r/K selection theory

A North Atlantic right whale with solitary calf. Whale

reproduction follows a K-selection strategy, with few
offspring, long gestation, long parental care, and a long
period until sexual maturity.
In ecology, r/K selection theory relates to
the selection of combinations of traits in
an organism that trade off between
quantity and quality of offspring. The
focus upon either increased quantity of
offspring at the expense of individual
parental investment of r-strategists, or
reduced quantity of offspring with a
corresponding increased parental
investment of K-strategists, varies widely,
seemingly to promote success in
particular environments.

The terminology of r/K-selection was

coined by the ecologists Robert MacArthur
and E. O. Wilson in 1970[1] based on their
work on island biogeography;[2] although
the concept of the evolution of life history
strategies has a longer history [3] (see e.g.
plant strategies).

The theory was popular in the 1970s and

1980s, when it was used as a heuristic
device, but lost importance in the early
1990s, when it was criticized by several
empirical studies.[4][5] A life-history
paradigm has replaced the r/K selection
paradigm but continues to incorporate
many of its important themes.[6]

Overview
A litter of mice with their mother. The reproduction of
mice follows an r-selection strategy, with many
offspring, short gestation, less parental care, and a
short time until sexual maturity.

In r/K selection theory, selective pressures

are hypothesised to drive evolution in one
of two generalized directions: r- or K-
selection.[1] These terms, r and K, are
drawn from standard ecological algebra as
illustrated in the simplified Verhulst model
of population dynamics:[7]
where N is the population, r is the
maximum growth rate, K is the carrying
capacity of the local environment, and
dN/dt denotes the derivative of N with
respect to time t. Thus, the equation
relates the growth rate of the population N
to the current population size,
incorporating the effect of the two
constant parameters r and K. (Note that
decrease is negative growth.) The choice
of the letter K came from the German
Kapazitätsgrenze (capacity limit), while r
came from rate.
r-selection

As the name implies, r-selected species

are those that emphasize high growth
rates, typically exploit less-crowded
ecological niches, and produce many
offspring, each of which has a relatively
low probability of surviving to adulthood
(i.e., high r, low K).[8] A typical r species is
the dandelion Taraxacum genus.

In unstable or unpredictable environments,

r-selection predominates due to the ability
to reproduce quickly. There is little
advantage in adaptations that permit
successful competition with other
organisms, because the environment is
likely to change again. Among the traits
that are thought to characterize r-selection
are high fecundity, small body size, early
maturity onset, short generation time, and
the ability to disperse offspring widely.

Organisms whose life history is subject to

r-selection are often referred to as r-
strategists or r-selected. Organisms that
exhibit r-selected traits can range from
bacteria and diatoms, to insects and
grasses, to various semelparous
cephalopods and small mammals,
particularly rodents.
K-selection

By contrast, K-selected species display

traits associated with living at densities
close to carrying capacity and typically are
strong competitors in such crowded
niches that invest more heavily in fewer
offspring, each of which has a relatively
high probability of surviving to adulthood
(i.e., low r, high K). In scientific literature, r-
selected species are occasionally referred
to as "opportunistic" whereas K-selected
species are described as "equilibrium".[8]

In stable or predictable environments, K-

selection predominates as the ability to
compete successfully for limited
resources is crucial and populations of K-
selected organisms typically are very
constant in number and close to the
maximum that the environment can bear
(unlike r-selected populations, where
population sizes can change much more
rapidly).

Traits that are thought to be characteristic

of K-selection include large body size, long
life expectancy, and the production of
fewer offspring, which often require
extensive parental care until they mature.
Organisms whose life history is subject to
K-selection are often referred to as K-
strategists or K-selected.[9] Organisms
with K-selected traits include large
organisms such as elephants, humans,
and whales, but also smaller, long-lived
organisms such as Arctic terns.[10]

Continuous spectrum

Although some organisms are identified

as primarily r- or K-strategists, the majority
of organisms do not follow this pattern.
For instance, trees have traits such as
longevity and strong competitiveness that
characterise them as K-strategists. In
reproduction, however, trees typically
produce thousands of offspring and
disperse them widely, traits characteristic
of r-strategists.[11]

Similarly, reptiles such as sea turtles

display both r- and K-traits: although sea
turtles are large organisms with long
lifespans (provided they reach adulthood),
they produce large numbers of unnurtured
offspring.

