In biology, coevolution occurs when two or more species reciprocally
affect each other's evolution.
Charles Darwin mentioned evolutionary interactions between flowering
plants and insects in
On the Origin of Species
On the Origin of Species (1859). The term
coevolution was coined by
Paul R. Ehrlich
Paul R. Ehrlich and
Peter H. Raven
Peter H. Raven in 1964.
The theoretical underpinnings of coevolution are now well-developed,
and demonstrate that coevolution can play an important role in driving
major evolutionary transitions such as the evolution of sexual
reproduction or shifts in ploidy. More recently, it has also been
demonstrated that coevolution influences the structure and function of
ecological communities as well as the dynamics of infectious
Each party in a coevolutionary relationship exerts selective pressures
on the other, thereby affecting each other's evolution. Coevolution
includes many forms of mutualism, host-parasite, and predator-prey
relationships between species, as well as competition within or
between species. In many cases, the selective pressures drive an
evolutionary arms race between the species involved. Pairwise or
specific coevolution, between exactly two species, is not the only
possibility; in guild or diffuse coevolution, several species may
evolve a trait in reciprocity with a trait in another species, as has
happened between the flowering plants and pollinating insects such as
bees, flies, and beetles.
Coevolution is primarily a biological concept, but researchers have
applied it by analogy to fields such as computer science, sociology,
1.1 Flowering plants
1.1.1 Insects and entomophilous flowers
1.1.2 Birds and ornithophilous flowers
1.2 Fig reproduction and fig wasps
1.3 Acacia ants and acacias
2 Hosts and parasites
2.1 Parasites and sexually reproducing hosts
2.2 Brood parasites
3 Predators and prey
5 Guild or diffuse coevolution
6 Outside biology
6.1 In algorithms
6.2 In architecture
6.3 In technology
6.4 In sociology
7 See also
10 External links
Main article: Mutualism (biology)
Coevolution is the evolution of two or more species which reciprocally
affect each other, sometimes creating a mutualistic relationship
between the species. Such relationships can be of many different
Flowers appeared and diversified relatively suddenly in the fossil
record, creating what
Charles Darwin described as the "abominable
mystery" of how they had evolved so quickly; he considered whether
coevolution could be the explanation. He first mentioned
coevolution as a possibility in On the Origin of Species, and
developed the concept further in Fertilisation of Orchids
Insects and entomophilous flowers
Further information: Entomophily
Honey bee taking a reward of nectar and collecting pollen in its
pollen baskets from white melilot flowers
Modern insect-pollinated (entomophilous) flowers are conspicuously
coadapted with insects to ensure pollination and in return to reward
the pollinators with nectar and pollen. The two groups have coevolved
for over 100 million years, creating a complex network of
interactions. Either they evolved together, or at some later stages
they came together, likely with pre-adaptations, and became mutually
adapted. The term coevolution was coined by Paul R. Ehrlich
Peter H. Raven
Peter H. Raven in 1964, to describe the evolutionary interactions
of plants and butterflies.
Several highly successful insect groups—especially the Hymenoptera
(wasps, bees and ants) and
Lepidoptera (butterflies) as well as many
Diptera (flies) and
Coleoptera (beetles)—evolved in
conjunction with flowering plants during the
Cretaceous (145 to 66
million years ago). The earliest bees, important pollinators today,
appeared in the early Cretaceous. A group of wasps sister to the
bees evolved at the same time as flowering plants, as did the
Lepidoptera. Further, all the major clades of bees first appeared
between the middle and late Cretaceous, simultaneously with the
adaptive radiation of the eudicots (three quarters of all
angiosperms), and at the time when the angiosperms became the world's
dominant plants on land.
At least three aspects of flowers appear to have coevolved between
flowering plants and insects, because they involve communication
between these organisms. Firstly, flowers communicate with their
pollinators by scent; insects use this scent to determine how far away
a flower is, to approach it, and to identify where to land and finally
to feed. Secondly, flowers attract insects with patterns of stripes
leading to the rewards of nectar and pollen, and colours such as blue
and ultraviolet, to which their eyes are sensitive; in contrast,
bird-pollinated flowers tend to be red or orange. Thirdly, flowers
such as some orchids mimic females of particular insects, deceiving
males into pseudocopulation.
