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Title: Coevolution  
Author: World Heritage Encyclopedia
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Subject: Ecological fitting, Host–parasite coevolution, Adaptation, Co-adaptation, Pseudocopulation
Collection: Biology Terminology, Ecological Processes, Environmental Terminology, Evolutionary Biology, Habitat
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Bumblebees and the flowers they pollinate have coevolved so that both have become dependent on each other for survival.

In biology, coevolution is "the change of a biological object triggered by the change of a related object".[1] In other words, when changes in at least two species' genetic compositions reciprocally affect each other’s evolution, coevolution has occurred.

There is evidence for coevolution at the level of populations and species. Charles Darwin briefly described the concept of coevolution in On the Origin of Species (1859) and developed it in detail in Fertilisation of Orchids (1862).[2][3][4] It is likely that viruses and their hosts coevolve in various scenarios.[5]

However, there is little evidence of coevolution driving large-scale changes in Earth's history, since abiotic factors such as mass extinction and expansion into ecospaces seem to guide the shifts in the abundance of major groups.[6] One proposed specific example was the evolution of high-crowned teeth in grazers when grasslands spread through North America - long held up as an example of coevolution. We now know that these events happened independently.[7]

Coevolution can occur at many biological levels: it can be as microscopic as correlated mutations between amino acids in a protein or as macroscopic as covarying traits between different species in an environment. Each party in a coevolutionary relationship exerts host species and its parasites (host–parasite coevolution[8]), and examples of mutualism evolving through time. Evolution in response to abiotic factors, such as climate change, is not biological coevolution (since climate is not alive and does not undergo biological evolution).

The general conclusion is that coevolution may be responsible for much of the genetic diversity seen in normal populations including: blood-plasma polymorphism, protein polymorphism, histocompatibility systems, etc.[9]

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 viability for fitness not seen in the asexual reproduction, which produces another generation of the organism susceptible to infection by the same parasite.[10][11]

Coevolution is primarily a biological concept, but researchers have applied it by analogy to fields such as computer science, sociology / international political economy[12] and astronomy.


  • Models 1
  • Coevolution types 2
  • Coevolution in the fossil record 3
  • Specific examples 4
    • Hummingbirds and ornithophilous flowers 4.1
    • Angraecoid orchids and African moths 4.2
    • Old world swallowtail and fringed rue 4.3
    • Garter snake and rough-skinned newt 4.4
    • California buckeye and pollinators 4.5
    • Acacia ant and bullhorn acacia tree 4.6
    • Yucca Moth and the yucca plant 4.7
  • Coevolution outside biology 5
    • Biological applications 5.1
    • Algorithms 5.2
    • Architecture 5.3
    • Cosmology and astronomy 5.4
    • Technological coevolution 5.5
  • See also 6
  • References 7
  • Further reading 8
  • External links 9


One model of coevolution was Leigh Van Valen's Red Queen's Hypothesis, which states that "for an evolutionary system, continuing development is needed just in order to maintain its fitness relative to the systems it is co-evolving with".[13] This hypothesis predicts that sexual reproduction allows a host to stay just ahead of its parasite by a generation, similar to the Red Queen in "Through the Looking Glass". …always running ….. Just ahead. The essence is that the host reproduces sexually giving it immunity over its parasite, which then evolves in response. This requires the next generation to repeat the sequence.[14] Emphasizing the importance of sexual conflict, Thierry Lodé described the role of antagonist interactions in evolution, giving rise to a concept of antagonist coevolution.[15] Coevolution branching strategies for asexual population dynamics in limited resource environments have been modeled using the generalized Lotka–Volterra equations.[16] A model based on adaptive dynamics and experimental data of floral and proboscis lengths, as well as nectar consumed and pollen deposited during the pollination of the long-tubed iris (Lapeirousia anceps) by the long-proboscid fly (Moegistorhynchus longirostris) has generated diverse coevolutionary dynamics, including two types of Red Queen dynamics, evolutionary branching (backed by observations of coexisting irises of short and long tubes in a single population) and trap.[17]

