1152:" has four possible outcomes: (a) they both betray each other, and are both sentenced to two years in prison; (b) A betrays B, which sets A free and B is sentenced to four years in prison; (c) B betrays A, with the same result as (b) except that it is B who is set free and the other spends four years in jail; (d) both remain silent, resulting in a six-month sentence each. Clearly (d) ("cooperation") is the best mutual strategy, but from the point of view of the individual betrayal is unbeatable (resulting in being set free, or getting only a two-year sentence). Remaining silent results in a four-year or six-month sentence. This is exemplified by a further example of the PDG: two strangers attend a restaurant together and decide to split the bill. The mutually best ploy would be for both parties to order the cheapest items on the menu (mutual cooperation). But if one member of the party exploits the situation by ordering the most expensive items, then it is best for the other member to do likewise. In fact, if the fellow diner's personality is completely unknown, and the two diners are unlikely ever to meet again, it is always in one's own best interests to eat as expensively as possible. Situations in nature that are subject to the same dynamics (rewards and penalties) as the PDG define cooperative behavior: it is never in the individual's fitness interests to cooperate, even though mutual cooperation rewards the two contestants (together) more highly than any other strategy. As described in the
1251:, better golfers have fewer strokes subtracted from their raw scores than the less talented players. The handicap therefore correlates with unhandicapped performance, making it possible, if one knows nothing about the horses, to predict which unhandicapped horse would win an open race. It would be the one handicapped with the greatest weight in the saddle. The handicaps in nature are highly visible, and therefore a peahen, for instance, would be able to deduce the health of a potential mate by comparing its handicap (the size of the peacock's tail) with those of the other males. The loss of the male's fitness caused by the handicap is offset by his increased access to females, which is as much of a fitness concern as is his health. A cooperative act is, by definition, similarly costly (e.g. helping raise the young at the nest of an unrelated pair of birds versus producing and raising one's own offspring). It would therefore also signal fitness, and is probably as attractive to females as a physical handicap. If this is the case, cooperation is evolutionarily stabilized by
1171:' 1971 reciprocal altruism theory) is a robust strategy which promotes altruism. In "tit-for-tat" both players' opening moves are cooperation. Thereafter each contestant repeats the other player's last move, resulting in a seemingly endless sequence of mutually cooperative moves. However, mistakes severely undermine tit-for-tat's effectiveness, giving rise to prolonged sequences of betrayal, which can only be rectified by another mistake. Since these initial discoveries, all the other possible IPD game strategies have been identified (16 possibilities in all, including, for instance, "generous tit-for-tat", which behaves like "tit-for-tat", except that it cooperates with a small probability when the opponent's last move was "betray".), but all can be outperformed by at least one of the other strategies, should one of the players switch to such a strategy. The result is that none is
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1502:). This also applies to gene B and so on through the alphabet. If a coin-flip "heads" means that gene X is the same in offspring-1 as it is in offspring-2, then in 26 flips of the coin approximately half are going to be "heads" and the rest "tails", i.e. half the genes inherited from parent "m" will be the same in the two offspring. The same will happen to the genes inherited from parent "f". Thus of the 52 genes inherited from the two parents, on average, 13 + 13 = 26 (or half) will be identical in the two sibs. Thus sibs are genetically as similar to one another as a parent is to an offspring. From a evolutionary genetic point of view it is therefore as advantageous to help with the upbringing of full sibs as it is to produce and raise one's own offspring.
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956:. Cancer cells divide uncontrollably, and at the cellular level, they are very successful, because they are (in the short term) reproducing very well and out competing other cells in the body. However, at the whole organism level, cancer is often fatal, and so may prevent reproduction. Therefore, changes to the genome which prevent cancer (for example, by causing damaged cells to act co-operatively by destroying themselves) are favoured. Multi-level selection theory contends that similar effects can occur, for example, to cause individuals to co-operate to avoid behaviours which favour themselves short-term, but destroy the community (and their descendants) long term.
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better than living alone, and cooperation arises passively as a result of many animals doing the same thing. By-product benefit can also arise as a consequence of subordinate animals staying and helping a nest that is dominated by leaders who often suffer high mortality rates. It has been shown that cooperation would be most advantageous for the sex that is more likely to remain and breed in the natal group. This is because the subordinate will have a higher chance to become dominant in the group as time passes. Cooperation in this scenario is often seen between non-related members of the same species, such as the wasp
837:. He illustrated this with a scenario having two hunters, each hunter having the choice of hunting (cooperate) or not hunting (free-riding). Assuming that cooperative hunting results in greater rewards than just a one-player hunt, when hunting is not rare, both hunters and non-hunters benefit because either player is likely to be with other hunters, and thus likely to reap the rewards of a successful hunt. This situation demonstrates "Prisoner's Delight" because the food of a successful hunt is shared between the two players regardless of whether or not they participated.
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1458:. This individual's genome therefore consists of 52 genes. The subscript indicates from which parent that copy of A has come. Mostly the two copies are identical, but occasionally they differ slightly. When this individual reproduces sexually, one or other copy of A (chosen randomly) is passed on to offspring-1, who gets its other copy of A from the sexual partner. The same happens with genes B, C, D, ..., Z. If we denote the two sexual partners by means of subscripts "m" and "f", then the genome of the offspring they produce might consist of A
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1005:—of which only mutualism can sometimes qualify as cooperation. Mutualism involves a close, mutually beneficial interaction between two different biological species, whereas "cooperation" is a more general term that can involve looser interactions and can be interspecific (between species) or intraspecific (within a species). In commensalism, one of the two participating species benefits, while the other is neither harmed nor benefitted. In parasitism, one of the two participating species benefits at the expense of the other.
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1498:. Each parent has contributed exactly half of the offspring's genome. So individual "m" shares only half of its genome with its offspring. Suppose individuals "m" and "f" produce a second offspring (offspring-2), whose genome is determined in exactly the same manner. There is a coin-flip 50% probability that offspring-2 will inherit the same copy of A from "m" as offspring-1 did (i.e. A
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917:, but with the green-beard effect, the actor has to instead identify which of its social partners share the gene for cooperation. A green-beard system must always co-occur within individuals and alleles to produce a perceptible trait, recognition of this trait in others, and preferential treatment to those recognized. Examples of green-beard behavior have been found in
937:, where blue-throated males preferentially establish territories next to each other. Results show that neighboring blue-throats are more successful at mate guarding. However, blue males next to larger, more aggressive orange males suffer a cost. This strategy blue has evolutionary cycles of altruism alternating with mutualism tied to the RPS game.
1287:. This will have the effect of a sexual population rapidly shedding peripheral phenotypic features, thereby canalizing the entire outward appearance and behavior of all of its members. They will all very quickly begin to look remarkably similar to one another in every detail, as illustrated in the accompanying photograph of the
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In a study of 79 students, participants played a game in which they could repeatedly give money to others and receive from others. They were told that they would never interact with the same person in the reciprocal role. A player's history of donating was displayed at each anonymous interaction, and
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Kin discrimination also influences cooperation because the actor can give aid preferentially towards related partners. Since kin usually share common genes, it is thought that this nepotism can lead to genetic relatedness between the actor and the partner's offspring, which affects the cooperation an
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showed individual differences in the extent to which they participated in group-territorial conflict. Some lions consistently 'cooperated' by approaching intruders, while others 'lagged' behind to avoid the risk of fighting. Although the lead female recognized the laggards, she failed to punish them,
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in that it assumes strategies are heritable and are subject to natural selection. Economic models are useful for analyzing cooperative relationships because they provide predictions on how individuals act when cooperation is an option. Economic models are not perfect, but they provide a general idea
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has to be evolutionarily costly. Thus, if a (low fitness) liar were to use the highly costly signal, which seriously eroded its real fitness, it would find it difficult to maintain a semblance or normality. Zahavi borrowed the term "handicap principle" from sports handicapping systems. These systems
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By-product benefit arises as a consequence of social partners having a shared interest in cooperation. For example, in meerkats, larger group size provides a benefit to all the members of that group by increasing survival rates, foraging success and conflict wins. This is because living in groups is
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will generally stand out in a population. This is because that altered appearance and functionality will be unusual, peculiar, and different from the norm within that population. The norm against which these unusual features are judged is made up of fit attributes that have attained their plurality
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Not all examples of mutualism are also examples of cooperation. Specifically, in by-product mutualism, both participants benefit, but cooperation is not involved. For example, when an elephant defecates, this is beneficial to the elephant as a way to empty waste, and it is also beneficial to a dung
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Multi-level selection theory suggests that selection operates on more than one level: for example, it may operate at an atomic and molecular level in cells, at the level of cells in the body, and then again at the whole organism level, and the community level, and the species level. Any level which
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Hamilton originally suggested that high relatedness could arise in two ways: direct kin recognition between individuals or limited dispersal, or population viscosity, which can keep relatives together. The easiest way to generate relatedness between social partners is limited dispersal, a mechanism
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Studies conducted on red wolves support previous researchers' contention that helpers obtain both immediate and long-term gains from cooperative breeding. Researchers evaluated the consequences of red wolves' decisions to stay with their packs for extended periods of time after birth. While delayed
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Contrary to the mainstream dogma, a recently published article. using agent-based models demonstrates that several crucial mechanisms, such as kin selection, punishment, multilevel selection, and spatial structure, cannot rescue the evolution of cooperation. The new findings revive a long-standing
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This mechanism can be relied to both within a species or social group and within species systems. It can also be applied to a multi-partner system, in which the owner of a resource has the power to choose its co-operation partner. This model can be applied in natural systems (examples exist in the
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There is an alternate strategy for identifying fit mates which does not rely on one gender having exaggerated sexual ornaments or other handicaps, but is probably generally applicable to most, if not all sexual creatures. It derives from the concept that the change in appearance and functionality
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In the late 1900s, some early research in animal cooperation focused on the benefits of group-living. While living in a group produces costs in the form of increased frequency of predator attacks and greater mating competition, some animals find that the benefits outweigh the costs. Animals that
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Cooperative hunting by wolves allows them to tackle much larger and more nutritious prey than any individual wolf could handle. However, such cooperation could, potentially, be exploited by selfish individuals who do not expose themselves to the dangers of the hunt, but nevertheless share in the
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enzyme. The legume benefits from a new supply of usable nitrogen from the rhizobia, and the rhizobia benefits from organic acid energy sources from the plant as well as the protection provided by the root nodule. Since the rhizobia live within the legume, this is an example of endosymbiosis, and
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Different theories explaining kin selection have been proposed, including the "pay-to-stay" and "territory inheritance" hypotheses. The "pay-to-stay" theory suggests that individuals help others rear offspring in order to return the favor of the breeders allowing them to live on their land. The
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However, some researchers, such as ecology professor Tim
Clutton-Brock, assert that cooperation is a more complex process. They state that helpers may receive more direct, and less indirect, gains from assisting others than is commonly reported. These gains include protection from predation and
1336:, but more recent studies focus on non-kin cooperation, where benefits may seem less obvious. Non-kin cooperation often involves many strategies that include manipulation and coercion, making these interactions more complicated to study. An example of manipulation is presented by the cuckoo, a
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The inclusive fitness theory provides a good overview of possible solutions to the fundamental problem of cooperation. The theory is based on the hypothesis that cooperation helps in transmitting underlying genes to future generations either through increasing the reproductive successes of the
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Cooperation in animals appears to occur mostly for direct benefit or between relatives. Spending time and resources assisting a related individual may at first seem destructive to an organism's chances of survival but is actually beneficial over the long-term. Since relatives share part of the
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generated by the photobiont, and the photobiont benefits from the increased water retention and increased surface area to capture water and mineral nutrients conferred by the mycobiont. Many lichens are examples of obligate symbiosis. In fact, one-fifth of all known extant fungal species form
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behavior often decreases the reproductive success of the actor (the individual performing the cooperative behavior). Hence, cooperation seemed to pose a challenging problem to the theory of natural selection, which rests on the assumption that individuals compete to survive and maximize their
861:, where dominant females will attack and evict subordinate females who become pregnant. The pregnancy is seen as a failure to cooperate because only the dominant females are allowed to bear offspring. Dominant females will attack and kill the offspring of subordinate females if they evade
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is a problem through enforcement, which is the mechanism where the actor is rewarded for cooperating or punished for not cooperating. This happens when cooperation is favored in aiding those who have helped the actors in the past. Punishment for noncooperation has been documented in
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dispersal helped other wolves' offspring, studies also found that it extended male helper wolves' life spans. This suggests that kin selection may not only benefit an individual in the long-term through increased fitness but also in the short-term through increased survival chances.
