Population Regulation

Regulation of populations results from extrinsic factors such as food supply,

weather, shelter, diseases,4and natural enemies, or intrinsic factors like
behavior, physiology, and genetic characteristics determining population
density. Some factors, such as behavior, are primarily density dependent and
tend to limit population growth more as its density rises. Other factors, such
as weather, are called density independent because they often affect the same
proportion of the population no matter how dense it is. Most populations
are limited by a combination of both types of factors, though density
dependence seems of greatest importance for vertebrates, and density
independence of greatest importance for insects and plants.
Regulation by competition sometimes produces a more constant equi-
librium than the oscillations of predation. Parasitism and disease often
function as population regulators very much like predators. A nonequi-
librium coexistence, in which disturbance or unpredictable (stochastic) events permit
continuation of the population, but with irregular fluctuations, is increasingly
considered common. There is virtually always a suitable site available for a species
because the disturbances are dispersed in time and space. Weather controls
population size, but the population normally has little feedback effect on the
weather.
As the population size builds, a number of density dependent results considered
to be intrinsic to the population may follow. Dispersal of individuals from an area
takes place. (Conversely, at low density, dispersal into the area may occur.) Some
physiological responses involving hormones somehow triggered by crowding have
also been identified in a number of vertebrates. In certain mammals and birds, the rate
of egg production (or ovulation) in the female may drop, or the rate of natural
abortion of embryos may rise, in either case limiting population growth. In the case
of certain salamanders, crowding has induced cannibalism, with a similar limiting
effect on population growth.
The on-site behavioral controls qf vertebrates are perhaps the most familiar
intrinsic regulatory mechanisms. Social dominance hierarchies and territoriality both
have the effect of limiting reproduction and the number of individuals in an area. In a
social dominance hierarchy, as occurs in the case of chickens or baboons, a pecking
order is established beginning with the most dominant individual and continuing
down to the least dominant individual. In the dominance hierarchy only certain of the
individuals are permitted enough food or resources to survive, and only certain of
the individuals reproduce. Territorial behavior, the establishment (and usually defense)
of a certain small area against other individuals in the population, results in a limited
number of possible individuals being present in a given area.
An example of how territoriality works to regulate population size is seen in
grouse (Figure 2.11). In late summer the old cocks defend their territories, but at the
end of the warm season the population is above the carrying capacity. The family
groups break up. One group of birds is displaced from the territories, and by January
almost all of them are dead. Of the birds remaining in the territories, some are unable
to defend their territories and in the autumn foim a second group of resident surplus

1 OVERVIEW
birds that gradually move to marginal habitats, do not breed, and almost all die by
spring. The third group successfully defends its territories, survives the winter well,
and is the breeding stock for the following spring and summer. In this manner
territorial behavior may regulate an equilibrium population level of grouse.
Population regulation is a fascinating, controversial, and highly significant
ecological research area, made all the more relevant by its relative applicability to the
exponential human population growth on earth.

ECOLOGICAL CONCEPTS IN BRIEF 2

 Family groups (territorial
Late summer behavior by old cocks at
dawn)

Conflict (family breakup) ■

(Displaced birds)|

Nonterritorial transient (Successful territorial

birds, Group 1 Territorial behavior defenders)
Autumn (young and old)
Group 1 birds do not Resident surplus birds, Territorial owners,
breed. Group 2 Group3.
(Some (Some High survival
replacement) replacement)
Winter Most Group 1 birds dead. Group 2 birds move
to marginal habitat

Spring
I
Group 2 birds do not Breeding birds
breed, survival poor
Figure 2.11 Territoriality and social behavior regulating population size in grouse. (From Smith,
1979; courtesy of Harper and Row, Inc., Publishers.)