Coevolution types

Coevolution can occur between pairs of entities (often referred to as pairwise coevolution) exists, with examples including predator and prey, host and symbiont or host and parasite. However, some instances of coevolution are less clearcut: a species may evolve in response to more than one species. If this coevolution is occurring in a non-additive way, then this type of coevolution is known as diffuse coevolution.[18]

A general characterization that can be made of many viruses, widely known to be obligate parasites, is that they coevolved alongside their respective hosts. This is suggested by the similar genetic arrangement between virus and host.[19] Correlated mutations between the two species enter them into an evolution arms race: the host must develop a defense mechanism to overcome the parasite, and the parasite must overcome the new defense mechanism in order to persist and reproduce. 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.

Coevolution in the fossil record

Many examples of coevolution have been observed among living species but coevolution has not been conclusively shown in the fossil record. This may be because the fossil record does not tend to preserve high resolution data (e.g. on the level of species) and because coevolution does not drive large-scale changes in Earth's history.[6] Even classic examples of coevolution in the fossil record such as the interaction between bees and the flowers they pollinate or the development of grazers in spreading grasslands are little supported by evidence. In the case of bees and flowers, these two organisms evolved at different times - Grit, not grass hypothesis.

Specific examples

Hummingbirds and ornithophilous 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 upon 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.[21]

Flowers have converged to take advantage of similar birds.[22] Flowers compete for pollinators, and adaptations reduce unfavourable effects of this competition. The fact that birds can fly during inclement weather can make them more efficient pollinators in cases in which bees and other insects are inactive. Ornithophily may have arisen for this reason in isolated environments with poor insect colonization or areas with plants which flower in the winter.[22][23] Bird-pollinated flowers usually have higher volumes of nectar and higher sugar production than those pollinated by insects.[24] This meets the birds' high energy requirements, which are the most important determinants of their flower choice.[24] In experiments with monkeyflowers, an increase in red pigment in petals and flower nectar volume has been shown to noticeably reduce the proportion of pollination by bees as opposed to solely attract hummingbirds; while greater flower surface area was shown to increase the amount of bee pollination. Additional red coloration of flowers significantly decreased bee visitation but seemingly had no effect on the frequency of hummingbird visitation. Thus, hummingbirds may not necessarily have an innate preference for red and the high concentration of these red pigments in the flowers of M. cardinalis could potentially function primarily to discourage bee visitation. In first generation hybrids of these two species, the composition of pollinator visitors (59% bees; 1,744 visits) was exactly in between the two parental species, implying a strong genetic component to pollinator coevolution.[25] In similar experiments with Penstemon flowers, pollen transfer by both bees and hummingbirds was recorded with two closely related species species differing in main pollinator. The results suggest that the flower traits that discourage bee pollination may be even more influential on the flowers' evolutionary change than ‘pro-bird’ adaptations are. However, adaptation 'towards' birds and adaptation 'away' from bees can and do happen simultaneously.[26] 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 seem to have converged to a common morphology and color.[24] Different lengths and curvatures of the corolla tubes can affect the efficiency of extraction in hummingbird species in relation to differences in bill morphology.[24] 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 also allows the plant to place pollen on a certain part of the bird's body.[24] This opens the door for a variety of morphological co-adaptations.

An important requirement for attraction is conspicuousness to birds, which reflects the properties of avian vision and habitat features.[24] Birds have their greatest spectral sensitivity and finest hue discrimination at the red end of the visual spectrum,[24] so red is particularly conspicuous to them. Hummingbirds may also be able to see ultraviolet "colors".[24] The prevalence of ultraviolet patterns and nectar guides in nectar-poor entomophilous (insect-pollinated) flowers warns the bird to avoid these flowers.[24]

Hummingbirds form the family Trochilidae, whose two subfamilies are the Phaethornithinae (hermits) and the Trochilinae. Each subfamily 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.[24]

Angraecoid orchids and African moths

Angraecoid orchids and African moths coevolve because the moths are dependent on the flowers for nectar and the flowers are dependent on the moths to spread pollen so they can reproduce also leading to evolutionary growth. Coevolution has led to deep flowers and moths with long proboscides.

