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文件:Dasyscolia ciliata.jpg
授粉的黄蜂 Dasyscolia ciliata 在与Ophrys speculum花进行拟交配[1]

模板:Evolutionary biology In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well as gene-culture coevolution.在生物学中,当两个或多个物种通过自然选择过程相互影响彼此各自的演化时,就会发生共同演化。该词语有时用在同一物种中存在相互影响和演化的两个特征上,例如基因和文化的共同演化。

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 after the middle 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. 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.

共同演化作为自然界研究的一个主要课题,在20世纪60年代中期之后迅速扩大,丹尼尔·H·詹森(Daniel H. Janzen)展示了金合欢和蚂蚁之间的共同演化(见下文),保罗·R·埃利希(Paul R. Ehrlich)和彼得·H·雷文(Peter H. Raven)提出植物和蝴蝶之间的共同演化可能促进了两个群体的物种多样化。如今共同进化的理论基础已经颇为成熟(例如共同进化的地理镶嵌理论),而且向我们表明了共同演化在推动主要的进化转变中扮演着重要的角色,例如有性生殖的演化或者倍性的变化。[2][3]最近,共同进化也被证实可以影响生态群落的结构和功能以及共生群体的演化,例如植物和它们的传粉者,以及传染病的动态过程。[3][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.


Coevolution is primarily a biological concept, but researchers have applied it by analogy to fields such as computer science, sociology, and astronomy.



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.



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. He first mentioned coevolution as a possibility in On the Origin of Species, and developed the concept further in Fertilisation of Orchids (1862).

花朵在化石记录中比较突发地出现和多样化,创造了被查尔斯 · 达尔文描述为如此迅速进化的“令人憎恶的神秘”; 他考虑是否可以以共同演化作为解释。[8]他可能在物种起源中首次提到了共同演化,并在兰花的传粉(1862)中进一步发展了这个概念。[9][10][11][12]



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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.


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

一些非常成功的昆虫群体——尤其是膜翅目(黄蜂、蜜蜂和蚂蚁)和鳞翅目(蝴蝶和飞蛾)以及许多种双翅目(苍蝇)和鞘翅目(甲虫)——在白垩纪(1.45亿至6.6亿年前)与被子植物共同演化。最早的蜜蜂,今天重要的传粉者,出现在白垩纪早期。[15]一群与蜜蜂相近亲的黄蜂与被子植物同时演化,鳞翅目也是如此。[15]此外,所有主要的蜜蜂分支种系均首次出现在白垩纪中到晚期间,同时出现的是真根植物的辐射适应(占所有被子植物的四分之三) ,正当被子植物成为世界上陆地的主要植物时。[9]

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.


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

对丝兰(Yucca whipplei)这种植物只有斑点豆斑蛾才能够为其授粉,而斑点豆斑蛾是一种依靠丝兰生存的丝兰蛾。[16][17]花粉会随附在蛾进食植物的种子的过程之中被采集。花粉已经进化得非常粘着,当飞蛾移动到下一朵花时仍然会留在其口腔的部分。丝兰则在花的深处为蛾提供了一个产卵的地方,来远离潜在的捕食者。




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. 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. 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. 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.


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.


Ornithophilous flowers need to be conspicuous to birds. Birds have their greatest spectral sensitivity and finest hue discrimination at the red end of the visual spectrum, 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. 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.



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一颗无花果树 露出它许多细小的结种成熟雌蕊群。它们被无花果蜂Blastophaga psenes授粉,在 栽培的无花果树中也有无性 品种。[25]

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.花的容器,里面装着雌花或花粉



The acacia ant (Pseudomyrmex ferruginea) is an obligate plant ant that protects at least five species of "Acacia" (Vachellia)模板:Efn from preying insects and from other plants competing for sunlight, and the tree provides nourishment and shelter for the ant and its larvae. 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.