Yucca whipplei, is pollinated exclusively by Tegeticula
maculata, a yucca moth that depends on the yucca for survival. The
moth eats the seeds of the plant, while gathering pollen. The pollen
has evolved to become very sticky, and remains on the mouth parts when
the moth moves to the next flower. The yucca provides a place for the
moth to lay its eggs, deep within the flower away from potential
Birds and ornithophilous flowers
Further information: Ornithophily
Purple-throated carib feeding from and pollinating a flower
Hummingbirds and ornithophilous (bird-pollinated) flowers have evolved
a mutualistic relationship. The flowers have nectar suited to the
birds' diet, their color suits the birds' vision and their shape fits
that of the birds' bills. The blooming times of the flowers have also
been found to coincide with hummingbirds' breeding seasons. The floral
characteristics of ornithophilous plants vary greatly among each other
compared to closely related insect-pollinated species. These flowers
also tend to be more ornate, complex, and showy than their insect
pollinated counterparts. It is generally agreed that plants formed
coevolutionary relationships with insects first, and ornithophilous
species diverged at a later time. There is not much scientific support
for instances of the reverse of this divergence: from ornithophily to
insect pollination. The diversity in floral phenotype in
ornithophilous species, and the relative consistency observed in
bee-pollinated species can be attributed to the direction of the shift
in pollinator preference.
Flowers have converged to take advantage of similar birds. Flowers
compete for pollinators, and adaptations reduce unfavourable effects
of this competition. The fact that birds can fly during inclement
weather makes them more efficient pollinators where bees and other
insects would be inactive.
Ornithophily may have arisen for this
reason in isolated environments with poor insect colonization or areas
with plants which flower in the winter. Bird-pollinated
flowers usually have higher volumes of nectar and higher sugar
production than those pollinated by insects. This meets the birds'
high energy requirements, the most important determinants of flower
choice. In Mimulus, an increase in red pigment in petals and
flower nectar volume noticeably reduces the proportion of pollination
by bees as opposed to hummingbirds; while greater flower surface area
increases bee pollination. Therefore, red pigments in the flowers of
Mimulus cardinalis may function primarily to discourage bee
visitation. In Penstemon, flower traits that discourage bee
pollination may be more influential on the flowers' evolutionary
change than 'pro-bird' adaptations, but adaptation 'towards' birds and
'away' from bees can happen simultaneously. However, some flowers
Heliconia angusta appear not to be as specifically
ornithophilous as had been supposed: the species is occasionally (151
visits in 120 hours of observation) visited by
Trigona stingless bees.
These bees are largely pollen robbers in this case, but may also serve
Following their respective breeding seasons, several species of
hummingbirds occur at the same locations in North America, and several
hummingbird flowers bloom simultaneously in these habitats. These
flowers have converged to a common morphology and color because these
are effective at attracting the birds. Different lengths and
curvatures of the corolla tubes can affect the efficiency of
extraction in hummingbird species in relation to differences in bill
morphology. Tubular flowers force a bird to orient its bill in a
particular way when probing the flower, especially when the bill and
corolla are both curved. This allows the plant to place pollen on a
certain part of the bird's body, permitting a variety of morphological
A fig exposing its many tiny matured, seed-bearing gynoecia. These are
pollinated by the fig wasp, Blastophaga psenes. In the cultivated fig,
there are also asexual varieties.
Ornithophilous flowers need to be conspicuous to birds. Birds have
their greatest spectral sensitivity and finest hue discrimination at
the red end of the visual spectrum, so red is particularly
conspicuous to them. Hummingbirds may also be able to see ultraviolet
"colors". The prevalence of ultraviolet patterns and nectar guides in
nectar-poor entomophilous (insect-pollinated) flowers warns the bird
to avoid these flowers. Each of the two subfamilies of
Phaethornithinae (hermits) and the Trochilinae, has
evolved in conjunction with a particular set of flowers. Most
Phaethornithinae species are associated with large monocotyledonous
herbs, while the
Trochilinae prefer dicotyledonous plant species.
