r-K SELECTION: THE DEVELOPMENT-REPRODUCTION

TRADE-OFF

The extensive interest in life-history evolution stems from the

predictions made first by R.H. MacArthur that the demographic

or life history characteristics of a population may vary depending

upon the type of environment it experienced.

For example, in a constant environment limited by resources, the

population may exist at or near the carrying capacity, K.

In other environments, in which conditions change in an

unpredictable manner, a population may seldom reach the carrying

capacity and is generally in a high-population-growth phase.

If we assume that these are two extremes of a continuum of

environmental conditions found in nature, then we can say that

populations in these two situations are subject to high-density

conditions, or K-selection, and low density conditions, or r-

selection, respectively (MacArthur, 1962).

In order to evaluate the evolutionary consequences of

environmental patterns on life-history characteristics, Pianka

(1970) categorized in a general way the population characteristics

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and the traits that selection would favor in the two types of

environments (Table 1). For example, he assumed that in an

unpredictable environment the population density would be low

and mortality type III (high early mortality), so that selection

would favor those individuals with rapid development (early

reproductive maturity) and high fecundity because these changes

would enable a population to recover quickly after an episode of

high mortality.

On the other hand, in a stable environment, the population

density would be near the carrying capacity and adult mortality

concentrated in the older age classes, so selection would favor

delayed reproduction and generally a more efficient use of

resources to promote a longer life.

Returning to the concept of energy allocation to reproduction

versus growth or maintenance, for populations in unpredictable

environments more energy is used for reproduction, increasing r,

while for populations in stable environments more energy is used

for growth and maintenance (survival), or increasing K .

In order to maximize fitness in a predictable environment, it pays

to invest resources in long-term development and long life (K

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selection); in a risky environment, it is better to produce as much

offspring as quickly as possible (r selection).

In the effort to attain generalities concerning life-history

evolution, these ideas have been overextended to classify species

as r-selected or K-selected, depending upon the population

characteristics of these species.

In addition, it has been suggested that whole groups of

organisms, such as insects or annual plants, can be categorized as

r-selected and that other groups, such as terrestrial vertebrates

or perennial plants, are K-selected.

Characteristics of r and K selection

Of course, the animal or plant is not thinking: "How do I change

my characteristics?" Natural selection is the force for change,

not the individual's conscious decision. But, natural selection has

produced a gradation of strategies, with extreme r-selection at

one end of the spectrum and extreme K-selection at the other

end.

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Table 1: Comparison of some characteristics of organisms which
are extreme r or K strategists:
r K
Unstable environment, density
independent
Environment: Variable and or
unpredictable
small size of organism
energy used to make each individual
is low
many offspring are produced
early maturity
short life expectancy
each individual reproduces only once
type III survivorship pattern
in which most of the individuals die
within a short time
but a few live much longer
Stable environment, density
dependent interactions
Environment: Constant and or
predictable
large size of organism
energy used to make each individual
is high
few offspring are produced
late maturity, often after a
prolonged period of parental care
long life expectancy
individuals can reproduce more than
once in their lifetime
type I or II survivorship pattern
in which most individuals live to near
the maximum life span
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