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world of apes, cleaner fish, and more). Easy for exemplifying, though, are systems from international trading. Arabic countries control vast amounts of oil, but seek technologies from western countries. These in turn are in need of Arab oil. The solution is co-operation by trade.
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codifies this problem and has been the subject of much research, both theoretical and experimental. In its original form the prisoner's dilemma game (PDG) described two awaiting trial prisoners, A and B, each faced with the choice of betraying the other or remaining silent. The
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individual (direct fitness) or of other individuals who carry the same genes (indirect fitness). Direct benefits can result from simple by-product of cooperation or enforcement mechanisms, while indirect benefits can result from cooperation with genetically similar individuals.
3345:"The phenotypic differences between carrion and hooded crows across the hybridization zone in Europe are unlikely to be due to assortative mating. Comment on Poelstra, J.W. et al. (2014). The genomic landscape underlying phenotypic integrity in the face of gene flow in crows"
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Another example of a hidden benefit is indirect reciprocity, in which a donor individual helps a beneficiary to increase the probability that observers will invest in the donor in the future, even when the donor will have no further interaction with the beneficiary.
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sounds an alarm call to warn other group members of a nearby coyote, it draws attention to itself and increases its own odds of being eaten. There have been multiple hypotheses for the evolution of cooperation, all of which are rooted in
Hamilton's models based on
844:, or reaping the benefits without any effort, is often a problem in collective action. Examples of free riding would be if an employee in a labor union pays no dues, but still benefits from union representation. In a study published in 1995, scientists found that
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This process contrasts with intragroup competition where individuals work against each other for selfish reasons. Cooperation exists not only in humans but in other animals as well. The diversity of taxa that exhibits cooperation is quite large, ranging from
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Some research suggests that individuals provide more help to closer relatives. This phenomenon is known as kin discrimination. In their meta-analysis, researchers compiled data on kin selection as mediated by genetic relatedness in 18 species, including the
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is a type of non-cooperative game theory that assumes an individual's decision is influenced by its knowledge of the strategies of other individuals. This theory was novel because it took into consideration the higher cognitive capabilities of animals. The
1340:, which lays its eggs in the nest of a bird of another species. That bird then is tricked into feeding and caring for the cuckoo offspring. Although this phenomenon may look like cooperation at first glance, it only presents benefits to one recipient.
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Clutton-Brock, T; Brotherton, P. N.; Russell, A. F.; O'Riain, M. J.; Gaynor, D.; Kansky, R.; Griffin, A.; Manser, M.; Sharpe, L.; McIlrath, G. M.; Small, T.; Moss, A.; Monfort, S. (2001). "Cooperation, control, and concession in meerkat groups".
1283:, while less well adapted attributes will be in the minority or frankly rare. Since the overwhelming majority of mutant features are maladaptive, and it is impossible to predict evolution's future direction, sexual creatures would be expected to
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between efficiency obtaining a desired resource and the amount of resources one can actively obtain. In that case, each partner in a system could benefit from specializing in producing one specific resource and obtaining the other resource by
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of the selfish individual who hangs back from the rest of the hunting pack, but who nevertheless joins in the spoils, that individual will be recognized as being different from the norm, and will therefore find it difficult to attract a mate
1308:). Its genes will therefore have only a very small probability of being passed on to the next generation, thus evolutionarily stabilizing cooperation and social interactions at whatever level of complexity is the norm in that population.
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in which genetic similarity correlates with spatial proximity. If individuals do not move far, then kin usually surrounds them. Hence, any act of altruism would be directed primarily towards kin. This mechanism has been shown in
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Symbiosis may be obligate or facultative. In obligate symbiosis, one or both species depends on the other for survival. In facultative symbiosis, the symbiotic interaction is not necessary for the survival of either species.
1112:, the presence of helpers at the nest does not lead to an increase in chick mass. However, the presence of helpers does confer a hidden benefit: it increases the chance that a mother will survive to breed in the next year.
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In the light of the iterated prisoner's dilemma game and the reciprocal altruism theory failing to provide full answers to the evolutionary stability of cooperation, several alternative explanations have been proposed.
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One theory suggesting a mechanism that could lead to the evolution of co-operation is the "market effect" as suggested by Noe and
Hammerstein. The mechanism relies on the fact that in many situations there exists a
734:). As explained below, direct benefits encompass by-product benefits and enforced reciprocity, while indirect benefits (kin selection) encompass limited dispersal, kin discrimination and the greenbeard effect.
1054:). In this interaction, rhizobia bacteria induce root nodule formation in legume plants via an exchange of molecular signals. Within the root nodules, rhizobia fix atmospheric nitrogen into ammonia using the
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models, were used to determine decisions made by animals in cooperative relationships. However, complicated interactions between animals have required the use of more complex economic models such as the
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are benefits from cooperation that are not obvious because they are obscure or delayed. (For example, a hidden benefit would not involve an increase in the number of offspring or offspring viability.)
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reproductive successes. Additionally, some species have been found to perform cooperative behaviors that may at first sight seem detrimental to their own evolutionary fitness. For example, when a
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practice group-living often benefit from assistance in parasite removal, access to more mates, and conservation of energy in foraging. Initially, the most obvious form of animal cooperation was
770:. They found that different species exhibited varying degrees of kin discrimination, with the largest frequencies occurring among those who have the most to gain from cooperative interactions.
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increased reproductive fitness. Furthermore, they insist that cooperation may not solely be an interaction between two individuals but may be part of the broader goal of unifying populations.
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because the act of cooperation is preferentially directed at individuals who have helped the actor in the past (directly), or helped those who have helped the actor in the past (indirectly).
1175:, and any prolonged series of the iterated prisoner's dilemma game, in which alternative strategies arise at random, gives rise to a chaotic sequence of strategy changes that never ends.
784:, a climbing plant, results show that kin groups have higher efficiency rates in growth than non-kin groups do. This is expected to rise out of reduced competition within the kin groups.
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helper's genetic make-up, enhancing each individual's chance of survival may actually increase the likelihood that the helper's genetic traits will be passed on to future generations.
730:. These models hypothesize that cooperation is favored by natural selection due to either direct fitness benefits (mutually beneficial cooperation) or indirect fitness benefits (
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Sinervo, Barry; Alexis Chaine; Jean
Clobert; Ryan Calsbeek; Lisa Hazard; Lesley Lancaster; Andrew G. McAdam; Suzanne Alonzo; Gwynne Corrigan; Michael E. Hochberg (1 May 2006).
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donations were significantly more frequent to receivers who had been generous to others in earlier interactions. Indirect reciprocity has only been shown to occur in humans.
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One specific form of cooperation in animals is kin selection, which involves animals promoting the reproductive success of their kin, thereby promoting their own fitness.
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is not competitive with others of the same level will be eliminated, even if the level below is highly competitive. A classic example is that of genes which prevent
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This is also called mutually beneficial cooperation as both actor and recipient depend on direct fitness benefits, which are broken down into two different types:
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noted that if the same contestants in the PDG meet repeatedly (in the so-called iterated prisoner's dilemma game, IPD) then tit-for-tat (foreshadowed by
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beetle that uses the elephant's dung. However, neither participant's behavior yields a benefit from the other, and thus cooperation is not taking place.