EVOLUTIONARY ECOLOGY
i
Variation and Selection
So far, we have treated all the individuals of a population as if they were
genetically identical, which is rarely the case. Variations among individuals
is the rule, as is vividly evident at a hog show or on a lively dance floor. Such
variations are partly due to physiological adjustment or acclimation, and
partly inherited. The genetically determined characteristics transmitted from
generation to generation are not identical. In fact, variation among
individuals in a population permits gradual change that results in adaptation.
Without such change, there would be no evolution. Changes in the
population through time may even lead up to the formation of new species,
called speciation.
What are the sources of genetic change? That is, how do the genes in the
DNA of cells change? We recognize two major ways, mutation and
recombination. Mutation, such as that caused by irradiation or certain
chemicals, is a change in the sequence of building blocks of a DNA molecule.
Most mutations are lethal or damaging, but reproductive cells that survive
mutations contain an altered DNA structure. Recombination

ECOLOGICAL CONCEPTS IN BRIEF

 causes variation in offspring by a different process. When male and female
parents reproduce, each offspring may receive a different combination of
chromosomes from the parents, and so each offspring differs. In this manner
no DNA is altered, but the existing DNA of the male and female parents is
simply combined in a different way in the offspring. Variation also results
from alterations in chromosome structure or number during the cell division
process.
Changes in the frequencies of genes in a population from generation to
generation are evolutionary changes. Four factors tend to make genetic change
and evolution ubiquitous.

1. Nonrandom mating among members of the population occurs, so

genes of frequently mating individuals increase.
2. Immigration brings new genes and emigration eliminates certain genes
from the population.
3. Mutations often occur and introduce or eliminate certain genes.
4. When populations are small, changes in the frequency of genes may
take place by chance (genetic drift and founder effect).

Any of these factors causes changes in gene frequencies from generation to

generation, and thereby the evolution of the species, through the general
process of natural selection.
If we go into the field, we may observe at least the essence of natural
selection as originally described by Darwin. An overpopulation of offspring
occurs. The population contains variability among the offspring.
Competition for resources ensues. Finally, survival of the fittest (best-
adapted) offspring results. This natural selection process, going on around
us all the time, does not eliminate the variability in a population. But it does
lead to organisms better adapted to their environment and it permits
adaptation to new or changing environments.
Natural selection as described focuses on individuals within a population.
Kin selection, in contrast, refers to changes in small groups of individuals,
such as family groups, that function as units relative to other units in the
population. In kin selection the fitness of small related groups changes,
intriguingly, the fitness of individuals may change in a direction opposite to
that of the kin group. Thus, in certain vertebrate species the group appears
to be more important than the individual.

Speciation

If there were random reproduction among all members of a population,

genetic change and evolution might not occur. But in real populations, many
factors prevent random reproduction. We call them isolating mechanisms or
barriers. Geographical barriers such as mountains and large

ECOLOGICAL CONCEPTS IN BRIEF 4

bodies of water are obvious. But other barriers may effectively separate patches at a
finer scale within a landscape. For example, various reproductive and other isolating
mechanisms are just as effective as geographic barriers in preventing random
reproduction in a population. An isolation mechanism causes two (or more) groups
of individuals to become increasingly different, because individuals interbreed and
mix genes within their group, but mating and gene flow between groups is inhibited.
When the gene pools of two groups are only slightly different (whether
geographically isolated or not), we may refer to them as populations within a species
(Figure 2.12). If gene flow between such populations remains limited over time, they
will diverge to the point at which we call them races, ecotypes, varieties, or
subspecies. Individuals of such populations can potentially interbreed if brought
together under appropriate conditions, so there is still only one species. Flowever, if
gene flow between these populations remains limited, they will diverge still further
genetically, and gene flow will cease altogether. When they are no longer able to
interbreed when brought together, we have two species. This process is called gradual
spéciation. A process of punctuated spéciation has also been proposed, in which a
population splits off from a species and becomes a new species rapidly, not by the
long, gradual process of reproduction over numerous generations.

i^Ty\/^y/Twospecles
Gene flow
_ One species
Time
Two races, ecotypes,
Figure 2.12 A gradual spéciation process. An
varieties, or subspecies
isolating mechanism appears and divides the
initial population in two parts. The amount of gene flow within
or between populations is indicated by the thickness of arrows.
Two populations

One population

ECOLOGICAL CONCEPTS IN BRIEF 5

 Many patterns of spéciation can be recognized around the globe; we note
here two particularly distinctive or frequent ones. A species is often found
in a series of populations with progressively different characteristics,
extending over a geographic area. This pattern, called a dine, may result
over time in the extremes being so different they become different species,
while intermediate forms are still present between them. In a second case,
adaptive radiation, a species encounters new environmental conditions and
diversifies into a number of species.

