Theory of Coevolution

Over billions of years, life has evolved into the extraordinarily diverse and complex organisms
that populate the Earth today. Although evolution often proceeds toward increasing complexity,
more complex traits do not necessarily make organisms more fit. The theory of biological
evolution is one of the most important, central theories of biology. Many evolution-theoretical
theses can be proven, owing to the enormous scientific progress recently made in many
biological fields. The most important evolutionary factors are mutation, recombination, and
selection.
Coevolution is defined as a process in the course of which two partners or entire partner
systems(animals, plants, bacteria, or fungi) depend on one another in their evolution. Both
acquire specific adaptations as a consequence of mutual selection pressure. Examples for a
gene–gene coevolution, for a close coevolution between species (specific coevolution), as well
as for a coevolution between species groups (diffuse or guild coevolution) will be presented.
These examples concern flower–pollinator systems, the dispersal of plant species by
animals,protection of plants by animal species, and coevolution between plant species and
phytophagous insects.

The term coevolution is used to describe cases where two (or more) species reciprocally affect
each other's evolution. So for example, an evolutionary change in the morphology of a plant,
might affect the morphology of an herbivore that eats the plant, which in turn might affect the
evolution of the plant, which might affect the evolution of the herbivore...and so on.

Coevolution is likely to happen when different species have close ecological interactions with
one another. These ecological relationships include:

1. Predator/prey and parasite/host

2. Competitive species
3. Mutualistic species

Plants and insects represent a classic case of coevolution — one that is often, but not always,
mutualistic. Many plants and their pollinators are so reliant on one another and their
relationships are so exclusive that biologists have good reason to think that the "match"
between the two is the result of a coevolutionary process.

But we can see exclusive "matches" between plants and insects even when pollination is not
involved. Some Central American Acacia species have hollow thorns and pores at the bases of
their leaves that secrete nectar. These hollow thorns are the exclusive nest-site of some species
of ant that drink the nectar. But the ants are not just taking advantage of the plant — they also
defend their acacia plant against herbivores.

This system is probably the product of coevolution: the plants would not have evolved hollow
thorns or nectar pores unless their evolution had been affected by the ants, and the ants would
not have evolved herbivore defense behaviors unless their evolution had been affected by the
plants.

Gene-by-Gene Coevolution

If a certain gene of a parasite, which codes for virulence,is complemented by a gene of its host,
which codes for resistance to the parasite, this is called gene-by-gene coevolution. This kind of
interaction especially occurs between plants and pathogenic fungi. In a population of Glycine
canescens, 11 different phenotypic resistance patterns, based on at least 12 genetic resistance
factors, could be identified as response to nine genotypes of the rust fungus Phakopsora
pachyrhizi.

Coevolution between Two Species

A close coevolution can be particularly shown for certain mutualistic or symbiotic systems.
Examples for the systems flower– pollinator and plant species–animal dispersal are given.

Flower–Pollinator Systems

The pollination of figs is very complicated .The urn-shaped inflorescences (syconia) of a fig tree,
in which the extremely reduced flowers line the inner cavity, attract thousands of fig wasps of
only a few millimeters in length. Each fig species (worldwide there are more than 1000) is
pollinated by its “own” wasp species. The female wasp, transporting the pollen in two thorax
bags,crawls through the very narrow, scale-covered opening of the inflorescence (ostiole) into
the cavity. The stigmata are covered with pollen. Then the wasp jabs its ovipositor into the style
of a flower and places an egg there. At this spot, as a consequence of the puncture and certain
substances secreted by the fig wasp, a cell growth (gall) develops. The larva lives in this gall
and feeds on the cell tissue. During this time, the carbon dioxide content within the syconium
increases; at the end of the larval development it amounts to about 10%. At this concentration,
only the male fig wasps are active. They fertilize the females in the gall, then leave it and bore a
hole into the wall of the syconium. With decreasing carbon dioxide concentration, the female
wasps become active. They leave the galls, fill both thorax bags with pollen, which has
meanwhile been secreted by the carpellate (male) flowers, and leave the syconium. The
process then starts a new. In order to avoid eggs being placed on all flowers, there are two
different female flower types:short styled and long styled. Eggs are only placed in short-styled
flowers; the ovipositor does not reach far enough into the long-styled ones.

