Chapter Four: Ecosystems: How They Change

4.1 Dynamics of Natural Populations

I. Each species in an ecosystem exists as a population, or reproducing group. In any
population, births and deaths will cause the population to grow or shrink.
A. If births and deaths are more or less equal over time, the population is said to be in
equilibrium.
Population Growth Curves
I. Every species has the capacity to increase its population when conditions are favorable.
The growth of a population under ideal conditions will be exponential.
A. Such a series is called an exponential increase. This sort of growth results in a
population explosion.
B. A basic feature of exponential increase is that the numbers increase faster and
faster as the population doubles and redoubles, with each doubling occurring in
the same amount of time. If we plot numbers over time during an exponential
increase, the pattern produced is commonly called a J- curve.
C. There are situations that lead to population explosions in a species. However, as
the population continues to expand, one of two things will occur: 1) natural
mechanisms may cause the population to level off and continue at dynamic
equilibrium. This pattern is known as an S-curve. 2) in the absence of natural
enemies, the population keeps growing until it exhausts essential resources and
then dies off.
II. After a J-curve crash, several things can occur.
A. For a herbivore species, any one of three scenarios may unfold. First, if the
ecosystem has not been seriously damaged, the plant food resource may recover,
allowing the herbivore population to recover, and the J-curve may be repeated.
Second, after the initial J, natural mechanisms may come into play as the
ecosystem recovers, thus bringing the population into an S-balance. In the third
scenario, damage to the ecosystem is so severe that recovery is limited, and small
surviving populations eke out an existence in a badly degraded environment.
III. The outstanding feature of natural ecosystems—ecosystems that are more or less
undisturbed by human activities—is that they are made up of populations that are
usually in dynamic equilibrium represented by S-curves. J-curves come about when
there are unusual disturbances, such as the introduction of foreign species, the
elimination of a predator, or the sudden alternation of a habitat.
A. The increases represented by J-curves are only temporary in animal populations,
because the animals inevitably die off as resources are exhausted.
Biotic Potential versus Environmental Resistance
I. The ability of a population to increase is known as biotic potential: it is the number of
offspring that a species may produce under ideal circumstances.
A. The biotic potential of different species varies tremendously.
 B. To have any effect on the size of subsequent populations, the young must survive
and reproduce in turn. Survival through the early growth stages to become part of
the breeding population is called recruitment.
II. There are two common reproductive strategies in the natural world. The first is to
produce massive amounts of young, but then leave survival to the whims of nature.
This strategy often results in very low recruitment. Thus, despite a high biotic
potential, a population may not increase at all because of low recruitment. However,
this strategy is highly successful if a species is adapted to an environment that can
suddenly change and become very favorable.
A. Organisms with this strategy are usually small, with rapid reproductive rates and
short life spans.
B. The second strategy is to have a much lower reproductive rate, but then care for
and protect the young until they can compete for resources with adult members of
the population. This strategy works best where the environment is stable and
already well populated by the species. Organisms with such a strategy are larger,
longer lived, and well adapted to normal environmental fluctuations.
III. Additional factors that influence population growth and geographic distribution are the
ability of animals to migrate, or of seeds to disperse, to similar habitats in other
regions; the ability to adapt to and invade new habitats; defense mechanisms; and
resistance to adverse conditions and disease.
A. All of these factors are components of an species’ life history, and represent a
particular strategy for reproduction and survival that enables the species to be
successful in a unique ecological niche in the ecosystem.
IV. Population explosions are seldom seen in natural ecosystems, because biotic and abiotic
factors tend to cause mortality in populations.
A. Among the biotic factors are predators, parasites, competitors, and lack of food.
Among the abiotic factors are unusual temperatures, moisture, light, salinity, pH,
lack of nutrients, and fire.
B. The combination of all the biotic and abiotic factors that may limit a population’s
increase is referred to as environmental resistance.
V. In general, the reproductive ability of a species remains fairly constant because that
ability is part of the genetic endowment of the species. What varies substantially is
recruitment. It is in the early stages of growth that individuals are most vulnerable.
A. Consequently, environmental resistance effectively reduces recruitment. If
recruitment is at the replacement level—with just enough to replace the adults—
then the population will remain at equilibrium. If recruitment is insufficient to
replace losses in the breeding population, then the population will decline.