ECOLOGICAL COMMUNITIES

Spatial Structure of a Community

In studying a population, we examine the individual organisms and how they
are clustered by age, sex, and genetic makeup. The ecological community, at
the next higher organizational level, may be described as the assemblage of
species populations at a particular place and time. Usually there are
interactions among the species of a community, and in many regions
distinctive species clusters are recognizable. Because both species and
species clusters differ greatly, a wide range of patterns and measurements
is found in community ecology. We study species composition (what particular
species are present), species diversity or richness (how many species are
present), dominance (which species are most abundant and by how much),
rare species, and the growth forms or life forms present (such as trees, shrubs,
succulents). We also look for guilds, groups of species which use a type of
resource in a similar way.
The vertical structure of an above ground plant community is a product
of the size and the branching and leaf characteristics of plants, and the
internal distribution of leaves is largely dependent on light penetration.
More or less distinct vegetation layers may be present, as in a forest with
canopy, subcanopy, understory, shrub layer, and herb layer, each receiving
progressively less light. The animal species also sort out vertically, with
certain animals mainly in the canopy, others mainly in the understory, and
so on. Vertical stratification in animal and microbial communities in the soil
is also marked. Oxygen level is a primary determinant of this vertical pattern,
though the amounts of organic matter, mineral nutrients, water, and heat
also strongly correlate with depth. In aquatic communities, vertical
stratification is pronounced and largely due to temperature, oxygen, and
light gradients.
The horizontal distributions of species exhibit complex patterns. At one
extreme, the boundary between two types of vegetation, such as field and
forest, may be relatively abrupt. In this case the overlap zone or ecotone is
narrow and composed mainly of species from both sides
intermixed. Sometimes the overlap zone is wide, and in it a mosaic of patches of each
vegetation type intermingle (Rapoport, 1982b). At the other extreme, no ecotone is

ECOLOGICAL CONCEPTS IN BRIEF 6

easily distinguishable, and we find a gradual change in species composition over
distance, called a continuum.
These different separations of vegetation types have sparked considerable
controversy and have given rise to two theories about the essential nature of a
community. If a continuum is present everywhere, interactions among species vary
continuously over space. Thus the integrated nature of a community is minimal; that
is, a species apparently does not require the presence of other specific species. From
this perspective, called the individualistic theory, the community is simply the
assemblage of species that happen to coexist at some point in time and at some
point along the continuum. The types of communities are as numerous as the points
along the continuum, and no community type is much more frequent than any other.
Alternatively, the so-called community unit theory recognizes the existence of primary
community types separated by eco- tones. This theory assumes some
interdependence among the species of a community.
In a continuum, when we plot the abundance of species against some controlling
environmental factor such as food supply or soil moisture, a series of broadly
overlapping bell-shaped curves (each representing a species) may appear (Figure
2.13a). When we plot the same species abundances against distance on land, we
sometimes find the same result (Figure 2.13b). Usually, though, we find flatter curves
with several species dropping out at, about the same place and less overlap at that
place, basically an ecotone between communities (Figure 2.13c). Why? Apparently the
continuum is present where the major controlling environmental factor varies evenly
(linearly) with distance, as when temperature changes evenly from the bottom to the
top of a mountain. Usually, however, environmental factors are unevenly or patchily
distributed over space. The result is a sharper ecotone, where the amount of a factor
changes abruptly in a short space (Figure 2.13c), such as that along the edge of
certain swamps or between a forest and adjacent cultivation.
When individual trees or small groups of trees within a community die, a gap is
produced where high light conditions prevail down to the ground. Larger patches in
the community may be produced by disturbances such as fire or pest defoliation.
Furthermore, the physical environment itself may be patchy, having rock outcrops or
wet spots. In all cases the patchiness of the community forms a mosaic.
Continuua observed over lengthy spans are primarily limited to natural
landscapes, in areas with little disturbance or environmental patchiness. Natural
landscapes with frequent environmental patchiness or natural disturbance primarily
have ecotones, and this pattern is even more pronounced in landscapes modified by
human influences.

