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 (1859). The term coevolution was coined by Paul R. Ehrlich and 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.[2] More recently, it has also been demonstrated that coevolution influences the structure and function of ecological communities as well as the dynamics of infectious disease.[2]

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, and astronomy.

Dasyscolia ciliata
The pollinating wasp Dasyscolia ciliata in pseudocopulation with a flower of Ophrys speculum[1]


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.[3][4]

Flowering plants

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.[5][6] He first mentioned coevolution as a possibility in On the Origin of Species, and developed the concept further in Fertilisation of Orchids (1862).[7][8][9]

Insects and entomophilous flowers

Apis mellifera - Melilotus albus - Keila
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.[10][11] The term coevolution was coined by Paul R. Ehrlich and Peter H. Raven in 1964, to describe the evolutionary interactions of plants and butterflies.[12]

Several highly successful insect groups—especially the Hymenoptera (wasps, bees and ants) and Lepidoptera (butterflies) as well as many types of 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.[13] A group of wasps sister to the bees evolved at the same time as flowering plants, as did the Lepidoptera.[13] 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.[5]

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.[13][1]

The yucca, Yucca whipplei, is pollinated exclusively by Tegeticula maculata, a yucca moth that depends on the yucca for survival.[14] 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 predators.[15]

Birds and ornithophilous flowers

Purple-throated carib hummingbird feeding
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.[16]

Flowers have converged to take advantage of similar birds.[17] 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.[17][18] Bird-pollinated flowers usually have higher volumes of nectar and higher sugar production than those pollinated by insects.[19] This meets the birds' high energy requirements, the most important determinants of flower choice.[19] 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.[20] 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.[21] However, some flowers such as 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 as pollinators.[22]

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 co-adaptations.[19]

Ficus plant
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.[23]

Ornithophilous flowers need to be conspicuous to birds.[19] Birds have their greatest spectral sensitivity and finest hue discrimination at the red end of the visual spectrum,[19] 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.[19] Each of the two subfamilies of hummingbirds, the 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.[19]

Fig reproduction and fig wasps

The genus Ficus 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.[23]

Acacia ants and acacias

Ant - Pseudomyrmex species, on Bull Thorn Acacia (Acacia cornigera) with Beltian bodies, Caves Branch Jungle Lodge, Belmopan, Belize - 8505045055
Pseudomyrmex ant on bull thorn acacia (Vachellia cornigera) with Beltian bodies that provide the ants with protein[24]

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.[24][25] 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.[26][27]

Hosts and parasites

Parasites and sexually reproducing hosts

Host–parasite coevolution is the coevolution of a host and a parasite.[28] A general characteristic of many viruses, as 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.[29] 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 in Through the Looking-Glass: "it takes all the running you can do, to keep in the same place".[30] The host reproduces sexually, producing some offspring with immunity over its parasite, which then evolves in response.[31]

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 parasite.[32][33][34] 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.[35]

Brood parasites

Brood parasitism demonstrates close coevolution of host and parasite, for example in some 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.[36][37]

Antagonistic coevolution

Antagonistic coevolution is seen in the harvester ant species Pogonomyrmex barbatus and Pogonomyrmex rugosus, in a relationship both parasitic and mutualistic. The queens are unable to produce worker ants by mating with their own species. Only by crossbreeding can they produce workers. The winged females act as parasites for the males of the other species as their sperm will only produce sterile hybrids. But because the colonies are fully dependent on these hybrids to survive, it is also mutualistic. While there is no genetic exchange between the species, they are unable to evolve in a direction where they become too genetically different as this would make crossbreeding impossible.[38]

Predators and prey

Leopard kill - KNP - 001
Predator and prey: a leopard killing a bushbuck

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.[39]

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.[40]

Sexual conflict has been studied in Drosophila melanogaster (shown mating, male on right).


Both intraspecific competition, with features such as sexual conflict[41] and sexual selection,[42] and interspecific competition, such as between predators, may be able to drive coevolution.[43]

Guild or diffuse coevolution

Amegilla cingulata on long tube of Acanthus ilicifolius flower
Long-tongued bees and long-tubed flowers coevolved, whether pairwise or "diffusely" in groups known as guilds.[44]

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 flowers.[44][45][46]

Outside biology

Coevolution is primarily a biological concept, but has been applied to other fields by analogy.