Old world swallowtail and fringed rue

Old world swallowtail caterpillar on fringed rue

An example of antagonistic coevolution is the old world swallowtail (Papilio machaon) caterpillar living on the fringed rue (Ruta chalepensis) plant. The rue produces etheric oils which repel plant-eating insects. The old world swallowtail caterpillar developed resistance to these poisonous substances, thus reducing competition with other plant-eating insects.

Garter snake and rough-skinned newt

Coevolution of predator and prey species is illustrated by the Rough-skinned newt (Taricha granulosa) and the common garter snake (Thamnophis sirtalis). The newts produce a potent neurotoxin that concentrates in their skin. Garter snakes have evolved resistance to this toxin through a series of genetic mutations, and prey upon the newts. The relationship between these animals has resulted in an evolutionary arms race that has driven toxin levels in the newt to extreme levels. This is an example of coevolution because differential survival caused each organism to change in response to changes in the other.

California buckeye and pollinators

When beehives are populated with bee species that have not coevolved with the California buckeye (Aesculus californica), sensitivity to aesculin, a neurotoxin present in its nectar, may be noticed; this sensitivity is only thought to be present in honeybees and other insects that did not coevolve with A. californica.[27]

Acacia ant and bullhorn acacia tree


  • Coevolution, video of lecture by Stephen C. Stearns (Open Yale Courses)
  • [2], Global Optimization of Some Difficult Benchmark Functions by Host-Parasite Coevolutionary Algorithm: An algorithm for Global Optimization.

External links

  • Dawkins, R. Unweaving the Rainbow.
  • Geffeney, Shana L., et al. "Evolutionary diversification of TTX-resistant sodium channels in a predator-prey interaction". Nature 434 (2005): 759–763.
  • Michael Pollan The Botany of Desire: A Plant's-eye View of the World. Bloomsbury. ISBN 0-7475-6300-4. Account of the co-evolution of plants and humans
  • Thompson, J. N. (1994). The Coevolutionary Process. Chicago: University of Chicago Press. pp. 376 pp.  
  • Mintzer, Alex; Vinson, S.B. "Kinship and incompatibility between colonies of the acacia ant Pseudomyrex ferruginea". Behavioral Ecology and Sociobiology 17 (1): 75–78.   Abstract
  • Armstrong, W.P. "The Yucca and its Moth". Wayne's Word.  