ECOLOGICAL CONCEPTS IN BRIEF 7

(b ) 0 10 20 30 40 50
Distance (m or km)

Environmental factor gradient

Environmental factor gradient

Environmental factor gradient

(c) 0 10 20 30 40 50
Distance (m or km)
Figure 2.13 Distribution of species along environmental and distance
gradients. Each bell-shaped or flat-topped curve represents a species. The
gradients of environmental factors such as temperature, phosphorus level, or
food supply range from low to high, (a) Response of each species to an
environmental factor determined by laboratory experiments, (b) Species
distributions where the controlling environmental factor is directly correlated
with distance, and (c) where the controlling environmental factor is patchily
distributed. Pattern c is common in landscapes.

Plant and animal communities are often mapped, and to do this they are named
and classified, as much as possible to show their relationships to one another. The
primary criteria used in classifications are appearance (physiognomy), species
composition, dominant species, and habitat. The

8 OVERVIEW
 major divisions in the various classification systems differ in their use of
these criteria, but at finer divisions, species composition is almost always
used as the classification criterion.

Ecological Niche and Species Diversity

Many ecologists agree that the ecological niche is the functional role of a
species in a community, including the environmental variables affecting the
species. The totality of environmental variables and functional roles to which
a species is adapted is called a fundamental niche. However, a species normally
does not occupy that totality but only a portion of it, called a realized niche.
The difference between the fundamental and realized niches is often
ascribed to competition, though other factors may be important.
Niche width, a measure of the variety of different resources exploited by an
organism, provides further insight into the roles of a species in a community.
A broad niche indicates that the species is a generalist, sacrificing efficiency
in the use of a narrow range of resources in order to use a wide range of
resources. A narrow niche indicates a specialist species, one that exploits
efficiently a narrow range of resources on which it depends. One way many
species coexist in a community is by specialization, since it reduces
competition.
Ecologists use species diversity as one important measure of the struc-
tural heterogeneity of a community. There are many ways a community can
be diverse in species. They may vary in number (richness) of species, degree
of dominance by one or a few species, relative abundance of all species
(evenness), number of rare species, number of nonnative species, vertical
stratification of species, horizontal patchiness, number of growth or life
forms, and so forth. Species diversity measurements are used to help
interpret mechanisms operating in the community.
Various indices have been constructed that combine two or more of these
measures into a single ijumber. For example, the Shannon-Wiener (or
information) index combines number and relative abundance of species
(Peet, 1974). A synthetic index is convenient in reducing two or more
variables to a single number, but because the variables are not separated
the mechanisms behind them are less likely to be identified. Number of
species is the single clearest measure of species diversity, although
measuring additional separate variables adds interpretive power.
Measurements of bird species diversity, plant species diversity, and the
like, turn out to be very coarse measurements. Thus, as a finer community
characteristic, guild diversity is measured, for example, birds that feed on
seeds, catch flying insects, or eat flesh. In this way a more penetrating view
of the several mechanisms operating in the community emerges than if
species diversity of birds as a whole, for example, is measured.
When disturbance causes the formation of a bare area, plants and animals
colonize this bare area (Figure 2.14). Then, over time, species replace one
another until the community consists primarily of species which can

ECOLOGICAL CONCEPTS IN BRIEF 9

 successfully reproduce where they are, that is, a climax community. This
directional species replacement process is called succession.
We mentioned the case of small clusters of trees blowing down or dying
in a forest, producing scattered gaps. If we study such gaps over time, we
may see an initial colonization by herbaceous plants, followed by takeover
by small woody plants, followed by takeover by trees, fol-

