Cooperation (evolution)

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 1207: 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. 1259: 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. 813:

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. 1025: 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 1135: 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. 49: 5192: 604: 5202: 591: 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 1195: 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 1119:

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

909:

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

848:

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,

1369:

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

1242:

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

812:

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

1278:

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

1085:

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

951:

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

899:

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

749:

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

1373:

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

981:

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

1270:

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

1331:

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

1138:

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

1058:

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

745:

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

696:

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 792:

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

692:

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

1081:

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

720:

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 856:

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

750:

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.

982:

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.

1147:

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

793:

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" 1115:

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.

725:

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 671:

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

753:

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

1360:

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. 2035:

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 969:

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

1303:

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. 900:

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

1008:

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. 1190:

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.

964:

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 1351:

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

1097:

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.)

721:

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

1332:

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. 697:

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.

872:

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. 693:

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 ( 2354:

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).

1120:

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.

742:

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.

952:

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

801:

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:

3616: 4924: 320: 2645: 1167:

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

1086:

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.

2203: 2413:

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. 993:

refers to two or more biological species that interact closely, often over a long period of time. Symbiosis includes three types of interactions—

635: 4673: 3344: 1202:

believes is a handicap, comparable to a race horse's handicap. The larger the handicap the more intrinsically fit the individual (see text).

3973: 746:"territory inheritance" theory contends that individuals help in order to have improved access to breeding areas once the breeders depart. 243: 2079:

Strassmann, J; Seppa, P.; Queller, D. C. (10 August 2000). "Absence of within-colony kin discrimination: foundresses of the social wasp,

554: 3808: 1185: 5177: 4632: 3818: 362: 17: 1450:

An individual's gene complement (or genome) can be represented by the letters of the alphabet. Each letter is represented twice: A

3983: 1226:. Both are costly in fitness terms, and both are generally conspicuous to other members of the population or species. This led 68: 1300: 4940: 4700: 3300:

Koeslag, J. H. (1990). "Koinophilia groups sexual creatures into species, promotes stasis, and stabilizes social behaviour".

3126: 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 906:

bacteria, where cooperation is disfavored when populations are well mixed, but favored when there is high local relatedness.

549: 1858:"Paying to stay or paying to breed? Field evidence for direct benefits of helping behavior in a cooperatively breeding fish" 1206: 4297: 2501:

Lutzoni, François; Pagel, Mark; Reeb, Valérie (2001). "Major fungal lineages are derived from lichen symbiotic ancestors".

1925:

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 1160: 731: 656:

is the process where groups of organisms work or act together for common or mutual benefits. It is commonly defined as any

496: 1374:

puzzle in the evolution theory. In addition, the work has potential therapeutic benefits for numerous incurable diseases.

5023: 3943: 4685: 4345: 3785: 3665: 2804: 1840: 1539: 628: 357: 188: 1243:

are aimed at reducing disparities in performance, thereby making the outcome of contests less predictable. In a horse

2465: 1415: 833:

in 2007 after he found that benefits can result as an automatic consequence of an otherwise "self-interested" act in

4752: 1328:

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. 4757: 4355: 1810:

Grafen, A. (1984) Natural selection, kin selection and group selection. In Krebs, J. R. & Davies, N. B. (Eds.)

90: 1239: 5002: 4992: 4916: 4191: 2151:

Heinsohn, R; Packer, C. (1 September 1995). "Complex cooperative strategies in group-territorial African lions".

1186:

Evolutionary mechanisms suggesting that reciprocity is the result, not the cause, of the evolution of cooperation

1143:

Even if all members of a group benefit from cooperation, individual self-interest may not favor cooperation. The

1059:

since both the bacteria and the plant can survive independently, it is also an example of facultative symbiosis.

347: 315: 5130: 4997: 4777: 4521: 4319: 4223: 4112: 3014: 1819: 1362: 1296: 1258: 1172: 569: 352: 2293:

Griffin, A; West, S. A.; Buckling, A. (28 August 2004). "Cooperation and competition in pathogenic bacteria".

5226: 5195: 4782: 4526: 4265: 2925:

Koeslag, J.H. (1997). "Sex, the Prisoner's Dilemma Game, and the evolutionary inevitability of cooperation".

621: 608: 4727: 2207: 5135: 4637: 539: 1016:, in which one species lives inside of another, and ectosymbiosis, in which one species lives on another. 5205: 4799: 595: 3378:

Koeslag, J.H. (2003). "Evolution of cooperation: cooperation defeats defection in the cornfield model".

4901: 4075: 3570:"The conditional defector strategies can violate the most crucial supporting mechanisms of cooperation" 1682:

Clutton-Brock, T. (2002). "Breeding together: Kin selection and mutualism in cooperative vertebrates".