In algorithms

Coevolutionary algorithms are used for generating artificial life as well as for optimization, game learning and machine learning.[47][48][49][50][51] Daniel Hillis added "co-evolving parasites" to prevent an optimization procedure from becoming stuck at local maxima.[52] Karl Sims coevolved virtual creatures.[53]

In architecture

The concept of coevolution was introduced in architecture by the Danish architect-urbanist Henrik Valeur as an antithesis to the concept of "star-architecture".[54] As the curator of the Danish Pavilion at the 2006 Venice Biennale of Architecture he conceived and orchestrated the exhibition-project CO-EVOLUTION: Danish/Chinese Collaboration on Sustainable Urban Development in China, which was awarded the Golden Lion for Best National Pavilion.[55]

The exhibition included urban planning projects for the cities of Beijing, Chongqing, Shanghai, and Xi'an, which had been developed in collaboration between young professional Danish architects and architecture students and professors and students of architecture from the four Chinese cities.[56] 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."[57] 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."[58]

At the School of Architecture, Planning and Landscape, Newcastle University, a coevolutionary approach to architecture has been defined as a design practice that engages students, volunteers and members of the local community in practical, experimental work aimed at "establishing dynamic processes of learning between users and designers."[59]

In cosmology and astronomy

In his book The Self-organizing Universe, Erich Jantsch attributed the entire evolution of the cosmos to coevolution.

In astronomy, an emerging theory proposes that black holes and galaxies develop in an interdependent way analogous to biological coevolution.[60]-->

In technology

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.[61] 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.[62]

In sociology

In Development Betrayed: The End of Progress and A Coevolutionary Revisioning of the Future (1994)[63] Richard Norgaard proposes a coevolutionary cosmology to explain how social and environmental systems influence and reshape each other.[64] In Coevolutionary Economics: The Economy, Society and the Environment (1994) John Gowdy suggests that: "The economy, society, and the environment are linked together in a coevolutionary relationship".[65]

See also


  1. ^ The acacia ant protects at least 5 species of "Acacia", now all renamed to Vachellia: V. chiapensis, V. collinsii, V. cornigera, V. hindsii, and V. sphaerocephala.


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External links

Autumn leaf color

Autumn leaf color is a phenomenon that affects the normal green leaves of many deciduous trees and shrubs by which they take on, during a few weeks in the autumn season, various shades of red, yellow, purple, black, blue, orange, magenta, and brown. The phenomenon is commonly called autumn colours or autumn foliage in British English and fall colors, fall foliage, or simply foliage in American English.

In some areas of Canada and the United States, "leaf peeping" tourism is a major contribution to economic activity. This tourist activity occurs between the beginning of color changes and the onset of leaf fall, usually around September and October in the Northern Hemisphere and April to May in the Southern Hemisphere.

CoEvolution Quarterly

CoEvolution Quarterly (1974–1985) was a journal descended from Stewart Brand's Whole Earth Catalog. Stewart Brand founded the CoEvolution Quarterly in 1974 using proceeds from the Whole Earth Catalog. It evolved out of the original Supplement to the Whole Earth Catalog. Fred Turner notes that in 1985, Brand merged CoEvolution Quarterly with The Whole Earth Software Review (a supplement to The Whole Earth Software Catalog) to create the Whole Earth Review.CoEvolution Quarterly became the first place to publish Ivan Illich's Vernacular Values.


Cospeciation is a form of coevolution in which the speciation of one species dictates speciation of another species and is most commonly studied in host-parasite relationships. In the case of a host-parasite relationship, if two hosts of the same species get within close proximity of each other, parasites of the same species from each host are able to move between individuals and mate with the parasites on the other host. However, if a speciation event occurs in the host species, the parasites will no longer be able to "cross over" because the two new host species no longer mate and, if the speciation event is due to a geographic separation, it is very unlikely the two hosts will interact at all with each other. The lack of proximity between the hosts ultimately prevents the populations of parasites from interacting and mating. This can ultimately lead to speciation within the parasite.According to Fahrenholz's rule, first proposed by Heinrich Fahrenholz in 1913, when host-parasite cospeciation has occurred, the phylogenies of the host and parasite come to mirror each other. In host-parasite phylogenies, and all species phylogenies for that matter, perfect mirroring is rare. Host-parasite phylogenies can be altered by host switching, extinction, independent speciation, and other ecological events, making cospeciation harder to detect. However, cospeciation is not limited to parasitism, but has been documented in symbiotic relationships like those of gut microbes in primates.