Figure 2.14 Succession on slides (landslips) in the boreal forest. The disturbance patch resulted from
vegetation and soil (and perhaps snow and ice) sliding down the slope, probably when the soil was
saturated with water. Note that the older slide to the right has revegetated with birches (Betula) and
aspens (Populus). The remainder of the forest is spruce (Picea) and fir (Abies). Parc de la Caspésie,
Québec, Canada. (R. Forman.)
lowed by a blowing down, and the cycle starts again. This process is called cyclic
succession since it progresses through a series of stages and returns to a former
stage. In this example, the tree stage lasts much longer than the other stages, but in
the cyclic successions of bogs, tundra, grassland, and shrubland the stages are more
equal in length.

ECOLOGICAL CONCEPTS IN BRIEF 10

 In the above case the primary energy source is plant photosynthesis. In
heterotrophic succession the primary energy source is nonphotosynthetic organic
matter such as elephant droppings, a fallen log, or organic waste dumped in a stream.
The organic matter is consumed by decomposers, and a series of stages then occurs
that involves a food web based on those decomposers. In all these examples of
succession, colonization, species composition changes, and local extinction rates are
high in the early phases and decline over time.
In an abandoned tobacco field in the southeastern United States, the main
successional stages progress from herbaceous annuals to herbaceous perennials, to
grass, to shrubs, to successional trees such as pine (Pinus), to climax trees such as
oak and hickory (Quercus, Carya). However, one specific point within the field may go
through this sequence but skip the shrub stage by having a pine seedling directly
colonize the grass there. In similar fashion, other points in the field could skip the
grass stage or even two stages. In other words, in point succession each adjacent
point may undergo a different sequence, while the field as a whole goes through all
the stages listed as a kind of average of all the points in the field.
Disturbance is an important mechanism of succession, as are external
modifications ol the environment such as weather or dust deposition. Each
community includes its own proportion of species that replace one another in various
ways, and each differs in the relative importance of disturbance and external factors
during succession.
The climax community is a mosaic containing patches with species of earlier
successional stages (Figure 2.14). The mosaic may simply include small patches
relating to natural deaths of organisms, by herbivore grazing, hunting by people, or
tree falls, for example. It may also include large patches caused by disturbances such
as fire and pest outbreaks.
When fires or pest outbreaks, for example, are frequent and widespread, species
adapt to them, and it becomes questionable what the climax is. In New Jersey (United
States) Pine Barrens fires have been frequent and widespread, so the area is always
dominated by pine. But in a few isolated spots near old human settlements wildfire
has been eliminated or minimized for a century or more, and an old oak forest is
present. Which is the climax, pine or oak forest? Oak is the theoretical climax if fire
could be artificially eliminated. Pine, though, is the actual climax, since it maintains
itself with fire as a part of its ecosystem. Indeed,

ECOLOGICAL CONCEPTS IN BRIEF 11

 here wildfire prevention is a significant disturbance because it effectively leads to elimination of the pine
forest.

Island Biogeography

Charles Darwin wrote a book on island biogeography, and island communities have always intrigued
ecologists. For understanding the species diversity (number of species) on an island, a model was recently
developed that interrelates the colonization and extinction rates of species (Figure 2.15). The hypothesis is
that with more species present on an island, hence increased competition, the rate of colonization of new
arrivals would be lower. Also, the rate of extinction of species would be higher. The point at which the
idealized colonization and extinction curves cross is thus the predicted number of species on an island.
It was additionally proposed that islands near a species source, such as a mainland, would have more
diversity than isolated islands, because the rate of arriving species and colonists would be higher on nearby
islands (Figure 2.15). Similarly, more species are predicted on large islands than small islands because the
extinction rate is lower on large islands. Confirmation of the island biogeographic model has proven elusive
despite much interest. Several studies have supported aspects of it and certain predictions from it, but other
studies have demonstrated quite different patterns. The model has been applied to patches on land with

Colonization Extinction Colonization Extinction

Species diversity Species diversity

(a) (b)
Figure 2.15 Hypothesized effects of colonization and extinction on species diversity of islands with different characteristics, (a) Island
size: Ss and SL are the number of species on small and large islands, respectively, (b) Island isolation: SF and SN are the number of
species on islands far from and near a species source, respectively. After MacArthur and Wilson (1967).