The r/K dichotomy can be re-expressed as

a continuous spectrum using the
economic concept of discounted future
returns, with r-selection corresponding to
large discount rates and K-selection
corresponding to small discount rates.[12]
Ecological succession
In areas of major ecological disruption or
sterilisation (such as after a major
volcanic eruption, as at Krakatoa or Mount
Saint Helens), r- and K-strategists play
distinct roles in the ecological succession
that regenerates the ecosystem. Because
of their higher reproductive rates and
ecological opportunism, primary
colonisers typically are r-strategists and
they are followed by a succession of
increasingly competitive flora and fauna.
The ability of an environment to increase
energetic content, through photosynthetic
capture of solar energy, increases with the
increase in complex biodiversity as r
species proliferate to reach a peak
possible with K strategies.[13]

Eventually a new equilibrium is

approached (sometimes referred to as a
climax community), with r-strategists
gradually being replaced by K-strategists
which are more competitive and better
adapted to the emerging micro-
environmental characteristics of the
landscape. Traditionally, biodiversity was
considered maximized at this stage, with
introductions of new species resulting in
the replacement and local extinction of
endemic species.[14] However, the
Intermediate Disturbance Hypothesis
posits that intermediate levels of
disturbance in a landscape create patches
at different levels of succession,
promoting coexistence of colonizers and
competitors at the regional scale.

Application
While usually applied at the level of
species, r/K selection theory is also useful
in studying the evolution of ecological and
life history differences between
subspecies, for instance the African honey
bee, A. m. scutellata, and the Italian bee, A.
m. ligustica.[15] At the other end of the
scale, it has also been used to study the
evolutionary ecology of whole groups of
organisms, such as bacteriophages.[16]

Some researchers, such as Lee Ellis, J.

Philippe Rushton, and Aurelio José
Figueredo, have applied r/k selection
theory to various human behaviors,
including crime,[17] sexual promiscuity,
fertility, IQ, and other traits related to life
history theory.[18][19] Rushton's work
resulted in him developing "differential k
theory" to attempt to explain many
variations in human behavior across
geographic areas, a theory which has been
criticized by many other researchers.[19][20]
Other researchers have proposed that the
evolution of human inflammatory
responses is related to r/k selection.[21]

Status
Although r/K selection theory became
widely used during the 1970s,[22][23][24][25] it
also began to attract more critical
attention.[26][27][28][29] In particular, a review
by the ecologist Stephen C. Stearns drew
attention to gaps in the theory, and to
ambiguities in the interpretation of
empirical data for testing it.[30]

In 1981, a review of the r/K selection

literature by Parry demonstrated that there
was no agreement among researchers
using the theory about the definition of r-
and K-selection, which led him to question
whether the assumption of a relation
between reproductive expenditure and
packaging of offspring was justified.[31] A
1982 study by Templeton and Johnson,
showed that in a population of Drosophila
mercatorum under K-selection the
population actually produced a higher
frequency of traits typically associated
with r-selection.[32] Several other studies
contradicting the predictions of r/K
selection theory were also published
between 1977 and 1994.[33][34][35][36]
When Stearns reviewed the status of the
theory in 1992,[37] he noted that from 1977
to 1982 there was an average of 42
references to the theory per year in the
BIOSIS literature search service, but from
1984 to 1989 the average dropped to 16
per year and continued to decline. He
concluded that r/K theory was a once
useful heuristic that no longer serves a
purpose in life history theory.[38]

More recently, the panarchy theories of

adaptive capacity and resilience promoted
by C. S. Holling and Lance Gunderson have
revived interest in the theory, and use it as
a way of integrating social systems,
economics and ecology.[39]

Writing in 2002, Reznick and colleagues

reviewed the controversy regarding r/K
selection theory and concluded that:

The distinguishing feature of the

r- and K-selection paradigm was
the focus on density-dependent
selection as the important agent
of selection on organisms’ life
histories. This paradigm was
challenged as it became clear that
other factors, such as age-specific
mortality, could provide a more
mechanistic causative link
between an environment and an
optimal life history (Wilbur et al.
1974;[26] Stearns 1976,[40]
1977[30]). The r- and K-selection
paradigm was replaced by new
paradigm that focused on age-
specific mortality (Stearns,
1976;[40] Charlesworth, 1980[41]).
This new life-history paradigm
has matured into one that uses
age-structured models as a
framework to incorporate many
 of the themes important to the r–
K paradigm.

