Extinction Risk: Theoretical Considerations

I. Introduction
A. Changes in the size of a wildlife population over time can have a wide range of
consequences depending on the species, the location, and many other factors.
1. Growth of white-tailed deer (Odocoileus virginianus) populations over the past 15
years has caused changes in native vegetation structure and composition in many
places, particularly in the upper Midwest.
2. On the other hand, the number of Florida panthers (Puma concolor) has declined
to the point that many biologists are skeptical that the species can avoid
extinction.
B. Here we deal with the body of theory that has been developed to account for why and
how populations become extinct. Most of that theory deals with extinction as a
consequence of low numbers, the various difficulties that a population can get into
when it is too small.
II. Demographic problems contribute to the risk of extinction
A. All populations experience some variation in their numbers over time, and some of
this variation is predicable.
1. If a population has a distinct breeding season (i.e., they rut in the fall and calve in
the spring), each year after the females give birth to their offspring, the population
will grow in size.
2. If there is a season (say, a tough winter), that is particularly hard on a population,
the population abundance will decline.
B. However, we cannot forecast all the variation in abundance.
1. Random, unpredictable variation in population abundance is called stochasticity.
2. Two types of random variation a population may experience are demographic and
environmental stochasticity.
a. Demographic stochasticity.
1) Demographic stochasticity is random variation in things like sex ratios,
birth rates, and death rates.
2) Wildlife populations are comprised of individuals that are not exactly alike.
3) As a result, often more or fewer individuals die or are born than would be
expected. For example, if the female black bears (Ursus americanus) in a
population with demographic stochasticity produce a mean of 2 offspring
per year, some females in the population may have no offspring, some may
have 1 offspring, and some may have 2, 3, or even 4 offspring. This
variation is random from year-to-year.
4) Demographic stochasticity occurs within a population even if the
environment does not vary. Here is an example of demographic
stochasticity in a group of mountain gorillas (Gorilla beringei).
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a) Female gorillas breed every 3.5-4.5 years; therefore, the mean