Further reading

  1. ^ Yip; Patel, P; Kim, PM; Engelman, DM; McDermott, D; Gerstein, M; et al. (2008). "An integrated system for studying residue coevolution in proteins". Bioinformatics 24 (2): 290–292.  
  2. ^ Thompson, John N. (1994). The coevolutionary process. Chicago:  
  3. ^ Darwin, Charles (1859). On the Origin of Species (1st ed.). London: John Murray. Retrieved 2009-02-07. 
  4. ^ Darwin, Charles (1877). On the various contrivances by which British and foreign orchids are fertilised by insects, and on the good effects of intercrossing (2nd ed.). London: John Murray. Retrieved 2009-07-27. 
  5. ^ C.Michael Hogan. 2010. . Encyclopedia of EarthVirus. Editors: Cutler Cleveland and Sidney Draggan
  6. ^ a b Sahney, S., Benton, M.J. and Ferry, P.A. (2010). "Links between global taxonomic diversity, ecological diversity and the expansion of vertebrates on land" (PDF). Biology Letters 6 (4): 544–547.  
  7. ^ Jardine, P.E., Janis, C.M., Sahney, S., and Benton, M.J. (2012), "Grit not grass: Concordant patterns of early origin of hypsodonty in Great Plains ungulates and Glires", Palaeogeography, Palaeoclimatology, Palaeoecology: 1–10 
  8. ^ Rabajante, J; et al. (2015). "Red Queen dynamics in multi-host and multi-parasite interaction system".  
  9. ^ Anderson, R., and May, R. (1982), Coevolution of hosts and parasites, Parasitology, Cambridge Journals, retrieved from
  10. ^ Editors (2011), Sexual reproduction works thanks to ever-evolving host, parasite relationships: study, Physorg, retrieved from
  11. ^ L.T. Morran; O.G. Schmidt; I.A. Gelarden; R.C. Parrish II; C.M. Lively. "Running with the Red Queen: Host-Parasite Coevolution Selects for Biparental Sex," Science, July 8, 2011. Document:Science.1206360. Indiana University.
  12. ^ Jessop, Bob (2004). "Critical Semiotic Analysis and Critical Political Economy". Critical Discourse Studies 1 (1): 1–16. 
  13. ^ Van Valen L. (1973): "A New Evolutionary Law", Evolutionary Theory 1, pp. 1–30. Cited in: The Red Queen Principle
  14. ^ Sterns, S. (2009), Coevolution, EEB-122: Principles of evolution, ecology, and behavior, Open Yale Courses, retrieved from
  15. ^ Lodé, Thierry (2007). La guerre des sexes chez les animaux, une histoire naturelle de la sexualité. Paris: Odile Jacob.  
  16. ^ G. S. van Doorn, F. J. Weissing (April 2002). "Ecological versus Sexual Selection Models of Sympatric Speciation: A Synthesis" (PDF). Selection (Budapest, Hungary: Akadémiai Kiadó) 2 (1-2): 17–40.  
  17. ^ Zhang, F.; Hui, C.; Pauw, A. (2013). "Adaptive divergence in Darwin's race: how coevolution can generate trait diversity in a pollination system". Evolution 67: 548–560.  
  18. ^ Juenger, Thomas, and Joy Bergelson. "Pairwise versus diffuse natural selection and the multiple herbivores of scarlet gilia, Ipomopsis aggregata." Evolution (1998): 1583-1592.
  19. ^ Adrian J.Gibbs, Charles H.Calisher and Fernando Garcia-Arenal. 1995. Molecular basis of virus evolution. 603 pages
  20. ^ Wilford, J.N. (1995). "Which Came First: Bees or Flowers? Find Points to Bees" (PDF). New York Times. 
  21. ^ Kay, Kathleen M.; Reeves, Patrick A.; Olmstead, Richard G.; Schemske, Douglas W. (2005). "Rapid speciation and the evolution of hummingbird pollination in neotropical Costus subgenus Costus (Costaceae): evidence from nrDNA ITS and ETS sequences.". American Journal of Botany 92: 1899–1910.  
  22. ^ a b Brown James H., Kodric-Brown Astrid (1979). "Convergence, Competition, and Mimicry in a Temperate Community of Hummingbird-Pollinated Flowers". Ecology 60 (5): 1022–1035.  
  23. ^ Cronk, Quentin; Ojeda, Isidro (2008). "Bird-pollinated flowers in an evolutionary and molecular context". Journal of Experimental Botany 59: 715–727.  
  24. ^ a b c d e f g h i j Stiles, F. Gary (1981). "Geographical Aspects of Bird Flower Coevolution, with Particular Reference to Central America". Annals of the Missouri Botanical Garden 68 (2): 323–351.  
  25. ^ Schemske, Douglas W.; Bradshaw, H.D. (1999). "Pollinator preference and the evolution of floral traits in monkeyflowers (Mimulus).". National Academy of Sciences 96 (21): 11910–11915.  
  26. ^ Castellanos, M. C.; Wilson, P.; Thomson, J.D. (2005). "'Anti-bee' and 'pro-bird' changes during the evolution of hummingbird pollination in Penstemon flowers". Journal of Evolutionary Biology 17: 876–885.  
  27. ^ C. Michael Hogan (13 September 2008). California Buckeye: Aesculus californica,
  28. ^ National Geographic. "Acacia Ant Video". 
  29. ^ Palmer TM, Doak DF, Stanton ML, Bronstein JL, Kiers ET, Young TP, Goheen JR, Pringle RM (2010). "Synergy of multiple partners, including freeloaders, increases host fitness in a multispecies mutualism". Proceedings of the National Academy of Sciences of the United States of America 107 (40): 17234–9.  
  30. ^ Hemingway, Claire (2004). "Pollination Partnerships Fact Sheet" (PDF). Flora of North America: 1–2. Retrieved 2011-02-18. Yucca and Yucca Moth 
  31. ^ Pellmyr, Olle; James Leebens-Mack (August 1999). "Forty million years of mutualism: Evidence for Eocene origin of the yucca-yucca moth association" (PDF). Proc. Natl. Acad. Sci. USA 96 (16): 9178–9183.  
  32. ^ "Henrik Valeur's biography". Retrieved 2015-08-29. 
  33. ^ "An interview with Henrik Valeur". Movingcities. Retrieved 2015-10-17. 
  34. ^ "About the exhibition". Danish Architecture Centre. Retrieved 2015-08-29. 
  35. ^ Valeur, Henrik (2006). CO-EVOLUTION: Danish/Chinese Collaboration on Sustainable Urban Development in China. Copenhagen: Danish Architecture Centre.  
  36. ^ Valeur, Henrik (2014). India: the Urban Transition - a Case Study of Development Urbanism. Copenhagen: Architectural Publisher B.  
  37. ^ Britt, Robert. "The New History of Black Holes: 'Co-evolution' Dramatically Alters Dark Reputation". 
  38. ^ Theo D’Hondt, Kris De Volder, Kim Mens and Roel Wuyts, Co-Evolution of Object-Oriented Software Design and Implementation, TheKluwer International Series in Engineering and Computer Science, 2002, Volume 648, Part 2, 207–224, doi:10.1007/978-1-4615-0883-0_7
  39. ^ Cherns, A. (1976). "The principles of sociotechnical design". In". Human Relations 29: 8.  