Fig reproduction and fig wasps
Main article: Reproductive coevolution in Ficus
Ficus genus is composed of 800 species of vines, shrubs, and
trees, including the cultivated fig, defined by their syconiums, the
fruit-like vessels that either hold female flowers or pollen on the
inside. Each fig species has its own fig wasp which (in most cases)
pollinates the fig, so a tight mutual dependence has evolved and
persisted throughout the genus.
Acacia ants and acacias
Pseudomyrmex ant on bull thorn acacia (
Vachellia cornigera) with
Beltian bodies that provide the ants with protein
Main article: Pseudomyrmex ferruginea
The acacia ant (Pseudomyrmex ferruginea) is an obligate plant ant that
protects at least five species of "Acacia" (Vachellia)[a] from preying
insects and from other plants competing for sunlight, and the tree
provides nourishment and shelter for the ant and its larvae.
Such mutualism is not automatic: other ant species exploit trees
without reciprocating, following different evolutionary strategies.
These cheater ants impose important host costs via damage to tree
reproductive organs, though their net effect on host fitness is not
necessarily negative and, thus, becomes difficult to forecast.
Hosts and parasites
Main article: Host–parasite coevolution
Parasites and sexually reproducing hosts
Host–parasite coevolution is the coevolution of a host and a
parasite. A general characterization of many viruses, obligate
parasites, is that they coevolved alongside their respective hosts.
Correlated mutations between the two species enter them into an
evolution arms race. Whichever organism, host or parasite, that cannot
keep up with the other will be eliminated from their habitat, as the
species with the higher average population fitness survives. This race
is known as the Red Queen hypothesis. The Red Queen hypothesis
predicts that sexual reproduction allows a host to stay just ahead of
its parasite, similar to the
Red Queen's race
Red Queen's race in Through the
Looking-Glass: "it takes all the running you can do, to keep in the
same place". The host reproduces sexually, producing some
offspring with immunity over its parasite, which then evolves in
The parasite/host relationship probably drove the prevalence of sexual
reproduction over the more efficient asexual reproduction. It seems
that when a parasite infects a host, sexual reproduction affords a
better chance of developing resistance (through variation in the next
generation), giving sexual reproduction variability for fitness not
seen in the asexual reproduction, which produces another generation of
the organism susceptible to infection by the same
Coevolution between host and parasite may
accordingly be responsible for much of the genetic diversity seen in
normal populations, including blood-plasma polymorphism, protein
polymorphism, and histocompatibility systems.
Eurasian reed warbler
Eurasian reed warbler raising a common cuckoo
Main article: Brood parasitism
Brood parasitism demonstrates close coevolution of host and parasite,
for example in cuckoos. These birds do not make their own nests, but
lay their eggs in nests of other species, ejecting or killing the eggs
and young of the host and thus having a strong negative impact on the
host's reproductive fitness. Their eggs are camouflaged as eggs of
their hosts, implying that hosts can distinguish their own eggs from
those of intruders and are in an evolutionary arms race with the
cuckoo between camouflage and recognition.
Cuckoos are counter-adapted
to host defences with features such as thickened eggshells, shorter
incubation (so their young hatch first), and flat backs adapted to
lift eggs out of the nest.
Predators and prey
Predator and prey: a leopard killing a bushbuck
Main article: Predation
Predators and prey interact and coevolve, the predator to catch the
prey more effectively, the prey to escape. The coevolution of the two
mutually imposes selective pressures. These often lead to an
evolutionary arms race between prey and predator, resulting in
The same applies to herbivores, animals that eat plants, and the
plants that they eat. In the Rocky Mountains, red squirrels and
crossbills (seed-eating birds) compete for seeds of the lodgepole
pine. The squirrels get at pine seeds by gnawing through the cone
scales, whereas the crossbills get at the seeds by extracting them
with their unusual crossed mandibles. In areas where there are
squirrels, the lodgepole's cones are heavier, and have fewer seeds and
thinner scales, making it more difficult for squirrels to get at the
seeds. Conversely, where there are crossbills but no squirrels, the
cones are lighter in construction, but have thicker scales, making it
more difficult for crossbills to get at the seeds. The lodgepole's
cones are in an evolutionary arms race with the two kinds of
Sexual conflict has been studied in
Drosophila melanogaster (shown
mating, male on right).
