in, 304-305 in ponds, 432-435

saline, 97 in tidal zone, 442-444
temperature, 130-131 modification of, 444-446

texture and structure of, 73-74 on fallen logs, 440
water, 112 primary, 432-435
Soil-group divisions, 84-89 secondary, 435-438
Solar constant, 186 Sugar cane, 227, 502
Solifluction, effect on tree line, 174 Sulphur, 280
Sound, 215 Sundew, 393
Soybean plant, 366 Sunfish, 203, 321
Spanish moss, 37.5376 Sunspot cycle, 463-464

Spawning of marine organisms in lunar Support by medium, 32-42
cycle, 220-221 Supralittoral /one, 30
Surface film of water, as a substratum,
Species composition, latitudinal differ-
ences, 487-489 62-63
of communities, 414424 Survivorship curves, 344
variation in microclimates, 123 Swallows, tree, 228
Index 533
Swamps, 99-100, 433-434 Timothy, 225

Swim bladder of fish, 31 Tolerant plants, 233-234
Symbiosis, 363-381 Torrents, see Streams
Synecology, 15 Trace elements, in animal nutrition,
297
Taiga, 183, 450-451 in ocean, 287-288
Tarantula, 389 in plant nutrition, 280-281
Taxis, 207 in soils, 285-286
Teleosts, osmotic pressure of, 95 Transformers, 467-468, 481

Temperature, biological action of, 141- Transparency of natural waters, 195-
159 198
changes, horizontal, 134 Transpiration, 114-116
in time, 131-134 ratio, 122
vertical, 134-141 Transport, by medium, 42-53
by 150-175
control of distribution of nutrients, in soil, 82-84
control of egg type and sex ratio, in water, 301-303, 305-308
158-159 production of aggregations by, 311
distribution of, 129-141 Tree line, 173-176
effect on activity, 156 Tree trunks, 118, 440
effect on biological rates, 152-155 Trees, competition in, 317-318
effect on calcium carbonate deposi- fluctuations in growth, 458
tion, 272-274 see also Forests

effect on evaporation, 116 Tribolium, 319, 323
extremes, adaptations for meeting, competition in, 396-397
144-152 humidity and reproductive cycle,
limitation by, 160-164 126-127
in relation to moisture, 181-184 Triclacna, 274, 368
morphological effects of, 157-158 Trophic level, 469-473
need for chilling, 168-171 Tropisms, 207
optimum, 155-156 Trout, relation to pll, 268
results of limitation by, 172-181 relation to temperature, 155
resistance at various life stages, 163- Tundra, 178, 183, 450-451
164 Turnover, 496-497
seasonal vertical changes in water, Turtles, temperature requirements, 168
138-141
tolerance of extremes, 129-131, 142- Ultraviolet radiation, 230-233
189,
143
Unmodifiable factors, 21
variation with altitude, 134-135
Upwelling, 305
variation with depth in water, 136
Utricularia, 393
Tent caterpillar, geotaxis in, 211-213
Teredo, 64
Vegetation of North America, 123-126
Termites, 63, 120, 332
350-357 Vegetation types of life zones, 178-180,
Territory,
183
Thermocline, 136, 138
Vegetative reproduction, 310
Thermoperiodism, 166
Thermotaxis, 156-157 Vernalization, 169
Vertical migration, of insects, 218-219
Thigmotaxis, 61
Tidal zonation, 103-107 of plankton, 217-218
Tidal zone, 101-107 Viscosity of medium, 34-42

succession in, 442-444, 446 Vision, 202-204
Tides, 101 maximum depth in water, 203-204
534 Index
Vital index, 334 Willow community, 419
Vitamin D, 231-233 Wilting coefficient, 111
Wind, effect, in aerating water, 246
Warm-blooded animals, 148 on vegetation, 54-55
Wasps, parasitic, 388-389 on water loss, 116
Water, abrasive action of, 55-58 Winter egg, 49
balance, 107 Wolves, 310, 462
in air, 112-117 Wood as substratum, 64-65
in soil, 111-112 Wood duck, 486
loss, 114->119
natural, composition of, 91-95
occurrence on land, 109-117 Xerarch succession, 432
passage through organism, 35-36 Xerophytes, 119

