First some definitions: coevolution is a change in the genetic composition
of one species (or group) in response to a genetic change in another. More
generally, the idea of some reciprocal evolutionary change in interacting
species is a strict definition of coevolution.
At first glance (or thought), it might seem that everything is
involved in coevolution. This assumption might stem from the fact that
virtually all organisms interact with other organisms and presumably influence
their evolution in some way. But this assumption depends entirely on ones
definition of the term Coevolution.
The term is usually attributed to Ehrlich and Raven's study of butterflies
on plants (1964) but the term was used by others prior to 1964 and the
idea was very present in the Origin of Species. Ehrlich and Raven documented
the association between species of butterflies and their host
plants noting that plants' secondary compounds (noxious compounds
produced by the plant) determined the usage of certain plants by butterflies.
The implication was that the diversity of plants and their "poisonous"
secondary compounds contributed to the generation of diversity of butterfly
Here we have a very general observation of one group of organisms having
an influence on another group of organisms. Is this coevolution? Some would
argue that it is not good evidence for coevolution because the reciprocal
changes have not been documented clearly. Like the issue of defining
an adaptation, we should not invoke coevolution without reasonable evidence
that the traits in each species were a result of or evolved from
the interaction between the two species.
Lets consider plants and insects: there is little evidence to
determine whether plants' secondary compounds arose for the purpose
of preventing herbivores from eating plant tissue. Certain plants may have
produced certain compounds as waste products and herbivores attacked those
plants that they could digest. Parasites and hosts: when a parasite
invades a host, it will successfully invade those hosts whose defense traits
it can circumvent because of the abilities it caries at that time.
Thus presence of a parasite on a host does not constitute
evidence for coevolution. These criticisms are quite distinct from
the opportunity for coevolution once a parasite has established
itself on a host.
The main point is that any old interaction, symbiosis, mutualism,
etc. is not synonymous with coevolution. In one sense there
has definitely been "evolution together" but whether this
fits our strict definition of coevolution needs to be determined by careful
1) observation, 2) experimentation and 3) phylogenetic
The classic analogy is the coevolutionary arms race: a plant
has chemical defenses, an insect evolves the biochemistry to detoxify these
compounds, the plant in turn evolves new defenses that the insect in turn
"needs" to further detoxify. At present the evidence for these
types of reciprocal adaptations is limited, but the suggestive
evidence of plant animal interactions is widespread. An important
point is the relative timing of the evolution of the various traits
that appear to be part of the coevolution. If the presumed reciprocally
induced, sequential traits actually evolved in the plant (host) before
the insect (parasite) became associated with it, we should not call it
coevolution. See different example figs. 22.6-22.7, pgs. 621-622 + text.
There are a variety of different modes of coevolution. In some cases
coevolution is quite specific such as those between two cellular functions.
The endosymbiont theory proposes that current day mitochondria and chloroplasts
were once free-living unicellular individuals. These cells entered the
cytoplasm of other cells, an example of the general phenomenon of endosymbiosis.
Current-day mitochondrial and chloroplast genomes are much smaller than
the genome sizes of their presumed free-living ancestors. Some of this
reduction in genome size is due to the transfer of genes from organelle
genomes to the nuclear genome. Thus, being in the cellular environment
has influenced the evolution of organelle genomes. There is evidence that
the faster rate of evolution of animal mitochondrial DNA has accelerated
the rate of evolution of some of the nuclear genes that function in the
mitochondria. Thus there is some evidence for reciprocal phenomena
Other modes of coevolution involve competitive interaction between two
specific species. The Plethodon salamander study is a good example:
two species are competing: in the Great Smoky mountains the two species
compete strongly as evidenced by the fact that each species will
increase population size if the other is removed. Here there is
a clear reciprocal interaction between the two populations (species), each
affecting the other.