Dual inheritance theory

Dual inheritance theory (DIT), also known as gene–culture coevolution or biocultural evolution, was developed in the 1960s through early 1980s to explain how human behavior is a product of two different and interacting evolutionary processes: genetic evolution and cultural evolution. Genes and culture continually interact in a feedback loop, changes in genes can lead to changes in culture which can then influence genetic selection, and vice versa. One of the theory's central claims is that culture evolves partly through a Darwinian selection process, which dual inheritance theorists often describe by analogy to genetic evolution.'Culture', in this context is defined as 'socially learned behavior', and 'social learning' is defined as copying behaviors observed in others or acquiring behaviors through being taught by others. Most of the modelling done in the field relies on the first dynamic (copying) though it can be extended to teaching. Social learning at its simplest involves blind copying of behaviors from a model (someone observed behaving), though it is also understood to have many potential biases, including success bias (copying from those who are perceived to be better off), status bias (copying from those with higher status), homophily (copying from those most like ourselves), conformist bias (disproportionately picking up behaviors that more people are performing), etc.. Understanding social learning is a system of pattern replication, and understanding that there are different rates of survival for different socially learned cultural variants, this sets up, by definition, an evolutionary structure: cultural evolution.Because genetic evolution is relatively well understood, most of DIT examines cultural evolution and the interactions between cultural evolution and genetic evolution.


Entomophily or insect pollination is a form of pollination whereby pollen of plants, especially but not only of flowering plants, is distributed by insects. Flowers pollinated by insects typically advertise themselves with bright colours, sometimes with conspicuous patterns (honey guides) leading to rewards of pollen and nectar; they may also have an attractive scent which in some cases mimics insect pheromones. Insect pollinators such as bees have adaptations for their role, such as lapping or sucking mouthparts to take in nectar, and in some species also pollen baskets on their hind legs. This required the coevolution of insects and flowering plants in the development of pollination behaviour by the insects and pollination mechanisms by the flowers, benefiting both groups.

Many plants, including flowering plants such as grasses, are instead pollinated by other mechanisms, such as by wind.

Escape and radiate coevolution

Escape and radiate coevolution is a multistep process that hypothesizes that an organism under constraints from other organisms will develop new defenses, allowing it to "escape" and then "radiate" into differing species. After a novel defense has been acquired, an organism is able to escape predation and rapidly multiply into new species because of relaxed selective pressure. There are many possible mechanisms available varying between different types of organisms, however they must be novel in order for escape to allow for radiation. This theory applies to predator-prey associations, but is most often applied to plant-herbivore associations.

This form of coevolution can be complex but is essential to understanding the vast biological diversity among organisms today. Out of the many forms of coevolution, escape and radiate is most likely responsible for providing the most diversity. This is due to the nature of the "evolutionary arms race" and the continuous cycle of counter adaptations. It is a relatively new field of study and is rapidly gaining credibility. To date, there has not been a formal study published specifically for escape and radiate coevolution.

Evolutionary arms race

In evolutionary biology, an evolutionary arms race is a struggle between competing sets of co-evolving genes, traits, or species, that develop adaptations and counter-adaptations against each other, resembling an arms race. These are often described as examples of positive feedback. The co-evolving gene sets may be in different species, as in an evolutionary arms race between a predator species and its prey (Vermeij, 1987), or a parasite and its host. Alternatively, the arms race may be between members of the same species, as in the manipulation/sales resistance model of communication (Dawkins & Krebs, 1979) or as in runaway evolution or Red Queen effects. One example of an evolutionary arms race is in sexual conflict between the sexes, often described with the term Fisherian runaway. Thierry Lodé emphasized the role of such antagonistic interactions in evolution leading to character displacements and antagonistic coevolution.

Evolutionary history of plants

The evolution of plants has resulted in a wide range of complexity, from the earliest algal mats, through multicellular marine and freshwater green algae, terrestrial bryophytes, lycopods and ferns, to the complex gymnosperms and angiosperms of today. While many of the earliest groups continue to thrive, as exemplified by red and green algae in marine environments, more recently derived groups have displaced previously ecologically dominant ones, e.g. the ascendance of flowering plants over gymnosperms in terrestrial environments.There is evidence that cyanobacteria and multicellular photosynthetic eukaryotes lived in freshwater communities on land as early as 1 billion years ago, and that communities of complex, multicellular photosynthesizing organisms existed on land in the late Precambrian, around 850 million years ago.Evidence of the emergence of embryophyte land plants first occurs in the mid-Ordovician (~470 million years ago), and by the middle of the Devonian (~390 million years ago), many of the features recognised in land plants today were present, including roots and leaves. By Late Devonian (~370 million years ago) some free-sporing plants such as Archaeopteris had secondary vascular tissue that produced wood and had formed forests of tall trees. Also by late Devonian, Elkinsia, an early seed fern, had evolved seeds.