VI. There is a definite upper limit to the population of any particular plant or animal that an
ecosystem can support. This limit is known as the carrying capacity.
 A. The carrying capacity is the maximum population of a species that a given habitat
can support without the habitat being degraded over the long term. If a population
greatly exceeds the habitat’s carrying capacity, it will undergo a J-curve crash.
B. Because conditions within a habitat can change from year to year, the habitat’s
carrying capacity varies accordingly.
VII. In general, a population’s biotic potential remains constant, so it is changes in
environmental resistance that allow population to increase or cause them to decrease.
Population balance is a dynamic balance, which means that additions and subtractions
are occurring continually and the population may fluctuate around a median.
A. Some populations fluctuate very little, whereas others fluctuate widely. As long as
decreased populations restore their numbers and the ecosystem’s carrying
capacity is not exceeded, the population is considered to be at equilibrium.
Density Dependence and Critical Number
I. The size of a population generally remains within a certain range when environmental
resistance factors are density dependent. That is, as population density increases,
increases, environmental resistance becomes more intense and causes such an
increase in mortality that the population growth ceases or declines.
A. As population density decreases, environmental resistance lessens, allowing the
population to recover.
II. Factors in the environment that cause mortality can also be density independent. That is,
their effect is independent of the density of the population. This is frequently true of
abiotic factors.
A. Although density-independent factors can be important sources of mortality, they
are not involved in maintaining population equilibria.
III. There is no guarantee that a population will recover from low numbers. The survival and
recovery of a population depends on a certain minimum population base, which is
referred to as the population’s critical number.
A. Often a group is necessary to provide protection and support for its members.
B. If a population is depleted below the critical number needed to provide supporting
interactions, the surviving members become more vulnerable, breeding fails, and
extinction is almost always inevitable.
C. Concern for declines in species led to the Endangered Species Act, which calls for
the recovery of two categories of species. Species whose populations are
declining rapidly are classified as threatened. If the population is near what
scientists believe to be its critical number, the species may be classified as
endangered.
4.2 Mechanisms of Population Equilibrium
I. Only those factors that are density-dependent are capable of actually regulating the
population, keeping it around an equilibrium.
 II. Top-down regulation is the control of a population by predation. In bottom-up regulation,
the most important control of a population occurs as a result of the scarcity of some
resource.
Predator-Prey Dynamics
I. In both predator and prey species, other factors may influence the observed fluctuations
in population densities.
II. As the prey population is culled down to those healthy individuals which can escape
attack, the predator population will decline, unless it can be switched to other prey.
A. The predators are limited by the availability of their food source.
B. A predator-prey relationship involves both top-down and bottom-up population
regulation. Density-independent factors can also play a role.
III. Parasitic organisms affect the populations of their host organisms in much the same way
that predators do their prey—in a density-dependent manner.
A. As the population of the host increases, parasites and their vectors have little
trouble finding new hosts, and infection rates increase, causing higher mortality.
Conversely, when the density of the host is low, the transfer of infection is less
efficient, so the levels of infection are greatly reduced, thus allowing the host
population to recover.
B. A parasite can work in conjunction with a predator to control a given herbivore
population.
IV. If herbivores eat plants faster than the plants can grow, the plants will eventually be
depleted, and the animals will suffer.
A. The consequences of overgrazing are not just to the herbivore in question. One or
more types of vegetation may be eliminated and replaced by other forms or not
replaced at all, leaving behind a degraded ecosystem.
V. Keystone species maintain a crucial role in an ecosystem’s biotic structure.
Competition
I. When species compete, their ecological niches are said to overlap. If any resource is in
short supply, the most intense competition will come from members of the same
species. This is called intraspecific competition.
A. Competition is a form of bottom-up regulation, because it occurs only when a
resource is limited.
II. Territoriality often controls the populations of carnivores and some herbivores.
Territoriality refers to individuals or groups defending a territory against the
encroachment of others of the same species.
A. Territoriality, therefore, is intraspecific competition.
B. In territoriality, what is really being protected by the defender or sought after by
the invader is the claim to an area suitable for nesting, for establishing a harem, or
for adequate food resources. Hence, the territory is defended only against others
that would cause the most direct competition for those resources—members of the
same species.
 C. As a consequence of territoriality, some members of the population are able to
nest, mate, or gain access to sufficient food resources or rear the next generation.