相思树蚁(Pseudomyrmex ferruginea)是一种能保护至少5种金合欢树(Vachellia)免受食用牛角相思树的昆虫和其他植物争夺阳光的专性植物蚂蚁,而这种树则为这种蚂蚁及其幼虫提供营养和庇护[26][27]。这种互利共生并不是自然而然的:其他蚂蚁种类遵循不同的进化策略,利用树木而不作回报;这些欺诈性的蚂蚁通过破坏树木的生殖器官对寄主施加重大伤害,不过它们对寄主健康的净影响并不一定是负面的,因此难以预测。[28][29]



Host–parasite coevolution is the coevolution of a host and a parasite. 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. 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". The host reproduces sexually, producing some offspring with immunity over its parasite, which then evolves in response.


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. 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.

寄生者和宿主的关系可能导致了有性生殖的流行,而不是更有效率的无性生殖。看起来,当寄生者感染宿主时,有性生殖提供了一个更好的机会来发展抵抗性(通过下一代的变异) ,这种可导致适应性的有性生殖变异性在无性生殖中则看不到,这样就更容易会产生另一代易受同一寄生者的感染的机体。[34][35][36]宿主和寄生者之间的共同演化可能相应导致了在普遍的群体中的许多遗传多样性,包括血浆多态性、蛋白多态性和组织相容性系统。[37]


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.



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.

拮抗性的共同演化关系在红收获蚁(Pogonomyrmex barbatus)和罗纹须蚁(Pogonomyrmex rugosus)之间可以看到,它们之间既有寄生关系也有互惠关系。蚁后无法通过与同类交配来繁殖工蚁。只有通过杂交,它们才能繁殖工蚁。有翅膀的雌性对其他物种的雄性像寄生一样,因为它们的精子只会繁殖不育的杂种。但由于殖民完全依赖于这些杂交种的生存,这也是互惠互利的。虽然两个物种之间没有基因交换,但它们不能朝向基因差异太大的方向演化,因为这将使杂交繁殖变得不可能。[41]


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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. Paul R. Ehrlich and Peter H. Raven in 1964 proposed the theory of escape and radiate coevolution to describe the evolutionary diversification of plants and butterflies. 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.


性冲突已经在黑腹果蝇(shown mating, male on right)的案例当中被研究


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



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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.



The geographic mosaic theory of coevolution was developed by John N. Thompson as a way of linking the ecological and evolutionary processes that shape interactions among species across ecosystems. It is based on three observations that are taken as assumptions: (1) species are usually groups of populations that are somewhat genetically distinct from each other, (2) interacting species often co-occur in only parts of their geographic ranges, and (3) interactions among species differ ecologically among environments.


From these assumptions, geographic mosaic theory suggests that natural selection on interactions among species is driven by three sources of variation:


1. Geographic selection mosaics occur in interactions among species, because genes are expressed in different ways in different environments and because different genes are favored in different environments. For example, natural selection on an interaction between a parasite population and a host population may differ between very dry environments and very wet environments. Alternatively, an interaction between two or more species may be antagonistic in some environments but mutualistic (beneficial to both or all species) in other environments.

  1. 地理选择镶嵌发生在物种的相互作用之间;因为基因在不同的环境中以不同的方式表达,而且不同的基因会在不同的环境中各受欢迎。例如,寄生虫种群和宿主种群之间相互作用的自然选择在非常干燥的环境和非常湿润的环境之间可能有所不同。或者,两个或两个以上物种之间的相互作用在某些环境中可能是对抗性的,但在其他环境中是互惠性的(对两个或所有物种都有益)。

2. Coevolutionary hotspots and coldspots occur because natural selection on interactions among species is reciprocal in some environments but not in others. For example, a symbiont population may decrease the survival or reproduction of its hosts in one environment, but it may have no effect on host survival or reproduction in another environment. When detrimental, natural selection will favor evolutionary responses in the host population, resulting in a coevolutionary hotspot of ongoing reciprocal evolutionary changes in the parasite and host populations. When the symbiont has no effect on the survival and reproduction of the host, natural selection on the symbiont population will not favor an evolutionary response by the host population (i.e, a coevolutionary coldspot).