physical-chemical properties of, 25-
27
Year-class analysis, 454-456
plants, see
Hydrophytes
Yield, 482-483, 490-493, 502
problem, in
amphibious situations,
99-107 per unit effort, 346, 491-492
Yucca, pollination of, 373
in aquatic environment, 91-99
in terrestrial environment, 107-129
transmission of light by ,
191- Zonation, of communities, 406-407
198 of continental vegetation, 123-126
transparency, changes in, 197-198 of marine environment, 30
transport by, 48 of plant communities with altitude,
see also Moisture 175-179
Weakfish, range extension of, 12 related to temperature, 172-181
Whales, 474-480 tidal, 103-107
diving adaptations 31-32
in, vertical, in algae, 235-237
Wheat, in relation to rainfall, 110 in bromeliads, 235
photosynthesis of, 260 Zones, life, 175-181
rust, transportation of, 46 Zoochlorellae, 368
Willow, relation of roots to oxygen, Zooplankton, 38, 476-479
254 Zooxanthellae, 368
CONVERSION SCALES
Oin.
- 1
-3
-4
-5
L-7
COMPARISON OF METRIC AND ENGLISH UNITS
Measures of Length
1 millimeter = 0.039 inch

1 centimeter = 0.394 inch
1 meter - 39.37 inches
1000 meters = 1 kilometer
1 kilometer = 0.621 English mile
1 nautical mile = 6080 feet
1 fathom = 6 feet = 1.83 m.
Measures of Area
1
sq.
cm. = 0.155 sq.
in.
1
sq. meter =
1.196 sq. yd.
10,000 sq. m. = 1 hectare
1 hectare = 2.47 acres
1
sq. kilometer
= 0.3861 sq.
mile
Measures of Volume
1 liter = 1.057 U. S.
quart
1 cu. m. = 1.308 cu. vd.
Measures of Weight
1
kilogram - 2.20 Ibs.
1000 kg.
= 1 metric ton
wr, f oon.ec iKe

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DO

JOHN WILEY & SONS, INC.

JOHN WILEY & SONS, INC.

All Rights Reserved

This book or any part thereof must not

THIRD PRINTING. APRIL. 1959

Contribution No. 704 from the Woods Hole Oceanographic Institution

Library of Congress Catalog Card Number: 54-12205

PRINTED IN THE UNITED STATES OF AMERICA

great, and no attempt at any exhaustive treatment of the

ecology given by the author at Harvard University. Many ideas and

Drury, E. H. Graham, D. R. Griffin, H. C. Hanson, A. G. Huntsman,

Chapter 1 VIEWPOINT OF MODERN ECOLOGY 1

The Meaning of Ecology 2

Chapter 2 THE MEDIUM 23

Contrasting Qualities of Air and Water 24

Significance of the Substratum 59

Build-Up of the Substratum 69

Chapter 5 TEMPERATURE 129

Control by Need for Minimum Amount of Heat 164

Chapter 6 LIGHT 185

Chapter 7 OXYGEN AND CARBON DIOXIDE 242

Hydrogen Ion Concentration 268

Chapter 8 NUTRIENTS 277

Origin of Groups 309

Chapter 10 RELATIONS BETWEEN SPECIES 362

Chapter 11 THE COMMUNITY 401

Chapter 12 SUCCESSION AND FLUCTUATION 125

Ecological Succession 125

Chapter 13 DYNAMICS OF THE ECOSYSTEM 465

Productivity of the Ecosystem 481

Contact with the environment is inescapable. Protoplasm, the essen-

supply of materials; (3) a removal of waste products. The exchanges

of water from their tissues. Animals in the environment of these

eventually destroy the population.

oxygen, as at the top of Mt. Everest, or if nutrients are not avail-

able, as on rocky plateaux, animals or plants cannot obtain their

light may be found, but the temperature may be so high as to exclude

because of high concentrations of carbon dioxide issuing from subter-

THE MEANING OF ECOLOGY

cludes the influences of other plants and animals present as well as

both of the organisms themselves and of the environments inhabited

sociology might be thought of as the "ecology of man" in a broad sense.

stated, "The discoveries of Darwin, himself a magnificent field

THE MEANING OF ENVIRONMENT

a sea-bird rookery (Fig. 1.2). In other situations the effects of the

biological influences whether conspicuousorobsc^iTe^

The Critical Environment

effectively. Others would, of course, succumb before they grew to

The U. S. Fish and Wildlife Service estimated that early in 1944

production of huge numbers of young. The prodigious fertility of

creasing. Many examples could be given of the huge excess number

(Fig. 1.3). Insects are notoriously prolific. The queen termite in an

progeny survived, after only four generations we would have a pile of

of pollen that a yellowish dustis often

ported to produce as many as 1,756,000 seeds per capsule. Spores are

an/ Eschmeyer, 1938

sequently be relatively low, but even slow breeders tend to increase.

by the California "big trees" (Sequoia gigantea] which do not pro-