fecundity rate = 0.25, which means you can expect 1 female mountain
gorilla to give birth to 1 offspring every 4 years.
b) If 3 of 4 females in a group give birth during the same year, and the
4th female gives birth the following year, there may be 3 years before
the next offspring is born.
c) If the mean fecundity rate is 0.25 and there are 4 females in the
group, we would expect 1 offspring per year. However, as we have
already seen, we can have multiple births in 1 year followed by several
years with no births.
d) Likewise, several or no individuals may die during the same year due
to chance.
e) Having more infants born or more individuals die than expected will
change the mean growth rate of the population. If a population is
very small, demographic variation can have very large consequences;
namely, if the population is small and the death rate is high, the
population could go extinct.
b. Environmental stochasticity.
1) Unpredictable variation in the environment, or environmental
stochasticity, also creates variation in population growth rate over time.
2) Although some environmental variation is predictable, like the changing
seasons, variation between different years tends to be less predictable.
3) Rainfall, temperature, snow pack amounts and conditions, frequency and
intensity of storms, humidity, and amount of cloud cover can all vary
between years, and these weather conditions directly affect the amount of
food available each year.
4) During bad years, birth rates may drop and death rates may increase. Taken
together, low birth rates and high death rates can substantially decrease the
population size.
3. In a large population with a wide distribution, stochastic effects generally are not
too harmful. However, if a population is very small, and/or is found only in a
small area, demographic and environmental stochasticity combined or alone can
drive a population to the brink of extinction. Here is the idea in different terms.
a. A large population is usually able to recover from bad years or catastrophes
because some individuals usually survive to reproduce in future years. For
example, if 90% of a population of 3,000 animals is killed during a
catastrophe, say a wildfire, 300 individuals would remain to continue to
reproduce.
b. However, if the same catastrophe kills 90% of a population of 30 individuals,
only 3 individuals would remain. If 2 are male and 1 is an old non-reproducing
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female, the population will become extinct. This is what happened to the
heath hen on Martha’s Vineyard in the early 20th century.
III. Genetic problems also contribute to the risk of extinction
A. Another problem faced by small populations is the loss of genetic variation at both
the individual and population level.
1. Individuals have genetic variation.
a. All diploid organisms are defined as having 2 copies of each gene. Each copy is
called an allele.
b. One copy is inherited from each parent.
c. Although each allele contains information about the same trait, they may not
be identical.
d. When an individual has corresponding alleles that are different, this individual
possesses genetic variation for that trait.
2. Populations also possess genetic variation.
a. This variation occurs when individuals in the same population have differences
in their genetic makeup.
b. A population with little genetic variation is less likely to survive environmental
change because all individuals may be equally susceptible. For example,
geneticists fear that the lack of genetic variation in cheetah (Acinonyx jubatus)
populations makes them more vulnerable to introduced diseases such as feline
leukemia.
3. Genetic variation is maintained primarily by mutation and by gene flow from
other localities in which different genes have a selective advantage.
B. Influences of population size on genetic variation.
1. In small populations, chance events can significantly alter allele frequencies. Such
alteration is called genetic drift.
a. If only a few individuals are drawn at random to form the next generation, the
alleles they carry are not likely to be in the same proportions as alleles in the
gene pool from which they were drawn.
b. Two important causes of genetic drift are bottlenecks and founder effects.
1) Bottlenecks.
a) Even organisms that normally have large populations may pass through
occasional periods when only a small number of individuals survive.
b) During these population bottlenecks, genetic variation can be lost by
chance.
c) Populations in nature pass through bottlenecks for many different
reasons. For example, predators may reduce populations of their prey
to very small sizes.
2) Founder effects.
a) When a species expands into new regions, a small number of pioneering
individuals may start populations.
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b) These pioneers are not likely to have all the alleles found in their source
population. Even if they do, the allele frequencies are likely to differ
from those in the source population.
c) The situation is equivalent to that for a large population reduced by a
bottleneck, but rather than a small surviving population, there is a small
founding population.
d) This type of genetic drift is called a founder effect.
e) Examples follow.
• Because many plant species reproduce sexually by self-fertilization, a
single seed—an extreme example of a founder effect, may start a
new plant population.
• Population geneticists studied the genetic composition of a
founding population when Drosophila subobscura, a well-studied
European species of fruit fly, was discovered near Puerto Montt,
Chile, in 1978 and at Port Townsend, Washington, in 1982.
¾ The founding insects probably reached Chile and the U.S. from
Europe aboard the same ship, because both populations are
genetically very similar.
¾ As expected from a small founding population, only a small part
of the total genetic variability found in Europe reached the
Americas.
¾ Geneticists estimate that a least 10, but no more than 100, flies
initially arrived in the New World.
2. Small populations lose genetic variation over time by inbreeding.
a. If a population is small enough, there are not many mates available from which
to choose. Eventually, individuals mate with others that are closely related.
b. The more closely related individuals are, the more genes they share.
c. If individuals with the same genes mate, the resulting offspring may inherit 2
identical alleles.
d. In very inbred populations, many individuals have identical copies of harmful
genes, often resulting in reduced fertility and unhealthy offspring. For
example, evidence has been found of reduced sperm fertility of inbred lions
and cheetahs.
e. The problems associated with low genetic diversity due to inbreeding are called
inbreeding depression.
3. The extinction risk of a small population may be affected by the genetic health of
the population.
a. From a management perspective, to protect the long-term viability of a
population, we need to ensure that there are enough individuals to protect it
from the loss of genetic variation.
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b. If a population is below the threshold necessary to maintain genetic variation,