[The role of competition between species, the coevolutionary responses
to this competition and the consequences for the evolution of communities
is illustrated in the Anolis lizard fauna of the Caribbean. There
is coevolution because the competitive interactions between resident and
invading species of Anolis involve reciprocal responses in the evolution
of body size. These affect the structure of the lizard community as evidenced
by the general pattern of there being a single species of lizard on each
Character displacement also provides and example of a pattern
we might interpret as the result of coevolution. Mud snails show pattern
of character displacement in sympatry due presumably to competition for
food items (don't confuse this with reinforcement; the selective agent
here is not reduced hybrid fitness). We might call this co
evolution because both species show a shift when compared to allopatric
samples of each species (mean of both ~ 3.2 in allopatry vs. ~ 4.0 and
~ 2.8 in sympatry). If only one species exhibited character displacement
and you were a really picky evolutionist you might not be convinced of
a reciprocal response.
Another strong case is the Ant - Acacia mutualism. Here specific
traits in each species appear to have evolved in response to the interaction.
The ant (Pseudomyrmex species) depends on the Acacia plant
for food and housing; acacia depends on ant for protection
from potential herbivores (species that eat plant tissue). Specific characters
of the plant appear to have evolved for the maintenance of this
mutualism: 1) swollen, ~ hollow thorns (= ant home), 2) extra-floral
nectaries (source of nectar outside the flower [i.e., the usual
location] providing ants with food), 3) leaflet tips = Beltian bodies (=
99% of solid food for larval/adult ants). Specific characters in the ant
that have evolved for the maintenance of this mutualism:
1) defense against herbivores 2) removal of fungal spores from Beltian
body break-point (prevents fungal pathogens from invading plant tissues).
The main point is that there are traits in both the ant and the
acacia that are traits not normally found in close relatives of each that
are not involved in similar mutualisms: mutualistic traits have
evolved for the interaction in reciprocal fashion. See another example
: fig. 22.1 & table 22.1, pg. 611.
Coevolution may be considered among broad groups of taxa, so called
diffuse coevolution (such as the general coevolution between plants
and insects [assuming it is real]). A nice idea, but in fact the real action
must be going on between pairs of species from each group. It is true that
the Pierid butterflies (family Pieridae) are associated with the plant
family Cruciferae, so there may be something general about each taxon that
allows the coevolution to proceed. But the true reciprocal events must
be mediated at the host species-insect species level.
Mimicry presents a context were coevolutionary phenomena should
be evident. Generally, we would expect that Mullerian mimicry would be
more likely to exhibit reciprocal evolutionary patterns since both
species involved are unpalatable and therefore have an opportunity to affect
the evolution of each other's color patters. This does not mean
that Batesian mimicry (one unpalatable model) will not involve coevolutionary
phenomena, but the evolution of warning coloration is certainly going to
be more asymmetrical since the palatable species will show a greater
response to the state of the model than will the model show to the evolving
state of the mimic.
The Mullerian mimics Heliconius erato and H. melpomene.
illustrate both the frequency dependent nature of mimicry and the fact
that each can influence the evolution of the other. One would expect that
the more abundant species would be the model in a mullerian system,
since it is what the selective agent (predation) is cueing on. In general
H. erato is the more abundant of the two species and H. melpomene
mimics the wing patterns of H. erato. In one area of overlap of
the two species, H. melpomene is the more abundant and H. erato
assumes the hindwing band pattern of H. melpomene (see figure below).
Thus depending on local conditions, both species are influencing the adaptive
responses of the other and thus fits strict definition of coevolution.
A crucial component of coevolution is phylogenetic analysis. If the cladograms of the host and the cladograms of the parasite are congruent (e.g., figs. 22.2 - 22.3, pg. 612-613) this certainly suggests coevolutionary phenomena. But again, be careful and think about it: cospeciation is just "association by descent". Have there been reciprocal phenomena?; maybe just the speciation of the host induced the speciation of the parasite and there was not parasite induced speciation of the host. One needs to know the evolutionary history before we can make firm statements about "co"evolution.