Evolutionary innovation continued throughout the rest of the Phanerozoic eon and still continues today. Most plant groups were relatively unscathed by the Permo-Triassic extinction event, although the structures of communities changed. This may have set the scene for the appearance of the flowering plants in the Triassic (~200 million years ago), and their later diversification in the Cretaceous and Paleogene. The latest major group of plants to evolve were the grasses, which became important in the mid-Paleogene, from around 40 million years ago. The grasses, as well as many other groups, evolved new mechanisms of metabolism to survive the low CO2 and warm, dry conditions of the tropics over the last 10 million years.

Fig wasp

Fig wasps are wasps of the superfamily Chalcidoidea which spend their larval stage inside figs. Most are pollinators but others simply feed off the plant. The non-pollinators belong to several groups within the superfamily Chalcidoidea, while the pollinators are in the family Agaonidae. While pollinating fig wasps are gall-makers, the remaining types either make their own galls or usurp the galls of other fig wasps; reports of them being parasitoids are considered dubious.

Herbivore adaptations to plant defense

Herbivores are dependent on plants for food, and have coevolved mechanisms to obtain this food despite the evolution of a diverse arsenal of plant defenses against herbivory. Herbivore adaptations to plant defense have been likened to "offensive traits" and consist of those traits that allow for increased feeding and use of a host. Plants, on the other hand, protect their resources for use in growth and reproduction, by limiting the ability of herbivores to eat them. Relationships between herbivores and their host plants often results in reciprocal evolutionary change. When a herbivore eats a plant it selects for plants that can mount a defensive response, whether the response is incorporated biochemically or physically, or induced as a counterattack. In cases where this relationship demonstrates "specificity" (the evolution of each trait is due to the other), and "reciprocity" (both traits must evolve), the species are thought to have coevolved. The escape and radiation mechanisms for coevolution, presents the idea that adaptations in herbivores and their host plants, has been the driving force behind speciation. The coevolution that occurs between plants and herbivores that ultimately results in the speciation of both can be further explained by the Red Queen hypothesis. This hypothesis states that competitive success and failure evolve back and forth through organizational learning. The act of an organism facing competition with another organism ultimately leads to an increase in the organism's performance due to selection. This increase in competitive success then forces the competing organism to increase its performance through selection as well, thus creating an "arms race" between the two species. Herbivores evolve due to plant defenses because plants must increase their competitive performance first due to herbivore competitive success.

Host–parasite coevolution

Host–parasite coevolution is a special case of coevolution, the reciprocal adaptive genetic change of a host and a parasite through reciprocal selective pressures.

It is characterized by reciprocal genetic change and thus changes in allele frequencies within populations. These are determined by three main types of selection dynamics: negative frequency-dependent selection when a rare allele has a selective advantage; overdominance caused by heterozygote advantage; and directional selective sweeps near an advantageous mutation.

Theories of host–parasite coevolution include the geographic mosaic theory, which assumes a selection mosaic, coevolutionary hotspots, and geographic mixing; the Red Queen hypothesis, which proposes that parasitism favours sexual reproduction in the host; and an evolutionary trade-off between transmission and virulence, since if the parasite kills its host too quickly, the parasite will not be able to reproduce.

Model systems include the nematode Caenorhabditis elegans with the bacterium Bacillus thuringiensis; the crustacean Daphnia and its numerous parasites; and Escherichia coli and the mammals (including humans) whose intestines it inhabits.

Interlocus sexual conflict

Interlocus sexual conflict is a type of sexual conflict that occurs through the interaction of a set of antagonistic alleles at two or more different loci in males and females, resulting in the deviation of either or both sexes from the fitness optima for the traits.Interlocus sexual conflict involves a co-evolutionary arms race between the two sexes in which either sex evolves a set of antagonistic adaptations that are detrimental to the fitness of the other sex. Interlocus sexual conflict can occur over aspects of male–female interactions such as mating frequency, fertilization, relative parental effort, female remating behavior, and female reproductive rate. The evolutionary pathways resulting from interlocus sexual conflict form part of interlocus contest evolution.


The Passifloraceae are a family of flowering plants, containing about 750 species classified in around 27 genera.They include trees, shrubs, lianas, and climbing plants, and are mostly found in tropical regions. The family takes its name from the passion flower genus (Passiflora) which includes the edible passion fruit (Passiflora edulis), as well as garden plants such as maypop and running pop.