the fitness of the individuals in the already small population may decrease.
c. With lower fecundities and/or higher mortalities, it will be increasingly
difficult for the small population to recover.
d. To maximize the amount of variation maintained in a population, zoos and
conservation organizations have organized gene banks.
IV. Allee effects and extinction vortexes
A. Another factor that may push populations toward extinction is the Allee effect.
1. At very low population densities, individuals may not have enough contact with
other individuals of the population.
2. It can be difficult for individuals to find mating partners, causing the reproductive
rate and the population growth rate to drop.
B. Once a population becomes too small to sustain itself, it may enter an extinction
vortex.
1. An extinction vortex is very much like a whirlpool or a black hole; once you begin
to be pulled toward its center, it becomes harder and harder to escape.
2. As populations fall below a critical threshold, it becomes too sensitive to
environmental and demographic stochasticity to maintain stable numbers.
3. Populations in this downward spiral cannot change its course without outside
interference and management. (An extinction vortex is a little like hypothermia.
Unless a person suffering from hypothermia receives heat from an external source,
their body temperature will continue to drop until they die.)
C. How large must a population be to sustain itself and avoid the problems faced by
small populations?
1. This is a difficult question with no clear answers but probably varies somewhat for
each species.
2. Some biologists speculate that populations should have at least 1,000 -10,000
individuals to persist long term; others estimate that populations should really
have at least 100,000 individuals or more to persist indefinitely.
V. Population viability analysis
A. One way to determine the long-term sustainability of a population is to perform a
population viability analysis (PVA).
1. A PVA is a model used to predict the probability of a population declining or
becoming extinct.
2. If a population is not likely to survive in its present condition, PVA can be used to
determine which aspects of the population (i.e., fecundity, mortality, or habitat
quality) will be the most sensitive to conservation efforts.
B. Effective population size.
1. Small populations may be in jeopardy of losing genetic variation via genetic drift,
and with it, their ability to respond to environmental change.
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2. Thus, biologists are keenly interested in the question of how large a population
must be in order to stave off the loss of genetic diversity.
3. This question is difficult to address, however, because not every member of a
population contributes to the genetic structure of the population in the same way.
a. Mating systems may prevent some individuals from mating with others, and
there may be age-specific differences in reproductive output.
b. Because of this, simple estimates of population density do not provide enough
information to evaluate the vulnerability of the population to genetic loss.
c. One way to overcome this difficulty is to calculate the genetic effective
population size, denoted Ne.
1) The EPS is the size of an idealized population that has the same amount of
genetic drift as the natural population under study.
2) An idealized population is one in which the number of reproducing males
and females is the same (1:1 sex ratio), random mating occurs among males
and females (that is, every male has an equal probability of mating with
every female), the rate of emigration and immigration is constant, and in
which there is no age structure. Usually Ne < N, though this is not always
the case.
3) The EPS depends on the relative reproductive success of males and females
and the variance in that success.
4) Thus, to determine Ne, we calculate separate effective population sizes for
males and females as follows:

NmK m − 1
Nmales =
⎛ K m + Vm ⎞
⎜ ⎟−1
⎜ K ⎟
⎝ m ⎠

Nf K f − 1
Nfemales = ,
⎛ K f + Vf ⎞
⎜ ⎟−1
⎜ K ⎟
⎝ f ⎠

where:

Nm and Nf = the numbers of breeding males and females, respectively,

Km and Kf = the average numbers of offspring produced by males and
females, respectively, in their lifetime, and
Vm and Vf = the variance in the number of offspring produced by males
and females, respectively.
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a) Inspection of these equations reveals how the variance in the number of

offspring produced influences the EPS of males and females.
b) For a given number of breeding males, Nm, producing an average
number of offspring Km, the EPS Nmales will decrease as the variance in
offspring Vm decreases. For example, for a population with Nm = 10,
and average number of offspring Km = 2, the effective male population
size, Nmales, will be 7.6 if Vm = 1, but 4.7 if Vm =2.
5) The separate values for males and females given in the equations above may
be combined in the following way to calculate the overall EPS:

⎛ 1 1 ⎞⎟
N = 4⎜ + - 1.
e ⎜N ⎟
⎝ males Nfemales ⎠

Clearly, the number, reproductive output, and variance of reproductive

output of males and females interact to determine the EPS of a population.

4. How large should the EPS be in order to maintain genetic diversity in the face of
genetic drift? It has been suggested that an effective population size of at least
500 is required, though it is likely that the minimum number will vary widely
among populations having sex ratios, age structures, and social arrangements.
5. Related to the concept of EPS is that of minimum viable population size (MVP).
a. The MVP is the smallest population that can persist for a specified time, usually
taken to be 1000 years.
b. The MVP is dependent both on the demographics of the population and the
genetic diversity within the population.
VI. How to prevent extinction
A. Of the 12 examples of extinction or steep decline mentioned in chapter 15 of the text,
probably the most important is modification or destruction of habitat.
B. The first step in averting extinction is to recognize the problem.
C. The second step is to discover how the population got into its present mess.
1. Is the cause of decline a single factor or a combination of factors?
2. Are those factors still operating?
3. If so can they be nullified?
D. Once a decline in a species is recognized and the causes are determined, the third step
is to treat the problem.
1. Sometimes all it takes is a legislative change such as a ban on hunting (as with the
Canadian muskoxen).
2. More usually, active management (e.g., predator control and captive breeding for
the Lord Howe Island woodhen) is necessary.
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3. The management actions needed to reverse the fortunes of a declining species are
seldom more than conventional management techniques unless a species is in
desperate straits.
4. Ex-situ techniques preserve and amplify a population of an endangered species
outside its natural habitat. Thereafter it can be reintroduced. The Lord Howe
Island woodhen and the Arabian oryx are good examples.