Dispersal of Plant Species by Animals (Zoochory)

There is a very close relationship between the nutcracker Nucifraga caryocatactes

caryocatactes and the arolla pine Pinus cembra. In Europe, the nutcracker is found in
mountainous habitats with fir- and larch-arolla forests. It mainly feeds on the nuts of the arolla
cone which it pecks open with its chisel-shaped peak. For the winter,the nutcracker hides cedar
nuts. One individual may have more than 10,000 hiding places. The “forgotten” hiding places
are essential for the young growth of arolla pines.The nutcracker also selects hiding places at
the timberline,thus promoting the growth of trees there. Similar relation-ships have been shown,
for example, for the Japanese nut-cracker N. caryocatactes japonicus and P. pumila in Japan.
On the island of Mauritius, today there remain only a few specimens of the endemic tree
species Calvaria major. All the Calvaria trees are about 300 years old,although every year
plenty of fruits and seeds are formed;thus, a rejuvenation should be possible. That it does not
occur is due to the bisystem of C. major and an indigenous bird species, the dodo Raphus
cucullatus, which became, as it can be proven,extinct in the Year 1681. The dodo ate the
approximately 5 cm long fruits of C. major. In its muscular stomach, with the help of stones
contained therein, it filed off part of the1.5 cm-thick seed coat; therefore, the seeds could
germinate. Since extinction of the dodo, artificially filing off the seeds of Calvaria has been
attempted to achieve germination.

Mutualism

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 types.

Birds and bird-pollinated flowers

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.
 Hosts and parasites

Hosts and parasites exert reciprocal selective pressures on each other, which may lead to rapid
reciprocal adaptation. For organisms with short generation times, host–parasite coevolution can
be observed in comparatively small time periods, making it possible to study evolutionary
change in real-time.

The dynamics of these interactions are summarized in the Red Queen hypothesis, namely that
both host and parasite have to change continuously to keep up with each other's adaptations.

Host–parasite coevolution is ubiquitous and of potential importance to all living organisms,

including humans, domesticated animals and crops. Major diseases such as malaria, AIDS and
influenza are caused by coevolving parasites. Better understanding of coevolutionary
adaptations between parasite attack strategies and host immune systems may assist in the
development of novel medications and vaccines.

Predator and prey

One particularly stark example of coevolutionary predator-prey system exists between the
African honey badger and the African honey bee. The African honey badger (Mellivora
capensis) primarily feeds on rodents, insects and arachnids in its native environment. However,
as one might expect from its name, the honey badger has a particular love of honey. It uses its
long claws to dislodge hives from their mounted positions, cutting teeth to tear open the hives,
and its thick, loose skin to ward off stings from the bees. In response, the African bee (Apis
meliffera scutellata) developed a behavioural adaptation: massive swarming. Honey badgers,
relatively immune to the stings of a bee, require a full-out attack by the hive. As the bees die
post-sting, they release a pheromone calling to others to continue the attack. They will also
pursue their quarry for up to a kilometer.

Therefore, predator-prey adaptation is the process of escalation in which a competitive

advantage given through natural selection for either predator or prey pushes for adaptation in its
counterpart. This shifts the advantage and begins a new round of competitive adaptation. By
nature these systems are antagonistic since the effectiveness of one species' adaptations
negatively impacts the survival of the other and are measured by the effectiveness of the prey's
defense against the predator.

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 anti-predator
adaptations.

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 herbivore.

Coevolution of Species Groups (Diffuse Coevolution)

Examples of a coevolution between species groups can be found particularly where certain
guilds are involved in the ecological structure (guild coevolution). A guild is defined as a group
of species using the same class of environmental resources in a similar manner —for instance,
phloem feeders (greenfly: Aphidina), pollinators, and predators. In this context, it is important
that several different species may be evolutionary vectors.In mutualism, there are numerous
examples of this form of coevolution: the many interactions between plants and their animal
pollinators, between plants and their animal dispersers, and between plants and their animal
“protectors".

Plants and Their Animal Dispersers: Myrmecochory

The phenomenon by which ants disperse the seeds of plants, an ecologically significant
ant-plant interaction with worldwide distribution.

Most myrmecochorous plants produce seeds rich in lipids, amino acids, or other nutrients that
are attractive to ants. The seed with its attached elaiosome is collectively known as a diaspore.
Seed dispersal by ants is typically accomplished when foraging workers carry diaspores back to
the ant colony, after which the elaiosome is removed or fed directly to ant larvae. Once the
elaiosome is consumed, the seed is usually discarded in underground middens or ejected from
the nest. Although diaspores are seldom distributed far from the parent plant, myrmecochores
also benefit from this predominantly mutualistic interaction through dispersal to favourable
locations for germination, as well as escape from seed predation.

Many plant species contain certain secondary substances, to protect themselves from being
eaten (chemical defence). Coevolution is a complex process that occurs on many levels. It may
appear in situations where one species interacts closely with several others, such as the
interaction between European cuckoos (Cuculus canorus) and the other species whose nests
they parasitize; it may involve many species, as in relationships between fruit-bearing plants
and birds; or it may take place in some subgroups of species but not others. It is important to
note that human activities often disrupt the process of coevolution by changing the nature and
the extent of the interactions between coevolving species.