Passiflora vines and Dryas iulia (among other heliconian butterflies) have demonstrated evidence of coevolution, in which the plants attempted to stop their destruction from larval feeding by the butterflies, while the butterflies tried to gain better survival for their eggs.The former Cronquist system of classification placed this family in the order Violales, but under more modern classifications systems such as that proposed by the Angiosperm Phylogeny Group, this is absorbed into the Malpighiales.

Pollinator Partnership

The Pollinator Partnership or P2 is a 501(c)(3) non-profit organization headquartered in San Francisco, California that works to protect the health of managed and native pollinating animals that are vital to wildland and agricultural ecosystems. The Pollinator Partnership’s mission of environmental stewardship and pollinator protection is achieved through conservation, policy, education, and research. Signature initiatives include the NAPPC (North American Pollinator Protection Campaign), National Pollinator Week, and EcoRegional Planting Guides that allow local citizens to plant gardens that provide habitats for important pollinating species.


Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the host) and parasitoidism (which always does, eventually). It is distinct from scavenging on dead prey, though many predators also scavenge; it overlaps with herbivory, as a seed predator is both a predator and a herbivore.

Predators may actively search for prey or sit and wait for it. When prey is detected, the predator assesses whether to attack it. This may involve ambush or pursuit predation, sometimes after stalking the prey. If the attack is successful, the predator kills the prey, removes any inedible parts like the shell or spines, and eats it.

Predators are adapted and often highly specialized for hunting, with acute senses such as vision, hearing, or smell. Many predatory animals, both vertebrate and invertebrate, have sharp claws or jaws to grip, kill, and cut up their prey. Other adaptations include stealth and aggressive mimicry that improve hunting efficiency.

Predation has a powerful selective effect on prey, and the prey develop antipredator adaptations such as warning coloration, alarm calls and other signals, camouflage, mimicry of well-defended species, and defensive spines and chemicals. Sometimes predator and prey find themselves in an evolutionary arms race, a cycle of adaptations and counter-adaptations. Predation has been a major driver of evolution since at least the Cambrian period.


The Prodoxidae are a family of moths, generally small in size and nondescript in appearance. They include species of moderate pest status, such as the currant shoot borer, and others of considerable ecological and evolutionary interest, such as various species of "yucca moths".

Reproductive coevolution in Ficus

The genus Ficus is composed of 800 species of vines, shrubs, and trees, defined by their syconiums, the fruit-like vessels that either hold female flowers or pollen on the inside. In addition to being cultivated by humans for thousands of years, Ficus is also known for their reproductive mutualism with the fig wasp.

Fig trees either produce hermaphrodite fruit known as caprifigs, or female figs; only the female figs are palatable to humans. In exchange for a safe place for their eggs and larvae, fig wasps help pollinate the ficus by crawling inside the tiny hole in the apex of the fig, called the ostiole, without knowing whether they crawled into a caprifig or a fig. If the female wasp crawls into the caprifig, she can successfully lay her eggs and die. The males hatch first, mate with the females, dig tunnels out of the caprifig, and die. The females, now covered in fig pollen from the caprifig, fly out to begin the cycle again. If the female wasp crawls into a female fig, she will not be able to successfully lay her eggs despite pollinating the fig with pollen from the caprifig she hatched in. The fig will absorb her body and her eggs as the fruit develops.

Stewart Brand

Stewart Brand (born December 14, 1938) is an American writer, best known as editor of the Whole Earth Catalog. He founded a number of organizations, including The WELL, the Global Business Network, and the Long Now Foundation. He is the author of several books, most recently Whole Earth Discipline: An Ecopragmatist Manifesto.

Whole Earth Review

Whole Earth Review (Whole Earth after 1997) was a magazine which was founded in January 1985 after the merger of the Whole Earth Software Review (a supplement to the Whole Earth Software Catalog) and the CoEvolution Quarterly. All of these periodicals are descendants of Stewart Brand's Whole Earth Catalog.

The last published hard copy issue of the magazine was the Winter 2002 issue. The next issue (Spring, 2003) was planned but never published in hard copy format. Bruce Sterling attempted to solicit funds for this issue by writing that "friends at Whole Earth Magazine have experienced a funding crunch so severe that the Spring 2003 special issue (#111) on Technological Singularity, edited by Alex Steffen of the Viridian curia, hasn't been printed and distributed. Whole Earth is soliciting donations to get the issue printed, and has put some of the content online." Eventually, elements of the 2003 issue appeared only in digital format on the Whole Earth website.

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