In this way, a healthy population of the species survives. If, by contrast, there
were an even rationing of inadequate resources to all members, the entire
population would become malnourished and might perish.
D. Through territoriality, breeding is restricted to only those individuals capable of
claiming and defending a territory; thus, population growth is curtailed in a
density dependent manner.
III. Individuals unable to claim a territory are usually the young of the previous generation.
Some may hang out of the fringes and seize the opportunity as they mature and older
members with territories weaken or perish. Some fall prey to various environmental
resistance factors.
A. Some may be driven to disperse, either to find another region where they can
successfully breed or to perish in conditions beyond their limit of tolerance along
the way.
B. Territoriality is a powerful force behind the dispersal, as well as the stabilization,
of populations.
IV. When plants produce a large number of seedlings in a limited area, self-thinning is the
result. Self-thinning is density-dependent.
A. Crowded conditions always lead to competition for resources between both plants
and animals. This leads to evolutionary changes in animals.
B. Every factor of environmental resistance is a selective pressure resulting in the
survival and reproduction of those individuals with a genetic endowment that
enables them to better cope with their surroundings. This is the essence of natural
selection.
V. Interspecific competition, therefore, has two distinct kinds of impact on the population of
a species. In the short term, it can lead to the density-dependent regulation of a
species population, through such factors as territoriality and self-thinning. It can also
lead, however, to long-term improvements, in that the species adapts to its
environment because those better able to compete are the ones who survive and
reproduce, and their superior traits are passed on to successive generations.
A. An important factor affecting the competition between plant species is the fact
that a single species generally cannot utilize all of the resources in a given area.
Therefore, any resources that remain may be claimed by other species having
different adaptations.
B. Another option is mutualism, an arrangement between two species whereby both
benefit.
C. Plants usually exist in communities with a rich mixture of species, so that a given
individual is likely to be in proximity with any of a number of species, as well as
with individuals of their own species. It is thus more likely that the size of
 neighboring plants will play a larger role than the actual species in determining
the intensity of competition.
D. Resources vary in intensity and over space and time, so there is a heterogeneous
environment for plants, allowing different species to coexist because they use
common resources in different ways. Ordinarily, no one species is able to
outcompete all the others because the different species have become specialized
in different ways in their use of resources and their tolerance to abiotic conditions.
E. When a competitor is removed competitive release occurs. That is, once the
strong competitor is absent, the weaker one thrives and exploits resources it is
normally unable to acquire because of the presence of its competitor.
F. Interspecific competition between animals may be a strong factor influencing the
distribution and abundance of species. In the short run, interspecific competition
can reduce the success of species and even eliminate them from the environment;
thus, it can help regulate a population. In the long run, interspecific competition
helps drive natural selection, bringing about greater specialization of ecological
niches and allowing resources to be divided up among species.
Introduced Species
I. Over the past 500 years, and especially now that there is a vast global commerce,
thousands of species of plants, animals, and microbes have been accidently or
deliberately introduced onto new continents and islands.
A. Most of the important insect pests in croplands and forests are species introduced
from other continents.
B. The problem of introduced species is increasing due to expanding world trade and
travel.
II. The regulation of populations is a matter of complex interactions among the members of
the biotic community. The relationships are specific to the organisms in each
particular ecosystem.
A. When a species is transported over a physical barrier from one ecosystem to
another, it is unlikely to fit into the framework of relationships in the new biotic
community. In most cases, it finds the environmental resistance of the new system
too severe and dies out.
B. In some instances, the introduced species simply joins the native flora and fauna
and does no measurable harm. The new species has then become naturalized.
C. In the worst cases, the transported species becomes invasive. It finds physical
conditions and a food supply that are hospitable, together with an insufficient
number of natural enemies to stop its population growth. Then its population
explodes, and it drives out native species by outcompeting them for space, food,
or other resources.
III. One solution to the takeover by an invasive species may be to introduce a natural enemy.
Unfortunately, a single enemy that will control a pest may not exist.
 A. There is no guarantee that the natural enemy, when introduced into the new
ecosystem, will focus its attention on the target pest.
IV. Most invasive species are not problems in their native lands. Their impact is so different
in a new setting because ecosystems on different continents or remote environments
have been isolated for millions of years. Consequently, the species within each
ecosystem have developed adaptations to other species within their own ecosystem,
and these are independent of adaptations that have developed in other ecosystems.