95: 3676:

Burtsev, Mikhail; Turchin, Peter (2006). "Evolution of cooperative strategies from first principles".

4695: 4492: 4302: 4070: 3963: 461: 436: 416: 396: 73: 4824: 3036: 2998: 2487: 1561: 5118: 4982: 4954: 4929: 4886: 4789: 4722: 4597: 4470: 4438: 4411: 4401: 3861: 1947: 1891:"Direct fitness benefits of delayed dispersal in the cooperatively breeding red wolf (Canis rufus)" 1399: 1288: 1263: 451: 446: 421: 376: 342: 336: 325: 4602: 5123: 5053: 4962: 4572: 4340: 3936: 3835: 3356: 1244: 1211: 1182:

show, however, that humans often act more cooperatively than strict self-interest would dictate.

564: 471: 466: 411: 372: 27:

Evolutionary process where groups of organisms work or act together for common or mutual benefits

5162: 4811: 4680: 4642: 4543: 4514: 4487: 4482: 4080: 3778: 3031: 1942: 1404: 1179: 902: 476: 431: 253: 148: 4668: 2977: 2034: 5103: 4977: 4896: 4891: 4876: 4861: 4851: 4767: 4742: 4577: 4533: 4497: 4465: 4406: 4384: 4365: 4228: 4181: 4132: 4127: 4085: 2985: 1409: 1348: 1144: 1129: 946: 574: 456: 401: 367: 280: 1214:

carry the largest weights, so the size of the handicap is a measure of the animal's quality.

5077: 4972: 4906: 4707: 4622: 4538: 4450: 4433: 4360: 4350: 4060: 3968: 3952: 3813: 3794: 3685: 3631: 3443: 3387: 3309: 3151: 3079: 3023: 2934: 2876: 2829: 2726: 2610: 2510: 2367: 2302: 2160: 2092: 2045: 1934: 1777: 1734: 1691: 1643: 1589: 1198:

A male peacock with its beautiful but clumsy, aerodynamically unsound erectile tail, which

975: 885: 778:

Cooperation exists not only in animals but also in plants. In a greenhouse experiment with

661: 426: 198: 83: 40: 4712: 4582: 884:

cooperation. There are three major mechanisms that generate this type of fitness benefit:

8: 5140: 5098: 5048: 4967: 4815: 4807: 4737: 4717: 4663: 4507: 4260: 4203: 4065: 4048: 4026: 3886: 3830: 3070:

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: 4418: 3689: 3635: 3498: 3447: 3391: 3313: 3155: 3083: 3027: 2938: 2880: 2833: 2730: 2614: 2514: 2371: 2306: 2164: 2096: 2049: 1938: 1781: 1738: 1695: 1647: 1593: 1024: 5113: 5065: 5058: 4655: 4567: 4428: 4389: 4213: 4157: 4147: 4107: 4021: 4016: 4011: 3929: 3881: 3840: 3825: 3752: 3709: 3652: 3594: 3569: 3550: 3467: 3279: 3275: 3202: 3115: 2845: 2777: 2676: 2578: 2553: 2534: 2475: 2430: 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: 5201: 5155: 4881: 4829: 4607: 4423: 4324: 4287: 4282: 4238: 4233: 4186: 4152: 3856: 3771: 3744: 3701: 3657: 3599: 3554: 3542: 3459: 3403: 3325: 3167: 3122: 3095: 3091: 3049: 3045: 2973: 2950: 2904: 2899: 2864: 2800: 2742: 2668: 2626: 2583: 2526: 2461: 2395: 2328: 2267: 2262: 2245: 2243: 2176: 2108: 2061: 2017: 1960: 1836: 1815: 1793: 1789: 1750: 1707: 1659: 1655: 1617: 1605: 1535: 1280: 1231: 1134: 1105: 1033: 815: 780: 763: 727: 713: 681: 677: 260: 138: 128: 123: 3756: 3471: 3283: 3206: 2781: 2680: 2644:

Bergmüller, 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: 3494: 3451: 3395: 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: 4084: 4082: 4079: 4077: 4074: 4072: 4069: 4067: 4064: 4062: 4059: 4057: 4054: 4050: 4047: 4046: 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: 3961: 3958: 3954: 3947: 3942: 3940: 3935: 3933: 3928: 3927: 3924: 3912: 3909: 3907: 3904: 3902: 3899: 3897: 3894: 3892: 3888: 3885: 3883: 3880: 3878: 3875: 3873: 3870: 3868: 3865: 3864: 3863: 3860: 3858: 3855: 3854: 3852: 3848: 3842: 3839: 3837: 3834: 3832: 3829: 3827: 3824: 3820: 3817: 3816: 3815: 3812: 3810: 3807: 3806: 3804: 3800: 3796: 3789: 3784: 3782: 3777: 3775: 3770: 3769: 3766: 3758: 3754: 3750: 3746: 3741: 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: 3571: 3564: 3556: 3552: 3548: 3544: 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 3172:2402153 3152:Bibcode 3080:Bibcode 3054:1195756 3024:Bibcode 2978:1592456 2955:9398503 2935:Bibcode 2909:8506355 2877:Bibcode 2850:4335611 2830:Bibcode 2747:7466396 2727:Bibcode 2719:Science 2611:Bibcode 2603:Science 2579:2936175 2539:4414913 2511:Bibcode 2391:1564281 2368:Bibcode 2341:4429250 2303:Bibcode 2181:7652573 2161:Bibcode 2153:Science 2093:Bibcode 2046:Bibcode 2038:Science 2013:3107641 1973:8338275 1935:Bibcode 1927:Science 1798:5875340 1778:Bibcode 1735:Bibcode 1727:Science 1692:Bibcode 1684:Science 1664:5875341 1644:Bibcode 1590:Bibcode 1224:peacock 1161:Axelrod 1139:spoils. 1063:Lichens 101:History 84:Outline 5206:Portal 4882:Hybrid 4718:Ethics 4560:organs 4522:Plants 4508:lemurs 4503:humans 4488:horses 4478:hyenas 4466:wolves 4461:canids 4395:origin 3755: 3747: 3712: 3704: 3678:Nature 3660: 3650: 3602: 3592: 3553: 3545: 3470: 3462: 3406: 3328: 3282: 3205: 3170: 3125: 3100:904334 3098: 3052: 3034: 2976: 2953: 2907: 2897: 2848: 2822:Nature 2803: 2780: 2745: 2679: 2671: 2629: 2586: 2576: 2537: 2529: 2503:Nature 2464: 2433: 2398: 2388: 2339: 2331: 2295:Nature 2278: 2270: 2233:: 1–8. 2187: 2179: 2119: 2111: 2064: 2020: 2010: 1971: 1963: 1945: 1839: 1818: 1796: 1753: 1710: 1662: 1616: 1608: 1582:Nature 1538: 1356:. The 1104:, the 1067:fungus 1001:, and 954:cancer 929:, and 766:, and 708:, and 607: 331:Darwin 4669:Death 4664:Aging 4643:brain 4429:Fungi 4390:Birds 4303:Fungi 4101:Event 3984:Index 3753:S2CID 3710:S2CID 3669:(PDF) 3620:(PDF) 3551:S2CID 3468:S2CID 3280:S2CID 3203:S2CID 2900:46660 2846:S2CID 2778:S2CID 2677:S2CID 2649:(PDF) 2535:S2CID 2431:S2CID 2337:S2CID 2276:S2CID 2185:S2CID 2117:S2CID 1969:S2CID 1614:S2CID 1490:... Z 1454:and A 1438:Notes 1301:above 1108:. In 1073:or a 972:trade 927:yeast 674:zebra 666:lions 69:Index 5156:Tree 4628:hair 4568:Cell 4471:dogs 4456:cats 4446:Life 4424:Fish 4377:taxa 3745:PMID 3702:PMID 3658:PMID 3600:PMID 3543:PMID 3460:PMID 3404:PMID 3365:2015 3326:PMID 3168:PMID 3123:ISBN 3096:PMID 3050:PMID 2999:help 2974:SSRN 2951:PMID 2905:PMID 2801:ISBN 2743:PMID 2669:PMID 2627:PMID 2584:PMID 2527:PMID 2488:help 2462:ISBN 2396:PMID 2329:PMID 2268:PMID 2216:2013 2177:PMID 2109:PMID 2062:PMID 2018:PMID 1961:PMID 1837:ISBN 1816:ISBN 1794:PMID 1751:PMID 1708:PMID 1660:PMID 1606:PMID 1562:help 1536:ISBN 1324:and 1163:and 1150:game 931:ants 805:and 79:Main 4654:Of 