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Noe, R.; Hammerstein, P. (1994). "Biological markets: supply and demand determine the effect of partner choice in cooperation, mutualism and mating".
1295:. Once a population has become as homogeneous in appearance as is typical of most species, its entire repertoire of behaviors will also be rendered
978:, and trade for the other. Trading for the resource requires co-operation with the other partner and includes a process of bidding and bargaining.
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refers to two or more biological species that interact closely, often over a long period of time. Symbiosis includes three types of interactions—
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believes is a handicap, comparable to a race horse's handicap. The larger the handicap the more intrinsically fit the individual (see text).
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746:"territory inheritance" theory contends that individuals help in order to have improved access to breeding areas once the breeders depart.
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Strassmann, J; Seppa, P.; Queller, D. C. (10 August 2000). "Absence of within-colony kin discrimination: foundresses of the social wasp,
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An individual's gene complement (or genome) can be represented by the letters of the alphabet. Each letter is represented twice: A
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1226:. Both are costly in fitness terms, and both are generally conspicuous to other members of the population or species. This led
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Koeslag, J. H. (1990). "Koinophilia groups sexual creatures into species, promotes stasis, and stabilizes social behaviour".
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1266:, showing details of appearance and coloration that are shared by all African pygmy kingfishers to a high degree of fidelity.
1247:, provenly faster horses are given heavier weights to carry under their saddles than inherently slower horses. Similarly, in
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bacteria, where cooperation is disfavored when populations are well mixed, but favored when there is high local relatedness.
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1858:"Paying to stay or paying to breed? Field evidence for direct benefits of helping behavior in a cooperatively breeding fish"
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Lutzoni, François; Pagel, Mark; Reeb, Valérie (2001). "Major fungal lineages are derived from lichen symbiotic ancestors".
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Griffin, A. S.; West, S. A. (2003). "Kin
Discrimination and the Benefit of Helping in Cooperatively Breeding Vertebrates".
1320:, who noted it as a potential problem for his theory of natural selection. In most of the 19th century, intellectuals like
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is the process where groups of organisms work or act together for common or mutual benefits. It is commonly defined as any
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puzzle in the evolution theory. In addition, the work has potential therapeutic benefits for numerous incurable diseases.
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are aimed at reducing disparities in performance, thereby making the outcome of contests less predictable. In a horse
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in 2007 after he found that benefits can result as an automatic consequence of an otherwise "self-interested" act in
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debated fervently on whether animals cooperate with one another and whether animals displayed altruistic behaviors.
684:. Many animal and plant species cooperate with both members of their own species and with members of other species.
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Grafen, A. (1984) Natural selection, kin selection and group selection. In Krebs, J. R. & Davies, N. B. (Eds.)
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Heinsohn, R; Packer, C. (1 September 1995). "Complex cooperative strategies in group-territorial
African lions".
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Evolutionary mechanisms suggesting that reciprocity is the result, not the cause, of the evolution of cooperation
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Even if all members of a group benefit from cooperation, individual self-interest may not favor cooperation. The
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since both the bacteria and the plant can survive independently, it is also an example of facultative symbiosis.
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Griffin, A; West, S. A.; Buckling, A. (28 August 2004). "Cooperation and competition in pathogenic bacteria".
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Koeslag, J.H. (1997). "Sex, the
Prisoner's Dilemma Game, and the evolutionary inevitability of cooperation".
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1016:, in which one species lives inside of another, and ectosymbiosis, in which one species lives on another.
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Koeslag, J.H. (2003). "Evolution of cooperation: cooperation defeats defection in the cornfield model".
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3570:"The conditional defector strategies can violate the most crucial supporting mechanisms of cooperation"
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Clutton-Brock, T. (2002). "Breeding together: Kin selection and mutualism in cooperative vertebrates".
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Burtsev, Mikhail; Turchin, Peter (2006). "Evolution of cooperative strategies from first principles".
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show, however, that humans often act more cooperatively than strict self-interest would dictate.
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carry the largest weights, so the size of the handicap is a measure of the animal's quality.
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A male peacock with its beautiful but clumsy, aerodynamically unsound erectile tail, which
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cooperation. There are three major mechanisms that generate this type of fitness benefit:
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Zahavi, Amotz (1977). "The cost of honesty (Further remarks on the handicap principle)".
1321:
1040:
994:
869:
834:
516:
506:
441:
406:
290:
193:
118:
53:
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1938:
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1024:
5113:
5065:
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2390:
2355:
2336:
2275:
2184:
2116:
2012:
1987:
1968:
1613:
1580:
Clutton-Brock, T (5 November 2009). "Cooperation between non-kin in animal societies".
1549:
1394:
1333:
1235:
914:
893:
889:
881:
853:
841:
559:
491:
275:
203:
168:
3399:
3321:
3163:
2457:
1299:, including any cooperative, altruistic and social interactions. Thus, in the example
5231:
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4881:
4829:
4607:
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2245:
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2176:
2108:
2061:
2017:
1960:
1836:
1815:
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1707:
1659:
1655:
1617:
1605:
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1280:
1231:
1134:
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815:
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681:
677:
260:
138:
128:
123:
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3471:
3283:
3206:
2781:
2680:
2644:
BergmĂĽller, Ralph; Johnstone, Rufus A.; Russell, Andrew F.; Bshary, Redouan (2007).
2434:
2279:
2188:
2120:
1222:
displayed by some animals, particularly certain birds, such as, amongst others, the
4747:
4314:
4292:
4100:
3734:
3713:
3693:
3647:
3639:
3589:
3581:
3534:
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3317:
3271:
3194:
3159:
3087:
3041:
2942:
2894:
2884:
2849:
2837:
2769:
2734:
2660:
2618:
2573:
2565:
2538:
2518:
2453:
2450:
Interorganismal signaling in suboptimum environments: The legume-rhizobia symbiosis
2422:
2385:
2375:
2340:
2318:
2310:
2257:
2168:
2100:
2053:
2007:
1999:
1972:
1952:
1902:
1869:
1785:
1742:
1699:
1651:
1597:
1531:
1527:
1366:
1357:
1353:
1252:
1219:
1153:
934:
755:
295:
4592:
3525:
Mesterton-Gibbons, M.; Adams, E.S. (2002). "The
Economics of Animal Cooperation".
2356:"Self-recognition, color signals, and cycles of greenbeard mutualism and altruism"
2229:
Gardner, Andy; Ashleigh S Griffin; Stuart A West (2009). "Theory of
Cooperation".
1889:
Sparkman, A. M.; Adams, J. R.; Steury, T. D.; Waits, L. P.; Murray, D. L. (2011).
1746:
1069:(the mycobiont) and a photosynthetic partner (the photobiont), which is usually a
880:
The second class of explanations for cooperation is indirect fitness benefits, or
5108:
4732:
4559:
4548:
4502:
4445:
4394:
4174:
3978:
2664:
2622:
2057:
1420:
1325:
1272:
830:
759:
722:
248:
238:
100:
2244:
Kummerli, Rolf; Andy
Gardner; Stuart A. West; Ashleigh S. Griffin (April 2009).
1703:
4948:
4820:
4762:
4122:
4117:
4055:
4033:
3876:
3585:
2869:
Proceedings of the National Academy of Sciences of the United States of America
2360:
Proceedings of the National Academy of Sciences of the United States of America
1337:
1317:
1238:. If a signal is to remain reliable, and generally resistant to falsification,
1168:
1164:
1078:
767:
709:
701:
330:
233:
3739:
3722:
3455:
2246:"Limited Dispersal, Budding Dispersal, and Cooperation: An Experimental Study"
1768:
Hamilton, W (1 July 1964). "The genetical evolution of social behaviour. II".
48:
5220:
5150:
5028:
4987:
4833:
4587:
4255:
4250:
2353:
1874:
1857:
1725:
Sherman, P (23 September 1977). "Nepotism and the evolution of alarm calls".
1389:
1384:
1248:
1074:
481:
153:
3643:
3538:
2889:
2738:
2380:
2172:
1956:
1907:
1890:
5145:
5093:
5038:
4871:
4866:
4460:
4218:
3866:
3748:
3705:
3661:
3603:
3546:
3463:
3407:
3198:
2946:
2672:
2646:"Integrating cooperative breeding into theoretical concepts of cooperation"
2630:
2587:
2569:
2530:
2399:
2332:
2271:
2112:
2065:
2021:
2003:
1964:
1754:
1711:
1609:
1227:
1199:
1013:
998:
845:
705:
501:
486:
270:
265:
183:
3434:
Dugatkin, L. A. (2002). "Animal cooperation among unrelated individuals".
3329:
3262:
Langlois, J. H.; Roggman, L. (1990). "Attractive faces are only average".
3185:
Johnstone, RA; Grafen, A (1993). "Dishonesty and the handicap principle".
3171:
3053:
2954:
2908:
2746:
2180:
2104:
1797:
1663:
1634:
Hamilton, W. D. (1964). "The genetical evolution of social behaviour. I".
1218:
There are striking parallels between cooperative behavior and exaggerated
664:
of the actor's social partners. For example, territorial choruses by male
5172:
5043:
4772:
4307:
4137:
4043:
4001:
3910:
3871:
3099:
1344:
1305:
1284:
1149:
1082:
obligate symbiotic associations with green algae, cyanobacteria or both.
1055:
922:
806:
717:
511:
228:
178:
4612:
3697:
3513:
Economics in Nature: Social Dilemmas, Mate Choice and Biological Markets
2314:
1601:
5033:
4934:
4856:
4843:
4477:
4277:
4164:
4142:
4095:
4090:
4038:
4006:
3921:
3900:
3763:
3223:, p. 176, 193. Trustees of the John Voelcker Bird Book Fund, Cape Town.