— Reznick, Bryant and Bashey,

2002[6]

See also
Evolutionary game theory
Semelparity and iteroparity
Ruderal species
Trivers–Willard hypothesis
Life history theory

References
1. Pianka, E.R. (1970). "On r and K
selection" (PDF). American Naturalist. 104
(940): 592–597. doi:10.1086/282697 .
2. MacArthur, R.; Wilson, E.O. (1967). The
Theory of Island Biogeography (2001 reprint
ed.). Princeton University Press. ISBN 0-
691-08836-5.
3. For example: Margalef, R. (1959). "Mode
of evolution of species in relation to their
places in ecological succession". XVth
International Congress of Zoology.
4. Roff, Derek A. (1993). Evolution Of Life
Histories: Theory and Analysis . Springer.
ISBN 978-0-412-02391-0.
5. Stearns, Stephen C. (1992). The Evolution
of Life Histories . Oxford University Press.
ISBN 978-0-19-857741-6.
6. Reznick, D; Bryant, MJ; Bashey, F (2002).
"r-and K-selection revisited: the role of
population regulation in life-history
evolution" (PDF). Ecology. 83 (6): 1509–
1520. doi:10.1890/0012-
9658(2002)083[1509:RAKSRT]2.0.CO;2 .
7. Verhulst, P.F. (1838). "Notice sur la loi que
la population pursuit dans son
accroissement" . Corresp. Math. Phys. 10:
113–121.
8. For example: Weinbauer, M.G.; Höfle,
M.G. (1 October 1998). "Distribution and
Life Strategies of Two Bacterial Populations
in a Eutrophic Lake" . Appl. Environ.
Microbiol. 64 (10): 3776–3783.
PMC 106546  . PMID 9758799 .
9. "r and K selection" . University of Miami
Department of Biology. Retrieved February
4, 2011.
10. John H. Duffus; Douglas M. Templeton;
Monica Nordberg (2009). Concepts in
Toxicology . Royal Society of Chemistry.
p. 171. ISBN 978-0-85404-157-2.
11. Hrdy, Sarah Blaffer (2000), "Mother
Nature: Maternal Instincts and How They
Shape the Human Species" (Ballantine
Books)
12. Reluga, T.; Medlock, J.; Galvani, A.
(2009). "The discounted reproductive
number for epidemiology". Mathematical
Biosciences and Engineering. 6 (2): 377–
393. doi:10.3934/mbe.2009.6.377 .
13. Gunderson, Lance H.; Holling, C.S.
(2001). Panarchy: Understanding
Transformations In Human And Natural
Systems . Island Press. ISBN 978-1-55963-
857-9.
14. McNeely, J. A. (1994). "Lessons of the
past: Forests and Biodiversity" . Biodiversity
and Conservation. 3: 3–20.
doi:10.1007/BF00115329 .
15. Fewell, Jennifer H.; Susan M. Bertram
(2002). "Evidence for genetic variation in
worker task performance by African and
European honeybees". Behavioral Ecology
and Sociobiology. 52: 318–25.
doi:10.1007/s00265-002-0501-3 .
16. Keen, E. C. (2014). "Tradeoffs in
bacteriophage life histories" .
Bacteriophage. 4 (1): e28365.
doi:10.4161/bact.28365 . PMC 3942329  .
PMID 24616839 .
17. Ellis, Lee (1987-01-01). "Criminal
behavior and r/k selection: An extension of
gene‐based evolutionary theory" . Deviant
Behavior. 8 (2): 149–176.
doi:10.1080/01639625.1987.9967739 .
ISSN 0163-9625 .
18. Figueredo, Aurelio José; Vásquez,
Geneva; Brumbach, Barbara Hagenah;
Schneider, Stephanie M. R. (2007-03-01).
"The K-factor, Covitality, and personality" .
Human Nature. 18 (1): 47–73.
doi:10.1007/bf02820846 . ISSN 1045-
6767 .
19. Weizmann, Fredric; Wiener, Neil I.;
Wiesenthal, David L.; Ziegler, Michael.
"Differential K theory and racial
hierarchies" . Canadian
Psychology/Psychologie canadienne. 31
(1): 1–13. doi:10.1037/h0078934 .
20. Peregrine, P. "Cross-cultural evaluation
of predicted associations between race and
behavior" . Evolution and Human Behavior.
24 (5): 357–364. doi:10.1016/s1090-
5138(03)00040-0 .
21. VAN BODEGOM, D.; MAY, L.; MEIJ, H. J.;
WESTENDORP, R. G. J. "Regulation of
Human Life Histories: The Role of the
Inflammatory Host Response" . Annals of
the New York Academy of Sciences. 1100
(1): 84–97. doi:10.1196/annals.1395.007 .
22. Gadgil, M.; Solbrig, O.T. (1972). "Concept
of r-selection and K-selection — evidence
from wild flowers and some theoretical