4623:eye 4573:DNA 4558:Of 4375:Of 3735:doi 3694:doi 3682:440 3648:PMC 3640:doi 3628:314 3590:PMC 3582:doi 3535:doi 3531:298 3495:doi 3452:doi 3396:doi 3384:224 3353:344 3318:doi 3306:144 3272:doi 3195:doi 3160:doi 3148:144 3088:doi 3042:doi 2943:doi 2931:189 2895:PMC 2885:doi 2838:doi 2826:355 2770:doi 2735:doi 2723:211 2661:doi 2619:doi 2607:288 2574:PMC 2566:doi 2562:365 2519:doi 2507:411 2454:doi 2423:doi 2386:PMC 2376:doi 2364:103 2319:hdl 2311:doi 2299:430 2258:doi 2169:doi 2157:269 2101:doi 2054:doi 2042:291 2008:PMC 2000:doi 1996:278 1953:doi 1931:302 1903:doi 1870:doi 1786:doi 1743:doi 1731:197 1700:doi 1688:296 1652:doi 1598:doi 1586:462 1528:doi 1482:, D 1474:, C 1466:, B 1419:by 1262:An 680:to 648:In 56:by 5223:: 3751:. 3743:. 3731:17 3729:. 3725:. 3708:. 3700:. 3692:. 3680:. 3656:. 3646:. 3638:. 3626:. 3622:. 3598:. 3588:. 3578:12 3576:. 3572:. 3549:. 3541:. 3529:. 3489:. 3466:. 3458:. 3450:. 3440:89 3438:. 3402:. 3394:. 3382:. 3351:. 3347:. 3324:. 3316:. 3304:. 3292:^ 3278:. 3266:. 3228:^ 3201:. 3191:46 3189:. 3166:. 3158:. 3146:. 3094:. 3086:. 3076:67 3074:. 3062:^ 3048:. 3040:. 3030:. 3020:53 3018:. 2990:: 2988:}} 2984:{{ 2949:. 2941:. 2929:. 2917:^ 2903:. 2893:. 2883:. 2873:90 2871:. 2867:. 2844:. 2836:. 2824:. 2776:. 2766:46 2764:. 2741:. 2733:. 2721:. 2709:^ 2689:^ 2675:. 2667:. 2657:76 2655:. 2651:. 2625:. 2617:. 2605:. 2582:. 2572:. 2560:. 2556:. 2533:. 2525:. 2517:. 2505:. 2480:: 2478:}} 2474:{{ 2460:. 2429:. 2419:35 2417:. 2394:. 2384:. 2374:. 2362:. 2358:. 2335:. 2327:. 2317:. 2309:. 2297:. 2274:. 2266:. 2254:63 2252:. 2248:. 2183:. 2175:. 2167:. 2155:. 2129:^ 2115:. 2107:. 2099:. 2089:87 2087:. 2060:. 2052:. 2040:. 2016:. 2006:. 1994:. 1990:. 1967:. 1959:. 1951:. 1941:. 1929:. 1917:^ 1899:22 1897:. 1893:. 1864:. 1860:. 1792:. 1784:. 1772:. 1749:. 1741:. 1729:. 1706:. 1698:. 1686:. 1672:^ 1658:. 1650:. 1638:. 1626:^ 1612:. 1604:. 1596:. 1584:. 1570:^ 1554:: 1552:}} 1548:{{ 1534:. 1526:. 1500:m2 1496:f2 1494:/Z 1492:m1 1488:f1 1486:/D 1484:m1 1480:f1 1478:/C 1476:m1 1472:f2 1470:/B 1468:m2 1464:f1 1462:/A 1460:m2 1291:, 1255:. 997:, 925:, 921:, 896:. 888:, 819:. 809:. 762:, 758:, 704:, 652:, 4832:/ 4823:/ 4814:/ 4810:/ 3945:e 3938:t 3931:v 3787:e 3780:t 3773:v 3759:. 3737:: 3716:. 3696:: 3688:: 3642:: 3634:: 3606:. 3584:: 3557:. 3537:: 3501:. 3497:: 3491:5 3474:. 3454:: 3446:: 3410:. 3398:: 3390:: 3367:. 3332:. 3320:: 3312:: 3286:. 3274:: 3268:1 3239:. 3209:. 3197:: 3174:. 3162:: 3154:: 3131:. 3102:. 3090:: 3082:: 3056:. 3044:: 3026:: 3001:) 2997:( 2980:. 2957:. 2945:: 2937:: 2911:. 2887:: 2879:: 2852:. 2840:: 2832:: 2809:. 2784:. 2772:: 2749:. 2737:: 2729:: 2683:. 2663:: 2633:. 2621:: 2613:: 2590:. 2568:: 2541:. 2521:: 2513:: 2490:) 2470:. 2456:: 2437:. 2425:: 2402:. 2378:: 2370:: 2343:. 2321:: 2313:: 2305:: 2282:. 2260:: 2218:. 2191:. 2171:: 2163:: 2123:. 2103:: 2095:: 2068:. 2056:: 2048:: 2024:. 2002:: 1975:. 1955:: 1937:: 1911:. 1905:: 1878:. 1872:: 1866:9 1845:. 1800:. 1788:: 1780:: 1774:7 1757:. 1745:: 1737:: 1714:. 1702:: 1694:: 1666:. 1654:: 1646:: 1640:7 1620:. 1600:: 1592:: 1564:) 1544:. 1530:: 1456:2 1452:1 1304:( 1148:" 637:e 630:t 623:v 379:) 375:( 20:)

Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.