2426:
1070:
1002:
918:
657:
285:
208:
163:
143:
57:
2558:
Philosophical Transactions of the Royal Society B: Biological Sciences
2228:
5167:
5018:
4627:
4245:
3993:
2841:
2554:"Cooperation beyond the dyad: on simple models and a complex society"
2522:
990:
966:
865:
and eviction often leads to increased stress and decreased survival.
802:
649:
544:
158:
78:
2717:
Axelrod, R.; Hamilton, W.D. (1981). "The evolution of cooperation".
2601:
Wedekind, C. (2000). "Cooperation Through Image Scoring in Humans".
2323:
1043:
is a form of symbiosis in which both participating species benefit.
5070:
4272:
3012:
Zahavi, Amotz (1975). "Mate selection—a selection for a handicap".
2773:
1430:
1425:
1275:
1051:
1047:
1028:
862:
858:
534:
133:
4455:
4169:
3905:
3895:
2795:
Maynard Smith, J. (1989). "Evolution in structured populations".
2594:
2078:
1831:
Maynard Smith, J. (1989). "Evolution in structured populations".
1223:
868:
Enforcement can also be mutually beneficial, and is often called
716:
favors those who achieve the greatest reproductive success while
668:
discourage intruders and are likely to benefit all contributors.
3342:
2643:
1988:"Evidence for competition and cooperation among climbing plants"
1522:
Gardner, Andy; Griffin, Ashleigh; West, Stuart (December 2009).
2141:. Davies, Nicholas B., John R. Krebs, and Stuart A. West. 2012.
1066:
1062:
953:
590:
2799:. Oxford: Oxford University Press. pp. 168–169, 181–183.
2760:
Trivers, R.L. (1971). "The evolution of reciprocal altruism".
2292:
1856:
Balshine-Earn, S.; Neat, F.C.; Reid, H.; Taborsky, M. (1998).
1316:
One of the first references to animal cooperation was made by
849:
suggesting that cooperation is not maintained by reciprocity.
712:, have found the evolution of cooperation fascinating because
4376:
2967:
2204:"Murderous Meerkat Moms Contradict Caring Image, Study Finds"
1855:
971:
926:
673:
3255:
2406:
1194:
974:. When only two partners exist, each can specialize in one
930:
665:
3485:
Alexander, R. (1974). "The evolution of social behavior".
3117:
The handicap principle: a missing piece of Darwin's puzzle
2820:
Godfray, H. C. J. (1992). "The evolution of forgiveness".
2134:
2132:
2130:
1575:
1573:
1571:
1046:
A classic example of mutualism is the interaction between
1285:
prefer mates with the fewest unusual or minority features
1888:
1814:(pp. 70–71). Oxford: Blackwell Scientific Publications.
2127:
1992:
Proceedings of the Royal Society B: Biological Sciences
1568:
1156:, cooperation cannot evolve under these circumstances.
1065:
are another example of mutualism. Lichens consist of a
3524:
3142:
Grafen, A. (1990). "Biological signals as handicaps".
2637:
1835:. Oxford: Oxford University Press. pp. 173–175.
2703:. Oxford: Oxford University Press. pp. 180–206.
2441:
1347:
models, such as the classic cooperative hunting and
1077:. The mycobiont benefits from the sugar products of
660:
that has evolved, at least in part, to increase the
2972:? Cooperative Behavior when the Stakes are Large".
913:This mechanism is similar to what happens with the
3720:
3114:
2150:
1521:
1311:
3721:West, S. A.; Griffin, A. S.; Gardner, A. (2007).
2819:
5218:
3295:
3293:
3219:Hockey, P.A.R., Dean, W.R.J., Ryan, P.G. (2005)
2500:
3261:
3184:
2716:
3675:
2412:
1812:Behavioural Ecology. An Evolutionary Approach.
852:Cooperation is maintained in situations where
3937:
3779:
3617:"Five rules for the evolution of cooperation"
3290:
2794:
2545:
1830:
1681:
1579:
629:
3511:Noë, R., van Hooff, J., and Hammerstein, P.
3336:
2862:
875:
3723:"Evolutionary explanations for cooperation"
3343:Unnikrishnan, M. K.; Akhila, H. S. (2014).
2968:van den Assem; van Dolder; Thaler (2012). "
2920:
2918:
1924:
1824:
3944:
3930:
3786:
3772:
2712:
2710:
2694:
2692:
2690:
2447:
2206:. National Geographic News. Archived from
1985:
1920:
1918:
1629:
1627:
796:
636:
622:
3809:Coloration evidence for natural selection
3738:
3651:
3593:
3484:
3237:The Genetical Theory of Natural Selection
3231:
3229:
3065:
3063:
3035:
2898:
2888:
2577:
2389:
2379:
2322:
2261:
2011:
1946:
1906:
1873:
1677:
1675:
1673:
1100:One example of a hidden benefit involves
5178:Transgenerational epigenetic inheritance
3951:
3793:
3487:Annual Review of Ecology and Systematics
3433:
2915:
2865:"Chaos and the evolution of cooperation"
2600:
1767:
1633:
1257:
1205:
1193:
1133:
1023:
940:
3567:
3424:Ch VIII (The Modern Library, 1859/1958)
3377:
3299:
2924:
2759:
2707:
2698:
2687:
2494:
1915:
1724:
1624:
1370:of how cooperative relationships work.
1012:Two special types of symbiosis include
14:
5219:
3226:
3141:
3112:
3069:
3060:
3011:
2551:
2201:
1670:
1234:rendered evolutionarily stable by his
4941:Dialogues Concerning Natural Religion
3925:
3767:
3614:
2139:An Introduction to Behavioral Ecology
1123:
3499:10.1146/annurev.es.05.110174.001545
3121:. Oxford: Oxford University Press.
2415:Behavioral Ecology and Sociobiology
2083:, do not prefer their own larvae".
24:
4346:Evolutionary developmental biology
3276:10.1111/j.1467-9280.1990.tb00079.x
3252:. Oxford: Oxford University Press.
2452:. Vol. 76. pp. 125–161.
1089:
25:
5243:
3515:Cambridge University Press (2001)
1416:Mutual Aid: A Factor of Evolution
5200:
5191:
5190:
3250:The Evolution of Human Sexuality
3221:Roberts Birds of Southern Africa
2263:10.1111/j.1558-5646.2008.00548.x
959:
737:
603:
602:
589:
47:
5003:Extended evolutionary synthesis
4192:Gene-centered view of evolution
3561:
3518:
3505:
3478:
3427:
3414:
3371:
3242:
3213:
3178:
3135:
3106:
3005:
2961:
2863:Novak, M.; Sigmund, K. (1993).
2856:
2813:
2788:
2753:
2347:
2286:
2237:
2222:
2202:Norris, Scott (15 March 2006).
2195:
2144:
2072:
2028:
1979:
1882:
1849:
1312:History of cooperation research
596:Evolutionary biology portal
5131:Hologenome theory of evolution
4998:History of molecular evolution
4224:Evolutionarily stable strategy
4113:Last universal common ancestor
3380:Journal of Theoretical Biology
3302:Journal of Theoretical Biology
3144:Journal of Theoretical Biology
3072:Journal of Theoretical Biology
3015:Journal of Theoretical Biology
2927:Journal of Theoretical Biology
2448:Zhang, F; Smith, D. L (2002).
1804:
1770:Journal of Theoretical Biology
1761:
1718:
1636:Journal of Theoretical Biology
1532:10.1002/9780470015902.a0021910
1515:
1444:
1363:evolutionarily stable strategy
1230:to suggest that both might be
787:
700:Prominent biologists, such as
555:Creation–evolution controversy
309:History of evolutionary theory
13:
1:
4925:Renaissance and Enlightenment
3400:10.1016/s0022-5193(03)00188-7
3322:10.1016/s0022-5193(05)80297-8
3164:10.1016/S0022-5193(05)80088-8
2458:10.1016/S0065-2113(02)76004-5
2231:Encyclopedia of Life Sciences
1747:10.1126/science.197.4310.1246
1509:
687:
5136:Missing heritability problem
4763:Gamete differentiation/sexes
3092:10.1016/0022-5193(77)90061-3
3046:10.1016/0022-5193(75)90111-3
2665:10.1016/j.beproc.2007.07.001
2623:10.1126/science.288.5467.850
2058:10.1126/science.291.5503.478
1790:10.1016/0022-5193(64)90039-6
1656:10.1016/0022-5193(64)90038-4
1365:is a refined version of the
1019:
985:
773:
540:Evolution as fact and theory
7:
3355:: 1410–1414. Archived from
2762:Quarterly Review of Biology
1704:10.1126/science.296.5565.69
1377:
1050:soil bacteria and legumes (
825:, another term to describe
10:
5248:
4768:Life cycles/nuclear phases
4320:Trivers–Willard hypothesis
3586:10.1038/s41598-022-18797-2
1986:Biernaskie, J. M. (2011).
1127:
944:
575:Nature-nurture controversy
5186:
5086:
5011:
4915:
4842:
4798:
4653:
4557:
4374:
4333:
4266:Parent–offspring conflict
4202:
4071:Earliest known life forms
3992:
3959:
3849:
3801:
3740:10.1016/j.cub.2007.06.004
3456:10.1007/s00114-002-0379-y
876:Indirect fitness benefits
462:Evolutionary neuroscience
437:Evolutionary epistemology
417:Evolutionary anthropology
397:Applications of evolution
5119:Cultural group selection
4983:The eclipse of Darwinism
4955:On the Origin of Species
4930:Transmutation of species
3862:Anti-predator adaptation
1437:
1400:Evolution of cooperation
1289:African pygmy kingfisher
1264:African pygmy kingfisher
452:Evolutionary linguistics
447:Evolutionary game theory
422:Evolutionary computation
18:Co-operation (evolution)
5124:Dual inheritance theory
4963:History of paleontology
3836:Paradox of the plankton
3644:10.1126/science.1133755
3568:Ibrahim, Ahmed (2022).