consideration". Am. Nat. 106 (947): 14–31.
doi:10.1086/282748 . JSTOR 2459833 .
23. Long, T.; Long, G. (1974). "Effects of r-
selection and K-selection on components of
variance for 2 quantitative traits" . Genetics.
76 (3): 567–573. PMC 1213086  .
PMID 4208860 .
24. Grahame, J. (1977). "Reproductive effort
and r-selection and K-selection in 2 species
of Lacuna (Gastropoda-Prosobranchia)".
Mar. Biol. 40 (3): 217–224.
doi:10.1007/BF00390877 .
25. Luckinbill, L.S. (1978). "r and K selection
in experimental populations of Escherichia
coli". Science. 202 (4373): 1201–1203.
doi:10.1126/science.202.4373.1201 .
PMID 17735406 .
26. Wilbur, H.M.; Tinkle, D.W.; Collins, J.P.
(1974). "Environmental certainty, trophic
level, and resource availability in life history
evolution". American Naturalist. 108 (964):
805–816. doi:10.1086/282956 .
JSTOR 2459610 .
27. Barbault, R. (1987). "Are still r-selection
and K-selection operative concepts?". Acta
Oecologica-Oecologia Generalis. 8: 63–70.
28. Kuno, E. (1991). "Some strange
properties of the logistic equation defined
with r and K – inherent defects or artifacts".
Researches on Population Ecology. 33: 33–
39. doi:10.1007/BF02514572 .
29. Getz, W.M. (1993). "Metaphysiological
and evolutionary dynamics of populations
exploiting constant and interactive
resources – r-K selection revisited".
Evolutionary Ecology. 7 (3): 287–305.
doi:10.1007/BF01237746 .
30. Stearns, S.C. (1977). "Evolution of life-
history traits – critique of theory and a
review of data" (PDF). Annu. Rev. Ecol. Syst.
8: 145–171.
doi:10.1146/annurev.es.08.110177.001045
.
31. Parry, G.D. (March 1981). "The
Meanings of r- and K-selection" . Oecologia.
48 (2): 260–4. doi:10.1007/BF00347974 .
32. Templeton A.R.; Johnson, J.S. (1982).
"Life History Evolution Under Pleiotropy and
K-selection in a Natural Population of
Drosophila mercatorum". In Barker, J.S.F.;
Starmer, W.T. Ecological genetics and
evolution: the cactus-yeast-drosophila
model system . Academic Press. pp. 225–
239. ISBN 978-0-12-078820-0.
33. Snell, Terry W.; King, Charles E.
(December 1977). "Lifespan and Fecundity
Patterns in Rotifers: The Cost of
Reproduction". Evolution. 31 (4): 882–890.
doi:10.2307/2407451 .
34. Taylor, Charles E.; Condra, Cindra
(November 1980). "r- and K-Selection in
Drosophila pseudoobscura". Evolution. 34
(6): 1183–93. doi:10.2307/2408299 .
35. Hollocher, H.; Templeton, A.R. (April
1994). "The molecular through ecological
genetics of abnormal abdomen in
Drosophila mercatorum. VI. The non-
neutrality of the Y chromosome rDNA
polymorphism" . Genetics. 136 (4): 1373–
84. PMC 1205918  . PMID 8013914 .
36. Templeton, A.R.; Hollocher, H.;
Johnston, J.S. (June 1993). "The molecular
through ecological genetics of abnormal
abdomen in Drosophila mercatorum. V.
Female phenotypic expression on natural
genetic backgrounds and in natural
environments" . Genetics. 134 (2): 475–85.
PMC 1205491  . PMID 8325484 .
37. Stearns, S.C. (1992). The Evolution of
Life Histories. Oxford University Press.
ISBN 978-0-19-857741-6.
38. Graves, J. L. (2002). "What a tangled
web he weaves Race, reproductive
strategies and Rushton's life history
theory" . Anthropological Theory. 2 (2): 2
131–154.
doi:10.1177/1469962002002002627 .
39. Gunderson, L. H. and Holling C. S.
(2001) Panarchy: Understanding
Transformations in Human and Natural
Systems Island Press.
ISBN 9781597269391.
40. Stearns, S.C. (1976). "Life history
tactics: a review of the ideas". Quarterly
Review of Biology. 51: 3–47.
doi:10.1086/409052 .
41. Charlesworth, B. (1980). Evolution in
age structured populations. Cambridge, UK:
Cambridge University Press.

Retrieved from
"https://en.wikipedia.org/w/index.php?
title=R/K_selection_theory&oldid=817422726"

Last edited 1 month ago by Kolbert…

Content is available under CC BY-SA 3.0 unless
otherwise noted.