IV.3 Evolution as a Force for Change

Adaptation Through Natural Selection
I. Predation and competition are two important mechanisms that keep natural populations
under control. Predators and prey become well adapted to each other’s presence.
A. Predators are rarely able to eliminate the prey species, largely because the prey
have various defenses against their predators.
B. Intraspecific competition also represents a powerful force that can lead to
improved adaptations of a species to its environment. Interspecific competition
promotes adaptations in the competitors that allow them to specialize in
exploiting a resource. This specialization can lead to resource partitioning that
allows many potential competitors to share a basic resource.
II. Most young plants and animals in nature do not survive; instead they fall victim to
various environmental resistance factors. These factors—predators, parasites,
drought, and other—are known as selective pressures. That is, each factor can affect
which individuals survive and reproduce and which are eliminated.
A. Predators may be seen as a selective force favoring the survival of traits that
enhance the prey’s ability to escape or protect itself and causing the elimination of
any traits handicapping those functions.
B. Every factor of environmental resistance is a selective pressure resulting in the
survival and reproduction of those individuals with a genetic endowment that
enables them to cope with their surroundings. In nature, is a constant selection
and, consequently, modification of a species’ gene pool toward features that
enhance survival and reproduction within the existing biotic community and
environment. Because this process occurs naturally, it is called natural selection.
C. The modification of the gene pool of a species by natural selection over the course
of many generations is the sum of biological evolution.
III. Under the selective pressures exerted by the factors of environmental resistance, the gene
pool of each population is continually tested. Virtually all traits of organisms can be
seen as features that adapt the organism for survival and reproduction, or fitness.
A. Essentially all characteristics of organisms can be grouped as follows:
1. Adaptations for coping with climatic and other abiotic factors.
2. Adaptations for obtaining food and water, or other nutrients.
 3. Adaptations for escaping from or protecting against predation and for
resistance to disease-causing or parasitic organisms.
4. Adaptations for finding or attracting mates or for pollinating and setting seed.
5. Adaptations migrating or for dispersing seeds.
B. When facing a new, powerful selective pressure, species have three alternatives:
1. Adaptation. The population of survivors may gradually adapt to the new
condition through natural selection
2. Migration. Surviving populations may migrate and find an area where
conditions are suitable to them
3. Extinction. Failing the first two possibilities, extinction is inevitable.
C. For adaptation to occur, there must be some individuals with traits that enable
them to survive and reproduce under the new conditions. There must also be
enough survivors to maintain a viable breeding population. If the breeding
population is maintained, the process of natural selection should lead to increased
adaptation over successive generations.
IV. There are four key variables among species that will affect whether or not a viable
population of individuals is likely to survive new conditions: 1) geographical
distribution, 2) specialization to given habitat or food supply, 3) genetic variation
within the gene pool of the species, and 4) the reproductive rate relative to the rate of
environmental change.
A. The rate at which changes in the environment occur is an important consideration.
If environmental changes are very slow, populations of slowly reproducing
species may be able to adapt. When changes in environmental conditions occur
more rapidly, however, more and more species will be eliminated in the contest to
adapt accordingly.
V. The infusion of new variations from mutations and the pressures of natural selection
serve to adapt a species to the biotic community and the environment in which it
exists. Over time, in this process of adaptation, the final product may be so different
from the population that started the process that it is considered a different species.
This is one aspect of the process of speciation.
A. Reproductive isolation is critical, because if subpopulations continue to
interbreed, all the genes will continue to mix through the entire population,
keeping it as one species.
B. Separated subpopulations must be exposed to different selective pressures. As the
separated species adapt to these pressures, they may gradually become so
different as to be considered different species and thus be unable to interbreed
with one another.
Drifting Continents
I. Geographic isolation is foundational to the process of speciation.
II. Around 22.5 million years ago, the continents were fused together into a “supercontinent”
we call Pangaea.
 A. The theory that the continents are in motion—plate tectonics—helps us
understand earthquakes and volcanic activity, and is a key to understanding the
geographic distribution of present-day biota.
III. Within Earth’s semimolten interior, hot material rises toward the surface and spreads out
at some location while cooler material sinks toward the interior at other locations.
Riding atop these convection currents, the tectonic plates move slowly with respect to
one another.