3539:10.1126/science.1080051
2890:10.1073/pnas.90.11.5091
2739:10.1126/science.7466396
2381:10.1073/pnas.0510260103
2173:10.1126/science.7652573
1957:10.1126/science.1089402
840:It has been shown that
797:Direct fitness benefits
565:Objections to evolution
472:Evolutionary psychology
467:Evolutionary physiology
412:Evolutionary aesthetics
391:Fields and applications
373:History of paleontology
4812:Punctuated equilibrium
4133:Non-adaptive radiation
4081:Evolutionary arms race
3199:10.1006/anbe.1993.1253
3113:Zahavi, Amotz (1997).
2993:Cite journal requires
2947:10.1006/jtbi.1997.0496
2570:10.1098/rstb.2010.0150
2552:Connor, R. C. (2010).
2004:10.1098/rspb.2010.1771
1875:10.1093/beheco/9.5.432
1405:Management cybernetics
1267:
1215:
1203:
1180:experimental economics
1140:
1037:
903:Pseudomonas aeruginosa
870:reciprocal cooperation
829:, is a term coined by
732:altruistic cooperation
497:Speciation experiments
477:Experimental evolution
432:Evolutionary economics
254:Recent human evolution
112:Processes and outcomes
5104:Evolutionary medicine
4978:Mendelian inheritance
4686:Biological complexity
4674:Programmed cell death
4366:Phenotypic plasticity
4086:Evolutionary pressure
4076:Evidence of evolution
3974:Timeline of evolution
3802:Patterns of evolution
3422:The Origin of Species
3264:Psychological Science
2797:Evolutionary Genetics
2653:Behavioural Processes
2105:10.1007/s001140050718
1908:10.1093/beheco/arq194
1833:Evolutionary Genetics
1524:Theory of Cooperation
1410:Microbial Cooperation
1297:evolutionarily stable
1261:
1210:The best horses in a
1209:
1197:
1173:evolutionarily stable
1137:
1027:
947:Multi-level selection
941:Multi-level selection
457:Evolutionary medicine
402:Biosocial criminology
368:History of speciation
281:Evolutionary taxonomy
244:Timeline of evolution
5227:Evolutionary biology
5078:Teleology in biology
4973:Blending inheritance
4351:Genetic assimilation
4214:Artificial selection
3953:Evolutionary biology
3814:Convergent evolution
3795:Evolutionary ecology
2699:Sigmund, K. (1993).
1343:In the past, simple
662:reproductive success
427:Evolutionary ecology
41:Evolutionary biology
5141:Molecular evolution
5099:Ecological genetics
4968:Transitional fossil
4758:Sexual reproduction
4598:endomembrane system
4527:pollinator-mediated
4483:dolphins and whales
4261:Parental investment
3887:Distraction display
3831:Divergent evolution
3698:10.1038/nature04470
3690:2006Natur.440.1041B
3684:(7087): 1041–1044.
3636:2006Sci...314.1560N
3630:(5805): 1560–1563.
3448:2002NW.....89..533D
3436:Naturwissenschaften
3392:2003JThBi.224..399K
3314:1990JThBi.144...15K
3156:1990JThBi.144..517G
3084:1977JThBi..67..603Z
3028:1975JThBi..53..205Z
2939:1997JThBi.189...53K
2881:1993PNAS...90.5091N
2834:1992Natur.355..206G
2731:1981Sci...211.1390A
2725:(4489): 1390–1396.
2615:2000Sci...288..850W
2564:(1553): 2687–2697.
2515:2001Natur.411..937L
2372:2006PNAS..103.7372S
2315:10.1038/nature02744
2307:2004Natur.430.1024G
2301:(7003): 1024–1027.
2165:1995Sci...269.1260H
2159:(5228): 1260–1262.
2097:2000NW.....87..266S
2085:Naturwissenschaften
2050:2001Sci...291..478C
1939:2003Sci...302..634G
1782:1964JThBi...7...17H
1739:1977Sci...197.1246S
1733:(4310): 1246–1253.
1696:2002Sci...296...69C
1648:1964JThBi...7....1H
1602:10.1038/nature08366
1594:2009Natur.462...51C
1322:Thomas Henry Huxley
935:side-blotch lizards
933:. An example is in
835:cooperative hunting
529:Social implications
517:Universal Darwinism
507:Island biogeography
442:Evolutionary ethics
407:Ecological genetics
353:Molecular evolution
291:Transitional fossil
119:Population genetics
35:Part of a series on
5114:Cultural evolution
4229:Fisher's principle
4158:Handicap principle
4148:Parallel evolution
4012:Adaptive radiation
3882:Deimatic behaviour
3841:Predator satiation
3826:Parallel evolution
3615:Nowak, MA (2006).
3574:Scientific Reports
3248:Symons, D. (1979)
2427:10.1007/BF00167053
1895:Behavioral Ecology
1862:Behavioral Ecology
1349:Prisoner's dilemma
1268:
1236:handicap principle
1216:
1204:
1145:prisoner's dilemma
1141:
1130:Prisoner's dilemma
1124:Prisoner's dilemma
1038:
915:green-beard effect
910:actor might give.
894:green-beard effect
890:kin discrimination
827:by-product benefit
823:Prisoner's Delight
803:by-product benefit
560:Theistic evolution
492:Selective breeding
204:Parallel evolution
169:Adaptive radiation
5214:
5213:
4830:Uniformitarianism
4783:Sex-determination
4288:Sexual dimorphism
4283:Natural selection
4187:Unit of selection
4153:Signalling theory
3919:
3918:
3857:Signalling theory
3733:(16): R661–R672.
3235:Fisher R. (1930)
3128:978-0-19-510035-8
2875:(11): 5091–5094.
2828:(6357): 206–207.
2609:(5467): 850–852.
2509:(6840): 937–940.
2366:(19): 7372–7377.
2081:Polistes carolina
2044:(5503): 478–481.
1998:(1714): 1989–96.
1933:(5645): 634–636.
1281:natural selection
1159:However, in 1981
1106:superb fairy-wren
1034:Vigna unguiculata
886:limited dispersal
816:Polistes dominula
781:Ipomoea hederacea
764:Australian magpie
728:inclusive fitness
714:natural selection
682:African elephants
646:
645:
337:Origin of Species
139:Natural selection
16:(Redirected from
5239:
5204:
5194:
5193:
4993:Modern synthesis
4753:Multicellularity
4748:Mosaic evolution
4633:auditory ossicle
4315:Social selection
4298:Flowering plants
4293:Sexual selection
3946:
3939:
3932:
3923:
3922:
3788:
3781:
3774:
3765:
3764:
3760:
3742:
3717:
3672:
3670:
3664:. Archived from
3655:
3621:
3608:
3607:
3597:
3565:
3559:
3558:
3533:(5601): 2146–7.
3522:
3516:
3509:
3503:
3502:
3482:
3476:
3475:
3431:
3425:
3418:
3412:
3411:
3375:
3369:
3368:
3366:
3364:
3340:
3334:
3333:
3297:
3288:
3287:
3259:
3253:
3246:
3240:
3233:
3224:
3217:
3211:
3210:
3187:Animal Behaviour
3182:
3176:
3175:
3139:
3133:
3132:
3120:
3110:
3104:
3103:
3067:
3058:
3057:
3039:
3009:
3003:
3002:
2996:
2991:
2989:
2981:
2965:
2959:
2958:
2922:
2913:
2912:
2902:
2892:
2860:
2854:
2853:
2842:10.1038/355206a0
2817:
2811:
2810:
2792:
2786:
2785:
2757:
2751:
2750:
2714:
2705:
2704:
2696:
2685:
2684:
2650:
2641:
2635:
2634:
2598:
2592:
2591:
2581:
2549:
2543:
2542:
2523:10.1038/35082053
2498:
2492:
2491:
2485:
2481:
2479:
2471:
2445:
2439:
2438:
2410:
2404:
2403:
2393:
2383:
2351:
2345:
2344:
2326:
2290:
2284:
2283:
2265:
2241:
2235:
2234:
2226:
2220:
2219:
2217:
2215:
2210:on 16 March 2006
2199:
2193:
2192:
2148:
2142:
2136:
2125:
2124:
2076:
2070:
2069:
2032:
2026:
2025:
2015:
1983:
1977:
1976:
1950:
1922:
1913:
1912:
1910:
1886:
1880:
1879:
1877:
1853:
1847:
1846:
1828:
1822:
1808:
1802:
1801:
1765:
1759:
1758:
1722:
1716:
1715:
1679:
1668:
1667:
1631:
1622:
1621:
1577:
1566:
1565:
1559:
1555:
1553:
1545:
1519:
1503:
1448:
1367:Nash equilibrium
1358:Nash equilibrium
1354:Nash equilibrium
1253:sexual selection
1220:sexual ornaments
1154:Nash equilibrium
756:western bluebird
638:
631:
624:
611:
606:
605:
598:
594:
593:
570:Level of support
363:Current research
348:Modern synthesis
343:Before synthesis
296:Extinction event
54:Darwin's finches
51:
32:
31:
21:
5247:
5246:
5242:
5241:
5240:
5238:
5237:
5236:
5217:
5216:
5215:
5210:
5182:
5109:Group selection
5082:
5007:
4911:
4838:
4800:Tempo and modes
4794:
4649:
4553:
4370:
4329:
4205:
4198:
4175:Species complex
3988:
3979:History of life
3955:
3950:
3920:
3915:
3845:
3797:
3792:
3727:Current Biology
3671:on 18 May 2011.