Intraspecific competition and Interspecific competition
Both intraspecific competition, with features such as sexual
conflict and sexual selection, and interspecific competition,
such as between predators, may be able to drive coevolution.
Guild or diffuse coevolution
Long-tongued bees and long-tubed flowers coevolved, whether pairwise
or "diffusely" in groups known as guilds.
The types of coevolution listed so far have been described as if they
operated pairwise (also called specific coevolution), in which traits
of one species have evolved in direct response to traits of a second
species, and vice versa. This is not always the case. Another
evolutionary mode arises where evolution is still reciprocal, but is
among a group of species rather than exactly two. This is called guild
or diffuse coevolution. For instance, a trait in several species of
flowering plant, such as offering its nectar at the end of a long
tube, can coevolve with a trait in one or several species of
pollinating insects, such as a long proboscis. More generally,
flowering plants are pollinated by insects from different families
including bees, flies, and beetles, all of which form a broad guild of
pollinators which respond to the nectar or pollen produced by
Coevolution is primarily a biological concept, but has been applied to
other fields by analogy.
See also: Evolutionary computation
Coevolutionary algorithms are used for generating artificial life as
well as for optimization, game learning and machine
Daniel Hillis added "co-evolving
parasites" to prevent an optimization procedure from becoming stuck at
Karl Sims coevolved virtual creatures.
The concept of coevolution was introduced in architecture by the
Henrik Valeur as an antithesis to the
concept of "star-architecture". As the curator of the Danish
Pavilion at the 2006 Venice Biennale of Architecture he conceived and
orchestrated an exhibition project named 'Co-evolution', awarded the
Golden Lion for Best National Pavilion.
The exhibition included urban planning projects for the cities of
Shanghai and Xi'an, which had been developed in
collaboration between young professional Danish architects and
students and professors and students from leading universities in the
four Chinese cities. By creating a framework for collaboration
between academics and professionals representing two distinct
cultures, it was hoped that the exchange of knowledge, ideas and
experiences would stimulate "creativity and imagination to set the
spark for new visions for sustainable urban development." Valeur
later argued that: "As we become more and more interconnected and
interdependent, human development is no longer a matter of the
evolution of individual groups of people but rather a matter of the
co-evolution of all people."
Further information: Software ecosystem
Computer software and hardware can be considered as two separate
components but tied intrinsically by coevolution. Similarly, operating
systems and computer applications, web browsers and web applications.
All of these systems depend upon each other and advance step by step
through a kind of evolutionary process. Changes in hardware, an
operating system or web browser may introduce new features that are
then incorporated into the corresponding applications running
alongside. The idea is closely related to the concept of "joint
optimization" in sociotechnical systems analysis and design, where a
system is understood to consist of both a "technical system"
encompassing the tools and hardware used for production and
maintenance, and a "social system" of relationships and procedures
through which the technology is tied into the goals of the system and
all the other human and organizational relationships within and
outside the system. Such systems work best when the technical and
social systems are deliberately developed together.
Models of coevolution have been proposed for sociology and
international political economy. Richard Norgaard's 2006 book
Development Betrayed proposes a "Co-Evolutionary Revisioning of the
Future" of social and economic life.
Escape and Radiate Coevolution
Genomics of domestication
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Stephen C. Stearns (Open Yale
Evolutionarily stable strategy
Darwinian literary studies
Evolution of emotion
W. D. Hamilton
Carel van Schaik
David Sloan Wilson
E. O. Wilson
George C. Williams
Jerome H. Barkow
Dominic D. P. Johnson
Justin L. Barrett
David F. Bjorklund
David C. Geary
Judith Rich Harris
Aurelio José Figueredo
Douglas T. Kenrick
Simon M. Kirby
Michael T. McGuire
Randolph M. Nesse
David P. Schmitt
Todd K. Shackelford
Peter K. Smith
Mark van Vugt
Center for Evolutionary Psychology
Human Behavior and
Max Planck Institute for Human Cognitive and Brain Sciences
New England Complex Systems Institute
The Adapted Mind
Evolution of Human Sexuality
Evolution and Human Behavior
Evolutionary psychology and culture
Criticism of evolutionary psychology
Evolutionary psychology research groups and centers
Bibliography of evolution and human behavior