Coevolution
In biology, coevolution occurs when two or more
Charles Darwin mentioned evolutionary interactions between flowering plants and insects in On the Origin of Species (1859). Although he did not use the word coevolution, he suggested how plants and insects could evolve through reciprocal evolutionary changes. Naturalists in the late 1800s studied other examples of how interactions among species could result in reciprocal evolutionary change. Beginning in the 1940s, plant pathologists developed breeding programs that were examples of human-induced coevolution. Development of new crop plant varieties that were resistant to some diseases favored rapid evolution in pathogen populations to overcome those plant defenses. That, in turn, required the development of yet new resistant crop plant varieties, producing an ongoing cycle of reciprocal evolution in crop plants and diseases that continues to this day.
Coevolution as a major topic for study in nature expanded rapidly from the 1960s, when Daniel H. Janzen showed coevolution between acacias and ants (see below) and Paul R. Ehrlich and Peter H. Raven suggested how coevolution between plants and butterflies may have contributed to the diversification of species in both groups. The theoretical underpinnings of coevolution are now well-developed (e.g., the geographic mosaic theory of coevolution), 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][3] More recently, it has also been demonstrated that coevolution can influence the structure and function of ecological communities, the evolution of groups of mutualists such as plants and their pollinators, and the dynamics of infectious disease.[2][4]
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 multi-species coevolution, which is sometimes called guild or diffuse coevolution, several to many species may evolve a trait or a group of traits in reciprocity with a set of traits in another species, as has happened between the flowering plants and pollinating insects such as bees, flies, and beetles. There are a suite of specific hypotheses on the mechanisms by which groups of species coevolve with each other.[5]
Coevolution is primarily a biological concept, but researchers have applied it by analogy to fields such as computer science, sociology, and astronomy.
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.[6][7]
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.[8][9] He first mentioned coevolution as a possibility in On the Origin of Species, and developed the concept further in Fertilisation of Orchids (1862).[7][10][11]
Insects and insect-pollinated 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.[12][13]
Several highly successful
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.[14][1]
The yucca, Yucca whipplei, is pollinated exclusively by Tegeticula maculata, a yucca moth that depends on the yucca for survival.[15] 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.[16]
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.[17]
Flowers have converged to take advantage of similar birds.[18] 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.[18][19] Bird-pollinated flowers usually have higher volumes of nectar and higher sugar production than those pollinated by insects.[20] This meets the birds' high energy requirements, the most important determinants of flower choice.[20] 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.[21] 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.[22] 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.[23]
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.[20]
Ornithophilous flowers need to be conspicuous to birds.
Fig reproduction and fig wasps
The genus

Acacia ants and acacias
The
Hosts and parasites
Parasites and sexually reproducing hosts
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.[33][34][35] 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.[36]
Brood parasites
Antagonistic coevolution
Antagonistic coevolution is seen in the
Predators and prey
The same applies to

Competition
Both intraspecific competition, with features such as sexual conflict[44] and sexual selection,[45] and interspecific competition, such as between predators, may be able to drive coevolution.[46]
Intraspecific competition can result in sexual antagonistic coevolution, an evolutionary relationship analogous to an arms race, where the evolutionary fitness of the sexes is counteracted to achieve maximum reproductive success. For example, some insects reproduce using traumatic insemination, which is disadvantageous to the female's health. During mating, males try to maximise their fitness by inseminating as many females as possible, but the more times a female's abdomen is punctured, the less likely she is to survive, reducing her fitness.[47]
Multispecies

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 reciprocal, but is among a group of species rather than exactly two. This is variously 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.[48][49][50]
Geographic mosaic theory
Mosaic coevolution is a theory in which
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
In architecture
The concept of coevolution was introduced in architecture by the Danish architect-urbanist Henrik Valeur as an antithesis to "star-architecture".[61] As the curator of the Danish Pavilion at the 2006 Venice Biennale of Architecture, he created an exhibition-project on coevolution in urban development in China; it won the Golden Lion for Best National Pavilion.[62][63][64][65]
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."[66]
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.[67]
In management and organization studies
Since year 2000, a growing number of management and organization studies discuss coevolution and coevolutionary processes. Even so, Abatecola el al. (2020) reveals a prevailing scarcity in explaining what processes substantially characterize coevolution in these fields, meaning that specific analyses about where this perspective on socio-economic change is, and where it could move toward in the future, are still missing.[68]
In sociology
In Development Betrayed: The End of Progress and A Coevolutionary Revisioning of the Future (1994)
In technology
See also
- Evolutionary arms race
- Bak–Sneppen model
- CoEvolution Quarterly
- Coextinction
- Ecological fitting
- Escape and radiate coevolution
- Genomics of domestication
Notes
- ^ 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
- Coevolution, video of lecture by Stephen C. Stearns (Open Yale Courses)