3668:
3619:
3611:
3566:
3562:
3523:
3519:
3510:
3506:
3483:
3479:
3432:
3428:
3419:
3415:
3376:
3372:
3362:
3360:
3359:on 4 March 2016
3341:
3337:
3298:
3291:
3260:
3256:
3247:
3243:
3234:
3227:
3218:
3214:
3183:
3179:
3140:
3136:
3129:
3111:
3107:
3068:
3061:
3037:10.1.1.586.3819
3010:
3006:
2994:
2992:
2983:
2982:
2966:
2962:
2923:
2916:
2861:
2857:
2818:
2814:
2807:
2793:
2789:
2758:
2754:
2715:
2708:
2697:
2688:
2648:
2642:
2638:
2599:
2595:
2550:
2546:
2499:
2495:
2483:
2482:
2473:
2472:
2468:
2446:
2442:
2411:
2407:
2352:
2348:
2291:
2287:
2242:
2238:
2227:
2223:
2213:
2211:
2200:
2196:
2149:
2145:
2137:
2128:
2077:
2073:
2033:
2029:
1984:
1980:
1923:
1916:
1887:
1883:
1854:
1850:
1843:
1829:
1825:
1809:
1805:
1766:
1762:
1723:
1719:
1690:(5565): 69–72.
1680:
1671:
1632:
1625:
1588:(7269): 51–57.
1578:
1569:
1557:
1556:
1547:
1546:
1542:
1520:
1516:
1512:
1507:
1506:
1501:
1497:
1493:
1489:
1485:
1481:
1477:
1473:
1469:
1465:
1461:
1457:
1453:
1449:
1445:
1440:
1435:
1421:Peter Kropotkin
1395:Dunbar's number
1380:
1326:Peter Kropotkin
1314:
1232:fitness signals
1188:
1132:
1126:
1102:Malarus cyaneus
1095:Hidden benefits
1092:
1090:Hidden benefits
1022:
988:
962:
949:
943:
878:
831:Kenneth Binmore
799:
790:
776:
760:pied kingfisher
740:
723:ground squirrel
690:
642:
601:
588:
587:
580:
579:
530:
522:
521:
392:
384:
383:
382:
310:
302:
301:
300:
249:Human evolution
239:History of life
223:
222:Natural history
215:
214:
213:
113:
105:
60:
28:
23:
22:
15:
12:
11:
5:
5245:
5235:
5234:
5229:
5212:
5211:
5209:
5208:
5198:
5187:
5184:
5183:
5181:
5180:
5175:
5170:
5165:
5160:
5159:
5158:
5148:
5143:
5138:
5133:
5128:
5127:
5126:
5121:
5116:
5106:
5101:
5096:
5090:
5088:
5084:
5083:
5081:
5080:
5075:
5074:
5073:
5068:
5063:
5062:
5061:
5051:
5046:
5041:
5036:
5031:
5021:
5015:
5013:
5009:
5008:
5006:
5005:
5000:
4995:
4990:
4985:
4980:
4975:
4970:
4965:
4960:
4959:
4958:
4949:Charles Darwin
4946:
4945:
4944:
4932:
4927:
4921:
4919:
4913:
4912:
4910:
4909:
4904:
4899:
4894:
4889:
4887:Non-ecological
4884:
4879:
4874:
4869:
4864:
4859:
4854:
4848:
4846:
4840:
4839:
4837:
4836:
4827:
4818:
4804:
4802:
4796:
4795:
4793:
4792:
4787:
4786:
4785:
4780:
4775:
4770:
4765:
4755:
4750:
4745:
4740:
4735:
4730:
4725:
4720:
4715:
4710:
4705:
4704:
4703:
4693:
4688:
4683:
4678:
4677:
4676:
4671:
4660:
4658:
4651:
4650:
4648:
4647:
4646:
4645:
4640:
4638:nervous system
4635:
4630:
4625:
4617:
4616:
4615:
4610:
4605:
4600:
4595:
4590:
4580:
4575:
4570:
4564:
4562:
4555:
4554:
4552:
4551:
4546:
4541:
4536:
4531:
4530:
4529:
4519:
4518:
4517:
4512:
4511:
4510:
4505:
4495:
4490:
4485:
4480:
4475:
4474:
4473:
4468:
4458:
4448:
4443:
4442:
4441:
4431:
4426:
4421:
4416:
4415:
4414:
4404:
4399:
4398:
4397:
4387:
4381:
4379:
4372:
4371:
4369:
4368:
4363:
4358:
4353:
4348:
4343:
4337:
4335:
4331:
4330:
4328:
4327:
4322:
4317:
4312:
4311:
4310:
4305:
4300:
4290:
4285:
4280:
4275:
4270:
4269:
4268:
4263:
4253:
4248:
4243:
4242:
4241:
4231:
4226:
4221:
4216:
4210:
4208:
4200:
4199:
4197:
4196:
4195:
4194:
4184:
4179:
4178:
4177:
4172:
4162:
4161:
4160:
4150:
4145:
4140:
4138:Origin of life
4135:
4130:
4125:
4123:Microevolution
4120:
4118:Macroevolution
4115:
4110:
4105:
4104:
4103:
4093:
4088:
4083:
4078:
4073:
4068:
4063:
4058:
4056:Common descent
4053:
4052:
4051:
4041:
4036:
4034:Baldwin effect
4031:
4030:
4029:
4024:
4014:
4009:
4004:
3998:
3996:
3990:
3989:
3987:
3986:
3981:
3976:
3971:
3966:
3960:
3957:
3956:
3949:
3948:
3941:
3934:
3926:
3917:
3916:
3914:
3913:
3908:
3903:
3898:
3893:
3891:
3890:
3889:
3884:
3879:
3877:Apparent death
3874:
3869:
3859:
3853:
3851:
3847:
3846:
3844:
3843:
3838:
3833:
3828:
3823:
3822:
3821:
3811:
3805:
3803:
3799:
3798:
3791:
3790:
3783:
3776:
3768:
3762:
3761:
3718:
3673:
3610:
3609:
3560:
3517:
3504:
3477:
3442:(12): 533–41.
3426:
3413:
3386:(3): 399–410.
3370:
3335:
3289:
3270:(2): 115–121.
3254:
3241:
3225:
3212:
3193:(4): 759–764.
3177:
3150:(4): 517–546.
3134:
3127:
3105:
3078:(3): 603–605.
3059:
3022:(1): 205–214.
3004:
2995:|journal=
2970:Split or Steal
2960:
2914:
2855:
2812:
2806:978-0198542155
2805:
2787:
2774:10.1086/406755
2752:
2706:
2686:
2636:
2593:
2544:
2493:
2484:|journal=
2466:
2440:
2405:
2346:
2285:
2256:(4): 939–949.
2236:
2221:
2194:
2143:
2126:
2091:(6): 266–269.
2071:
2027:
1978:
1948:10.1.1.561.819
1914:
1901:(1): 199–205.
1881:
1868:(5): 432–438.
1848:
1842:978-0198542155
1841:
1823:
1803:
1760:
1717:
1669:
1623:
1567:
1558:|journal=
1541:978-0470016176
1540:
1513:
1511:
1508:
1505:
1504:
1499:
1495:
1491:
1487:
1483:
1479:
1475:
1471:
1467:
1463:
1459:
1455:
1451:
1442:
1441:
1439:
1436:
1434:
1433:
1428:
1423:
1412:
1407:
1402:
1397:
1392:
1387:
1381:
1379:
1376:
1338:brood parasite
1318:Charles Darwin
1313:
1310:
1293:Ispidina picta
1187:
1184:
1169:Robert Trivers
1128:Main article:
1125:
1122:
1091:
1088:
1079:photosynthesis
1021:
1018:
987:
984:
961:
958:
945:Main article:
942:
939:
877:
874:
798:
795:
789:
786:
775:
772:
768:dwarf mongoose
739:
736:
710:W. D. Hamilton
702:Charles Darwin
689:
686:
676:herds to pied
644:
643:
641:
640:
633:
626:
618:
615:
614:
613:
612:
599:
582:
581:
578:
577:
572:
567:
562:
557:
552:
550:Social effects
547:
542:
537:
531:
528:
527:
524:
523:
520:
519:
514:
509:
504:
499:
494:
489:
484:
479:
474:
469:
464:
459:
454:
449:
444:
439:
434:
429:
424:
419:
414:
409:
404:
399:
393:
390:
389:
386:
385:
381:
380:
370:
365:
360:
355:
350:
345:
340:
333:
328:
323:
318:
312:
311:
308:
307:
304:
303:
299:
298:
293:
288:
283:
278:
276:Classification
273:
268:
263:
258:
257:
256:
246:
241:
236:
234:Common descent
231:
229:Origin of life
225:
224:
221:
220:
217:
216:
212:
211:
206:
201:
196:
191:
186:
181:
176:
171:
166:
161:
156:
151:
146:
141:
136:
131:
126:
121:
115:
114:
111:
110:
107:
106:
104:
103:
98:
93:
87:
86:
81:
76:
71:
65:
62:
61:
52:
44:
43:
37:
36:
26:
9:
6:
4:
3:
2:
5244:
5233:
5230:
5228:
5225:
5224:
5222:
5207:
5203:
5199:
5197:
5189:
5188:
5185:
5179:
5176:
5174:
5171:
5169:
5166:
5164:
5161:
5157:
5154:
5153:
5152:
5151:Phylogenetics
5149:
5147:
5144:
5142:
5139:
5137:
5134:
5132:
5129:
5125:
5122:
5120:
5117:
5115:
5112:
5111:
5110:
5107:
5105:
5102:
5100:
5097:
5095:
5092:
5091:
5089:
5085:
5079:
5076:
5072:
5069:
5067:
5064:
5060:
5057:
5056:
5055:
5054:Structuralism
5052:
5050:
5047:
5045:
5042:
5040:
5037:
5035:
5032:
5030:
5029:Catastrophism
5027:
5026:
5025:
5022:
5020:
5017:
5016:
5014:
5010:
5004:
5001:
4999:
4996:
4994:
4991:
4989:
4988:Neo-Darwinism
4986:
4984:
4981:
4979:
4976:
4974:
4971:
4969:
4966:
4964:
4961:
4957:
4956:
4952:
4951:
4950:
4947:
4943:
4942:
4938:
4937:
4936:
4933:
4931:
4928:
4926:
4923:
4922:
4920:
4918:
4914:
4908:
4905:
4903:
4902:Reinforcement
4900:
4898:
4895:
4893:
4890:
4888:
4885:
4883:
4880:
4878:
4875:
4873:
4870:
4868:
4865:
4863:
4860:
4858:
4855:
4853:
4850:
4849:
4847:
4845:
4841:
4835:
4834:Catastrophism
4831:
4828:
4826:
4825:Macromutation
4822:
4821:Micromutation
4819:
4817:
4813:
4809:
4806:
4805:
4803:
4801:
4797:
4791:
4788:
4784:
4781:
4779:
4776:
4774:
4771:
4769:
4766:
4764:
4761:
4760:
4759:
4756:
4754:
4751:
4749:
4746:
4744:
4741:
4739:
4736:
4734:
4731:
4729:
4728:Immune system
4726:
4724:
4721:
4719:
4716:
4714:
4711:
4709:
4706:
4702:
4699:
4698:
4697:
4694:
4692:
4689:
4687:
4684:
4682:
4679:
4675:
4672:
4670:
4667:
4666:
4665:
4662:
4661:
4659:
4657:
4652:
4644:
4641:
4639:
4636:
4634:
4631:
4629:
4626:
4624:
4621:
4620:
4618:
4614:
4611:
4609:
4606:
4604:
4601:
4599:
4596:
4594:
4591:
4589:
4588:symbiogenesis
4586:
4585:
4584:
4581:
4579:
4576:
4574:
4571:
4569:
4566:
4565:
4563:
4561:
4556:
4550:
4547:
4545:
4542:
4540:
4537:
4535:
4532:
4528:
4525:
4524:
4523:
4520:
4516:
4513:
4509:
4506:
4504:
4501:
4500:
4499:
4496:
4494:
4491:
4489:
4486:
4484:
4481:
4479:
4476:
4472:
4469:
4467:
4464:
4463:
4462:
4459:
4457:
4454:
4453:
4452:
4449:
4447:
4444:
4440:
4437:
4436:
4435:
4432:
4430:
4427:
4425:
4422:
4420:
4417:
4413:
4410:
4409:
4408:
4405:
4403:
4400:
4396:
4393:
4392:
4391:
4388:
4386:
4383:
4382:
4380:
4378:
4373:
4367:
4364:
4362:
4359:
4357:
4354:
4352:
4349:
4347:
4344:
4342:
4339:
4338:
4336:
4332:
4326:
4323:
4321:
4318:
4316:
4313:
4309:
4306:
4304:
4301:
4299:
4296:
4295:
4294:
4291:
4289:
4286:
4284:
4281:
4279:
4276:
4274:
4271:
4267:
4264:
4262:
4259:
4258:
4257:
4256:Kin selection
4254:
4252:
4251:Genetic drift
4249:
4247:
4244:
4240:
4237:
4236:
4235:
4232:
4230:
4227:
4225:
4222:
4220:
4217:
4215:
4212:
4211:
4209:
4207:
4201:
4193:
4190:
4189:
4188:
4185:
4183:
4180:
4176:
4173:
4171:
4168:
4167:
4166:
4163:
4159:
4156:
4155:
4154:
4151:
4149:
4146:
4144:
4141:
4139:
4136:
4134:
4131:
4129:
4126:
4124:
4121:
4119:
4116:
4114:
4111:
4109:
4106:
4102:
4099:
4098:
4097:
4094:
4092:
4089:
4087:
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4082:
4079:
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4064:
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4059:
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4047:
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4045:
4042:
4040:
4037:
4035:
4032:
4028:
4025:
4023:
4020:
4019:
4018:
4015:
4013:
4010:
4008:
4005:
4003:
4000:
3999:
3997:
3995:
3991:
3985:
3982:
3980:
3977:
3975:
3972:
3970:
3967:
3965:
3962:
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3902:
3899:
3897:
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3883:
3880:
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3875:
3873:
3870:
3868:
3865:
3864:
3863:
3860:
3858:
3855:
3854:
3852:
3848:
3842:
3839:
3837:
3834:
3832:
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3827:
3824:
3820:
3817:
3816:
3815:
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3810:
3807:
3806:
3804:
3800:
3796:
3789:
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3777:
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3766:
3758:
3754:
3750:
3746:
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3736:
3732:
3728:
3724:
3719:
3715:
3711:
3707:
3703:
3699:
3695:
3691:
3687:
3683:
3679:
3674:
3667:
3663:
3659:
3654:
3649:
3645:
3641:
3637:
3633:
3629:
3625:
3618:
3613:
3612:
3605:
3601:
3596:
3591:
3587:
3583:
3579:
3575:
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3556:
3552:
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3540:
3536:
3532:
3528:
3521:
3514:
3508:
3500:
3496:
3492:
3488:
3481:
3473:
3469:
3465:
3461:
3457:
3453:
3449:
3445:
3441:
3437:
3430:
3423:
3417:
3409:
3405:
3401:
3397:
3393:
3389:
3385:
3381:
3374:
3358:
3354:
3350:
3346:
3339:
3331:
3327:
3323:
3319:
3315:
3311:
3307:
3303:
3296:
3294:
3285:
3281:
3277:
3273:
3269:
3265:
3258:
3251:
3245:
3238:
3232:
3230:
3222:
3216:
3208:
3204:
3200:
3196:
3192:
3188:
3181:
3173:
3169:
3165:
3161:
3157:
3153:
3149:
3145:
3138:
3130:
3124:
3119:
3118:
3109:
3101:
3097:
3093:
3089:
3085:
3081:
3077:
3073:
3066:
3064:
3055:
3051:
3047:
3043:
3038:
3033:
3029:
3025:
3021:
3017:
3016:
3008:
3000:
2987:
2979:
2975:
2971:
2964:
2956:
2952:
2948:
2944:
2940:
2936:
2932:
2928:
2921:
2919:
2910:
2906:
2901:
2896:
2891:
2886:
2882:
2878:
2874:
2870:
2866:
2859:
2851:
2847:
2843:
2839:
2835:
2831:
2827:
2823:
2816:
2808:
2802:
2798:
2791:
2783:
2779:
2775:
2771:
2767:
2763:
2756:
2748:
2744:
2740:
2736:
2732:
2728:
2724:
2720:
2713:
2711:
2702:
2701:Games of Life
2695:
2693:
2691:
2682:
2678:
2674:
2670:
2666:
2662:
2658:
2654:
2647:
2640:
2632:
2628:
2624:
2620:
2616:
2612:
2608:
2604:
2597:
2589:
2585:
2580:
2575:
2571:
2567:
2563:
2559:
2555:
2548:
2540:
2536:
2532:
2528:
2524:
2520:
2516:
2512:
2508:
2504:
2497:
2489:
2477:
2469:
2467:9780120007943
2463:
2459:
2455:
2451:
2444:
2436:
2432:
2428:
2424:
2420:
2416:
2409:
2401:
2397:
2392:
2387:
2382:
2377:
2373:
2369:
2365:
2361:
2357:
2350:
2342:
2338:
2334:
2330:
2325:
2320:
2316:
2312:
2308:
2304:
2300:
2296:
2289:
2281:
2277:
2273:
2269:
2264:
2259:
2255:
2251:
2247:
2240:
2232:
2225:
2209:
2205:
2198:
2190:
2186:
2182:
2178:
2174:
2170:
2166:
2162:
2158:
2154:
2147:
2140:
2135:
2133:
2131:
2122:
2118:
2114:
2110:
2106:
2102:
2098:
2094:
2090:
2086:
2082:
2075:
2067:
2063:
2059:
2055:
2051:
2047:
2043:
2039:
2031:
2023:
2019:
2014:
2009:
2005:
2001:
1997:
1993:
1989:
1982:
1974:
1970:
1966:
1962:
1958:
1954:
1949:
1944:
1940:
1936:
1932:
1928:
1921:
1919:
1909:
1904:
1900:
1896:
1892:
1885:
1876:
1871:
1867:
1863:
1859:
1852:
1844:
1838:
1834:
1827:
1821:
1817:
1813:
1807:
1799:
1795:
1791:
1787:
1783:
1779:
1775:
1771:
1764:
1756:
1752:
1748:
1744:
1740:
1736:
1732:
1728:
1721:
1713:
1709:
1705:
1701:
1697:
1693:
1689:
1685:
1678:
1676:
1674:
1665:
1661:
1657:
1653:
1649:
1645:
1641:
1637:
1630:
1628:
1619:
1615:
1611:
1607:
1603:
1599:
1595:
1591:
1587:
1583:
1576:
1574:
1572:
1563:
1551:
1543:
1537:
1533:
1529:
1525:
1518:
1514:
1447:
1443:
1432:
1429:
1427:
1424:
1422:
1418:
1417:
1413:
1411:
1408:
1406:
1403:
1401:
1398:
1396:
1393:
1391:
1390:Collaboration
1388:
1386:
1385:Agreeableness
1383:
1382:
1375:
1371:
1368:
1364:
1359:
1355:
1350:
1346:
1341:
1339:
1335:
1334:kin selection
1329:
1327:
1323:
1319:
1309:
1307:
1302:
1298:
1294:
1290:
1286:
1282:
1277:
1274:
1265:
1260:
1256:
1254:
1250:
1246:
1245:handicap race
1241:
1237:
1233:
1229:
1225:
1221:
1213:
1212:handicap race
1208:
1201:
1196:
1192:
1183:
1181:
1178:Results from
1176:
1174:
1170:
1166:
1162:
1157:
1155:
1151:
1146:
1136:
1131:
1121:
1117:
1113:
1111:
1107:
1103:
1098:
1096:
1087:
1083:
1080:
1076:
1075:cyanobacteria
1072:
1068:
1064:
1060:
1057:
1053:
1049:
1044:
1042:
1036:
1035:
1030:
1026:
1017:
1015:
1014:endosymbiosis
1010:
1006:
1004:
1000:
996:
992:
983:
979:
977:
973:
968:
960:Market effect