A. The spreading process of the last 22.5 million years has brought the continents to
their present positions and accounts for the other interactions between tectonic
plates.
B. Movement of the crust itself is not gradual, for the plate boundaries are locked by
friction and hence are regions of major disturbances.
IV. Tectonic movement may gradually lead to major shifts in climate in three ways. First, as
continents gradually move to different positions on the globe, their climates change
accordingly. Second, the movement of continents alters the direction and flow of
ocean currents, which in turn have an effect on climate. Third, the uplifting of
mountains alters the movement of air currents.

IV.4 Ecosystem Responses to Disturbance

I. According to equilibrium theory, ecosystems are stable environments in which species
interact constantly in well-balanced predator-prey and competitive relationships.
Thus, biotic interactions determine the structure of living communities within
ecosystems. However, time can bring disturbances.
A. Ecosystems are very patchy environments. Conditions and resources within them
vary with temperature, moisture, exposure to sunlight, soil conditions, etc. the
distribution of species is similarly patchy, reflecting the patchiness in conditions.
B. The patchiness of many ecosystems has caused many ecologists to think of them
as nonequilibrium systems that seldom exhibit the characteristics of a true
equilibrium.
Ecological Succession
I. The transition from one biotic community to another is called ecological succession.
Succession occurs because the physical environment may be gradually modified by
the growth the biotic community itself.
A. Pioneer species start the process, but as these grow, they create conditions that are
more favorable to more longer-lived colonizers, and the process in general is
driven by the changing conditions that pave the way for other species. This
process is known as facilitation.
II. The succession of species does not go on indefinitely. A stage of development is
eventually reached in which there appears to be a dynamic balance between all the
species and the physical environment.
 A. This final state is called a climax ecosystem because the assemblage of species
continues on in space and time.
B. All balances are relative to the current biotic community and the existing climatic
conditions. Therefore, even climax ecosystems are subject to change if climatic
conditions change or if a new species is introduced or old ones are removed.
C. Natural succession may be seen as a progression toward a relatively more stable
climax—one that no longer changes over time.
III. If the area has not been occupied previously, the process of initial invasion and then
progression from one biotic community to the next is called primary succession.
A. The nature of the climax ecosystem differs according to the prevailing abiotic
factors of the region, yielding the biomes typical of different climatic regions.
B. The major difference between primary and secondary succession is that secondary
succession starts with preexisting soil.
C. Natural succession also occurs in lakes or ponds. Succession occurs because the
soil particles inevitably erode from the land and settle out in ponds or lakes,
gradually filling them. Aquatic vegetation also creates detritus that contributes to
the filling process. As the buildup occurs, terrestrial species from the surrounding
ecosystem can advance in the ecotone and aquatic species must move further out
into the lake.
Disturbance and Resilience
I. In order for natural succession to occur, the spores and seeds of the various invading
plants and the breeding populations of the various invading animals must already be
present in the vicinity. Ecological succession is a matter of populations of existing
species taking advantage of a new area as conditions become favorable.
A. In any given ecosystem, all stages of succession are likely to be represented. This
is so because disturbances constantly create gaps or patches in the landscape.
B. When a variety of ecological stages is present in a landscape, a greater diversity
of species can be expected. If certain species have been eliminated, natural
succession will be blocked or modified.
II. Fire is an abiotic factor that has particular relevance to succession. It is a major form of
disturbance common to terrestrial ecosystems.
A. Different species have different degrees of tolerance to fire. Some plant species
even depend on fire. Ecosystems that depend on the recurrence of fire to maintain
their existence are called fire climax systems.
III. Disturbances and shifting biotic relationships not only may have little detrimental effect
on an ecosystem, but may actually contribute to its ongoing functioning. Ecologists
refer to this condition as resilience. Thus, a resilient ecosystem is an ecosystem that
maintains its normal functioning even through a disturbance.
A. The processes of replenishment of nutrients, dispersion by surrounding plants and
animals, rapid regrowth of plant cover, and succession to a forest can all be
 thought of as resilience mechanisms. Resilience, therefore, helps maintain the
sustainability of ecosystems.
Evolving Ecosystems?
I. All the species in an ecosystem are simultaneously adapting to each other.
II. If relationships within an ecosystem are affected by the removal or introduction of a
dominant organism or by a change in climatic conditions, the entire ecosystem may
develop into one that bears little resemblance to its precursor.