See also


Technological coevolution

In astronomy, an emerging theory states that black holes and galaxies develop in an interdependent way analogous to biological coevolution.[37]

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

Cosmology and astronomy

Henrik 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."[36]

By creating a framework for collaboration between academics and professionals representing two distinct cultures it was thought that new solutions to shared problems (i.e. global pollution, ecological destruction and resource depletion resulting from the processes of rapid urbanization in China) could be developed and that the exchange of knowledge, ideas and experiences could stimulate creativity and imagination in order "to set the spark for new visions for sustainable urban development."[35]

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 professors and students from universities in the four Chinese cities.[34]

The concept of coevolution was introduced in architecture by the Danish architect-urbanist Henrik Valeur as an antithesis to the concept of "star-architecture".[32] 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.[33]


Coevolutionary algorithms are a class of algorithms used for generating artificial life as well as for optimization, game learning and machine learning. Coevolutionary methods have been applied by Daniel Hillis, who coevolved sorting networks, and Karl Sims, who coevolved virtual creatures.


The study of coevolution in natural populations could help in fields such as conservation, human epidemiology, and improved agriculture.

Biological applications

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

Coevolution outside biology

In this mutualistic symbiotic relationship, the yucca plant (Yucca whipplei) is pollinated exclusively by Tegeticula maculata, a species of yucca moth that in turn relies on the yucca for survival.[30] Yucca moths tend to visit the flowers of only one species of yucca plant. In the flowers, the moth eats the seeds of the plant, while at the same time gathering pollen on special mouth parts. The pollen is very sticky, and will easily remain on the mouth parts when the moth moves to the next flower. The yucca plant also provides a place for the moth to lay its eggs, deep within the flower where they are protected from any potential predators.[31] The adaptations that both species exhibit characterize coevolution because the species have evolved to become dependent on each other.

A flowering yucca plant that would be pollinated by a yucca moth

Yucca Moth and the yucca plant


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