ECOLOGICAL CONCEPTS IN BRIEF 12

limited success. The problems in testing the model are the limited knowledge of extinction rates, the difficulty of
eliminating other major variables, and the difficulty of doing appropriate experiments.
The model has stimulated empirical studies on archipelagoes and the plotting of species-area curves (Figure 2.16).
These curves agree with the prediction that larger islands will have more species, but the mechanism is quite different,
namely, there is more habitat diversity on larger islands. The curves seem to better support the idea that islands near
species sources have more species. By plotting an archipelago curve based on samples of a few islands one may predict
the species diversity on other islands with some accuracy.
An equation relating species diversity 5 to island area A

S = cA2

has generated interest because z, the slope of the line relating species and area (Figure 2.16), appears to be relatively
similar in different archipelagoes and for different types of organisms, (c is a constant measuring the number of species
in a unit area of island). The variation in z may simply be a mathematical property or may be indicative of different
ecological patterns on archipelagoes.

Patch area Logarithm of patch area

(a) (b)
Figure 2.16 Species-area curve for an archipelago, plotted (a) linearly and (b) logarithmically. The
linear plot shows only a slight gain in species between the minimum patch point M and the asymptote diversity level or species pool P. A steep G|
slope (initial species gain) characterizes relative homogeneity in the distribution of species over the archipelago, whereas a lower slope is produced
by relative heterogeneity or an abundance of rare species.

ECOLOGICAL CONCEPTS IN BRIEF 13

Biogeography in Geologic Time
The animal and plant communities of today are the latest word in a long history of evolutionary developments
and species migrations. Violent geological events took place in the middle of the Tertiary period, about 20
million years ago (see Chapter 7, Appendix). Huge mountain chains were thrown up: the Rocky Mountains, the
Andes, the Himalayas, and the Alps. Extensive dry regions formed downwind of these mountains as rain shadows,
often near the centers of continents. Pockets of arid evergreens and grasses spread rapidly, forming deserts,
grasslands, and Mediterranean type vegetation. Temperate forests at high latitudes and tropical forests at low
latitudes were separated from one another.
During the Tertiary, most of the plant species that are dominant on earth today evolved. Birds, mammals,
and insects also became widespread around the globe, replacing the dinosaurs and other reptiles which had
previously predominated (Figure 2.17). By the end of the Tertiary, many of today's families and genera
(subdivisions of families) of these animals had evolved, though most of the species were different.
Between two million years ago and about 8000 B.C. came the Pleistocene period, during which at least four
massive glaciations occurred, separated by warmer periods when much of the ice melted. Faunas and floras were
constantly moving or migrating across the land, at times being compressed into one or many isolated pockets
and at times spreading to cover large expanses. During this process many of our present terrestrial animal
species evolved (Figure 2.17).
The past ten or so millennia since the Pleistocene compose the present period, the Holocene, a time marked
by two major factors affecting animal and plant communities. First, climate has changed, primarily with a general
warming trend over the whole period, but also with a hot phase that began about 7500 years ago and later
cooled. Second, human activity has become widespread. Deforestation and agriculture, in particular, which
began some 8000 years ago, accelerated in the past four millennia. Very recently the effect of an industrial
society has changed the earth's flora and fauna at a pace even more rapid than that of glaciation, climatic change,
or agriculture.

ENERGY AND MATTER IN ECOSYSTEMS

Energy
Energy flows in a predictable way through an ecosystem. Solar energy is converted by photosynthesis to chemical
energy in plants. A portion of the energy then returns directly to the atmosphere by the respiration of plants.
Another portion of the energy goes to herbivores (see Figure 2.17),

ECOLOGICAL CONCEPTS IN BRIEF 14