957:
955:
948:
938:
936:
932:
928:
924:
920:
916:
911:
907:
905:
904:
897:
895:
891:
887:
883:
873:
871:
866:
864:
860:
855:
850:
847:
843:
838:
836:
832:
828:
824:
820:
818:
817:
810:
808:
804:
794:
785:
783:
782:
771:
769:
765:
761:
757:
751:
747:
743:
738:Kin selection
735:
733:
729:
724:
719:
715:
711:
707:
703:
698:
694:
685:
683:
679:
675:
669:
667:
663:
659:
655:
651:
639:
634:
632:
627:
625:
620:
619:
617:
616:
610:
600:
597:
592:
586:
585:
584:
583:
576:
573:
571:
568:
566:
563:
561:
558:
556:
553:
551:
548:
546:
543:
541:
538:
536:
533:
532:
526:
525:
518:
515:
513:
510:
508:
505:
503:
500:
498:
495:
493:
490:
488:
485:
483:
482:Phylogenetics
480:
478:
475:
473:
470:
468:
465:
463:
460:
458:
455:
453:
450:
448:
445:
443:
440:
438:
435:
433:
430:
428:
425:
423:
420:
418:
415:
413:
410:
408:
405:
403:
400:
398:
395:
394:
388:
387:
378:
374:
371:
369:
366:
364:
361:
359:
356:
354:
351:
349:
346:
344:
341:
339:
338:
334:
332:
329:
327:
326:Before Darwin
324:
322:
319:
317:
314:
313:
306:
305:
297:
294:
292:
289:
287:
284:
282:
279:
277:
274:
272:
269:
267:
264:
262:
259:
255:
252:
251:
250:
247:
245:
242:
240:
237:
235:
232:
230:
227:
226:
219:
218:
210:
207:
205:
202:
200:
197:
195:
192:
190:
187:
185:
182:
180:
177:
175:
172:
170:
167:
165:
162:
160:
157:
155:
154:Genetic drift
152:
150:
147:
145:
142:
140:
137:
135:
132:
130:
127:
125:
122:
120:
117:
116:
109:
108:
102:
99:
97:
94:
92:
89:
88:
85:
82:
80:
77:
75:
72:
70:
67:
66:
64:
63:
59:
55:
50:
46:
45:
42:
39:
38:
34:
33:
30:
19:
5163:Polymorphism
5146:Astrobiology
5094:Biogeography
5049:Saltationism
5039:Orthogenesis
5024:Alternatives
4953:
4939:
4872:Cospeciation
4867:Cladogenesis
4816:Saltationism
4773:Mating types
4696:Color vision
4690:
4681:Avian flight
4603:mitochondria
4341:Canalisation
4219:Biodiversity
3964:Introduction
3867:Alarm signal
3730:
3726:
3681:
3677:
3666:the original
3627:
3623:
3580:(1): 15157.
3577:
3573:
3563:
3530:
3526:
3520:
3512:
3507:
3490:
3486:
3480:
3439:
3435:
3429:
3421:
3416:
3383:
3379:
3373:
3361:. Retrieved
3357:the original
3352:
3348:
3338:
3308:(1): 15–35.
3305:
3301:
3267:
3263:
3257:
3249:
3244:
3236:
3220:
3215:
3190:
3186:
3180:
3147:
3143:
3137:
3116:
3108:
3075:
3071:
3019:
3013:
3007:
2986:cite journal
2969:
2963:
2933:(1): 53–61.
2930:
2926:
2872:
2868:
2858:
2825:
2821:
2815:
2796:
2790:
2765:
2761:
2755:
2722:
2718:
2700:
2659:(2): 61–72.
2656:
2652:
2639:
2606:
2602:
2596:
2561:
2557:
2547:
2506:
2502:
2496:
2449:
2443:
2418:
2414:
2408:
2363:
2359:
2349:
2298:
2294:
2288:
2253:
2249:
2239:
2230:
2224:
2212:. Retrieved
2208:the original
2197:
2156:
2152:
2146:
2138:
2088:
2084:
2080:
2074:
2041:
2037:
2030:
1995:
1991:
1981:
1930:
1926:
1898:
1894:
1884:
1865:
1861:
1851:
1832:
1826:
1811:
1806:
1776:(1): 17–52.
1773:
1769:
1763:
1730:
1726:
1720:
1687:
1683:
1639:
1635:
1585:
1581:
1523:
1517:
1446:
1414:
1372:
1342:
1330:
1315:
1292:
1271:caused by a
1269:
1249:amateur golf
1228:Amotz Zahavi
1217:
1200:Amotz Zahavi
1189:
1177:
1158:
1142:
1118:
1114:
1109:
1101:
1099:
1094:
1093:
1084:
1061:
1045:
1039:
1032:
1011:
1007:
999:commensalism
989:
980:
963:
950:
912:
908:
901:
898:
879:
867:
851:
846:female lions
839:
826:
822:
821:
814:
811:
800:
791:
779:
777:
752:
748:
744:
741:
706:E. O. Wilson
699:
695:
691:
670:
653:
647:
502:Sociobiology
487:Paleontology
335:
271:Biogeography
266:Biodiversity
184:Coextinction
174:Co-operation
173:
149:Polymorphism
74:Introduction
29:
5173:Systematics
5044:Mutationism
4862:Catagenesis
4790:Snake venom
4723:Eusociality
4701:in primates
4691:Cooperation
4619:In animals
4439:butterflies
4412:Cephalopods
4402:Brachiopods
4334:Development
4308:Mate choice
4061:Convergence
4044:Coevolution
4002:Abiogenesis
3911:Unkenreflex
3872:Aposematism
3420:Darwin, C.
2214:20 November
1642:(1): 1–16.
1345:game theory
1306:koinophilia
1056:nitrogenase
1031:nodules on
923:slime molds
854:free-riding
842:free riding
807:enforcement
788:Explanation
718:cooperative
654:cooperation
512:Systematics
321:Renaissance
199:Convergence
189:Contingency
179:Coevolution
5221:Categories
5034:Lamarckism
5012:Philosophy
4935:David Hume
4897:Peripatric
4892:Parapatric
4877:Ecological
4857:Anagenesis
4852:Allopatric
4844:Speciation
4808:Gradualism
4733:Metabolism
4593:chromosome
4583:Eukaryotes
4361:Modularity
4278:Population
4204:Population
4165:Speciation
4143:Panspermia
4096:Extinction
4091:Exaptation
4066:Divergence
4039:Cladistics
4027:Reciprocal
4007:Adaptation
3901:Camouflage
3493:: 325–83.
1820:0632009985
1510:References
1273:non-silent
1240:the signal
1110:M. cyaneus
1071:green alga
1003:parasitism
919:hydrozoans
882:altruistic
688:In animals
658:adaptation
286:Cladistics
209:Extinction
194:Divergence
164:Speciation
144:Adaptation
58:John Gould
5168:Protocell
5019:Darwinism
4907:Sympatric
4656:processes
4544:Tetrapods
4493:Kangaroos
4419:Dinosaurs
4356:Inversion
4325:Variation
4246:Gene flow
4239:Inclusive
4049:Mutualism
3994:Evolution
3555:146590101
3032:CiteSeerX
2768:: 35–57.
2486:ignored (
2476:cite book
2250:Evolution
1943:CiteSeerX
1618:205218102
1560:ignored (
1550:cite book
1041:Mutualism
1020:Mutualism
995:mutualism
991:Symbiosis
986:Symbiosis
967:trade-off
774:In plants
650:evolution
545:Dysgenics
261:Phylogeny
159:Gene flow
129:Diversity
124:Variation
5232:Ethology
5196:Category
5071:Vitalism
5066:Theistic
5059:Spandrel
4743:Morality
4738:Monogamy
4613:plastids
4578:Flagella
4534:Reptiles
4515:sea cows
4498:primates
4407:Molluscs
4385:Bacteria
4273:Mutation
4206:genetics
4182:Taxonomy
4128:Mismatch
4108:Homology
4022:Cheating
4017:Altruism
3819:examples
3757:14869430
3749:17714660
3706:16625195
3662:17158317
3604:36071078
3547:12481126
3472:11639874
3464:12536274
3408:12941597
3363:10 March
3284:18557871
3207:53191998
2782:19027999
2681:15044219
2673:17703898
2631:10797005
2588:20679112
2531:11418855
2435:37085820
2421:: 1–11.
2400:16651531
2333:15329720
2324:1842/698
2280:13882735
2272:19154373
2189:35849910
2121:12637238
2113:10929290
2066:11161200
2022:21147795
1965:14576431
1755:17781971
1712:11935014
1610:19890322
1431:Teamwork
1426:Polytely
1378:See also
1279:through
1276:mutation
1165:Hamilton
1052:Fabaceae
1048:rhizobia
1029:Rhizobia
976:resource
892:and the
863:eviction
859:meerkats
678:babblers
609:Category
535:Eugenics
377:timeline
358:Evo-devo
316:Overview
134:Mutation
96:Evidence
91:Glossary
5087:Related
4917:History
4778:Meiosis
4713:Empathy
4708:Emotion
4608:nucleus
4549:Viruses
4539:Spiders
4451:Mammals
4434:Insects
4234:Fitness
4170:Species
3969:Outline
3906:Mimicry
3896:Crypsis
3850:Signals
3714:4340926
3686:Bibcode
3653:3279745
3632:Bibcode
3624:Science
3595:9449918
3527:Science
3444:Bibcode
3388:Bibcode
3349:Science
3330:2200930
3310:Bibcode
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