2. 共同演化的热点和冷点的出现是因为物种间相互作用的自然选择在某些环境中是相互的,而在其他环境中则不然。例如,一个共生的生物种群可能会损害其宿主在一个环境中的生存或繁殖,但它可能对宿主在另一个环境中的生存或繁殖没有影响。当这一影响是负面的时候,自然选择将有利于宿主种群的进化反应,从而导致寄生虫和宿主种群中正在进行的相互演变的共同演化热点。当共生对宿主的生存和繁殖没有影响时,共生生物种群的自然选择则会不利于宿主种群的进化反应(即共同演化的冷点)。

3. Finally, there is constant remixing of the traits on which natural selection acts both locally and regionally. At any moment in time, a local population will have a unique combination of genes on which natural selection acts. These genetic differences among populations occur because each local population has a unique history of new mutations, genomic alterations (e.g., whole genome duplications), gene flow among populations from individuals arriving from other populations or going to other populations, random loss or fixation of genes at times when populations are small (random genetic drift), hybridization with other species, and other genetic and ecological processes that affect the raw genetic material on which natural selection acts. More formally, then, the geographic mosaic of coevolution can be viewed as a genotype by genotype by environment interaction (GxGxE) that results in the relentless evolution of interacting species.

3. 最后,自然选择作用在局部和区域两方面的特征-不断-重新混合。在任何时候,当地的种群都会有一个自然选择作用的独特基因组合。这些种群之间的遗传差异之所以会出现,是因为每个当地种群都有新的突变、基因组的改变(例如全基因组复制)、来自其他种群个体或个体朝向其他种群的种间基因流、在种群规模较小时随机丢失或加强的基因(随机遗传漂变)、与其他种群的杂交,以及影响自然选择之作用的原始遗传物质的其他遗传和生态过程。更正式地说,共同演化的地理镶嵌可被看作是一种基因型与环境的相互作用(GxGxE),这种作用导致了相互作用物种的无向演化。

Geographic mosaic theory has been explored through a wide range of mathematical models, studies of interacting species in nature, and laboratory experiments using microbial species and viruses.



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



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



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

共同进化的概念被丹麦建筑师兼城市规划专家亨利克 · 瓦勒尔在建筑学引入,作为“星架构”[58]的对立面。作为2006年威尼斯建筑双年展丹麦馆的馆长,他创建了一个关于中国城市发展共同进化的展览项目,并获得了最佳国家馆的金狮奖。[59][60][61][62]

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."



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.



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.



In Development Betrayed: The End of Progress and A Coevolutionary Revisioning of the Future (1994) Richard Norgaard proposes a coevolutionary cosmology to explain how social and environmental systems influence and reshape each other. 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".

发展的背叛:进步的终结和未来的共同演化修正(1994)[66]中,理查德·诺尔加尔提出了一个共同演化的宇宙学来解释社会和环境系统是如何相互影响和重塑的。[67]共同演化经济学:经济、社会和环境(1994)中,约翰 · 高迪认为: “经济、社会和环境在共同进化的关系中联系在一起。”。[68]



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. 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.


See also

  • Bak–Sneppen model
  • Coextinction
  • Ecological fitting
  • Escape and radiate coevolution
  • Genomics of domestication


  • Bak-Sneppen模型
  • 共同灭绝
  • 生态适应
  • 逃逸和辐射共同进化
  • 驯化的基因组学




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External links

  • Coevolution, video of lecture by Stephen C. Stearns (Open Yale Courses)


  • 共同演化,讲座视频 Stephen c. Stearns(耶鲁大学开放课程)

模板:Evolutionary psychology

Category:Ecological processes Category:Environmental terminology Category:Evolution of the biosphere Category:Evolutionary biology Category:Habitat

类别: 生态过程类别: 环境术语类别: 生物圈的进化类别: 进化生物学类别: 栖息地

This page was moved from wikipedia:en:Coevolution. Its edit history can be viewed at 共同演化/edithistory