247:, "Hyla arborea". In many sexually reproducing species such as this, some males can access mates by exploiting resources of more competitive males. Many species have dynamic reproductive strategies that can change in response to changes in the environment. In these instances, several factors contribute to the decision to switch between mating strategies. For example, in the European tree frog, a sexually competitive (as in, perceived to be attractive by females) male tend to call to attract mates. This is often referred to as the "bourgeois" tactic. On the other hand, a smaller male that would likely fail to attract mates using the bourgeois tactic will tend to hide near attractive males and attempt to access females. In this instance, the males can gain access to females without having to defend territories or acquiring additional resources (which often serve as the basis for attractiveness). This is referred to as the "parasitic" tactic, where the smaller male effectively cheats its way to accessing females, by reaping the benefit of sexual reproduction without contributing resources that normally attract females. Models such as this provide valuable tools for research aimed at energetic constraints and
230:
cleaners will feed on mucus when their clients are unable to control the cleaner's behavior; however, in natural settings, client fish often jolt, chase after cheating cleaners, or terminate interactions of swimming way, effectively controlling the cheating behavior. Studies on cleaning mutualisms generally suggest that cheating behavior is often adjusted depending on the species of the client. In cleaning shrimp, cheating is predicted to occur less often because shrimps bear a higher cost if the clients use aggression to control the cleaner's behavior. Studies have found that cleaner species can strategically adjust cheating behavior according to the potential associated risk. For example, predatory clients, which present a significantly high cost for cheating, experience less cheating behavior. On the other hand, nonpredatory clients present a lower cost for cheating, and thus experience more cheating behaviors from the cleaners. Some evidence suggest that physiological processes can mediate the cleaners' decision to switch from cooperating to cheating in mutualistic interactions. For example, in the bluestreak cleaner wrasse, changes in
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have potential for cheating to occur. Eusocial insects in the order
Hymenoptera, which includes bees and wasps, exhibit good examples of conflicts of interest present in insect societies. In these systems, queen bees and wasps can mate and lay fertilized eggs that hatch into females. On the other hand, workers of most species in Hymenoptera can produce eggs, but cannot produce fertilized eggs due to loss of mating ability. Workers that lay eggs represent a cost for the colony because workers that lay eggs often do significantly less work, and thus negatively impact the health of the colony (for example: decreased amount of collected food, or less attention given to tending the queen's eggs). In this case, a conflict of interest arises between the workers and the colony. The workers should lay eggs in order to pass on their genes; however, as a colony, having only the queen reproduce leads to better productivity. If workers sought to pass their own genes by laying eggs, foraging activities would diminish, leading to decreased resources for the entire colony. This, in turn, can cause a
58:, or mutually beneficial interactions between species, is common in ecological systems. These interactions can be thought of "biological markets" in which species offer partners goods that are relatively inexpensive for them to produce and receive goods that are more expensive or even impossible for them to produce. However, these systems provide opportunities for exploitation by individuals that can obtain resources while providing nothing in return. Exploiters can take on several forms: individuals outside a mutualistic relationship who obtain a commodity in a way that confers no benefit to either mutualist, individuals who receive benefits from a partner but have lost the ability to give any in return, or individuals who have the option of behaving mutualistically towards their partners but chose not to do so.
198:, where selfish behavior lead to the depletion of resources, with long-term negative consequences for the group. However, in natural bee and wasp societies, only 0.01â0.1% and 1%, respectively, of the workers lay eggs, suggesting that strategies exist to combat cheating to prevent tragedy of the commons. These insect systems have given scientists opportunities to study strategies that keep cheating in check. Such strategies are commonly referred to as "policing" strategies, generally where additional costs are imposed on cheaters to discourage or eliminate cheating behaviors. For example, honeybees and wasps may eat eggs produced by workers. In some ant species and yellowjackets, policing may occur via aggression towards or killing egg-laying individuals to minimize cheating.
521:"Cheating" sometimes happens when the yucca moth deposits too many eggs in one plant. In this case, the yucca plant has little to no benefits from this interaction. However, the plant has a unique way of constraining this behavior. While the constraint against cheating often occurs directly to the individual, in this case, the constraint occurs to the individual's offspring. The yucca plant can "abort" the moths by aborting the flowers. Pellmyr and Huth found that there is selective maturation for flowers that have low egg loads and high number of scars (and therefore a high amount of pollen). In this way, there is selection against the "cheaters" who try to use the yucca plant without providing the benefits of pollination.
101:
159:. However, when the scarcity of food sources causes individual cells to starve, roughly 10â´ to 10âľ cells aggregate to form a mobile, multicellular structure dubbed a "slug". In the wild, aggregates generally contain multiple genotypes, resulting in chimeric mixtures. Unlike clonal (genetically identical) aggregates typically found in multicellular organisms, the potential for competition exists in chimeric aggregates. For example, because individuals in the aggregate contain different genomes, differences in fitness can result in conflict of interest among cells in the aggregate, where different genotypes could potentially compete against each other for resources and reproduction. In
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the individual that initiated the cooperative behavior, mutualism should be selected for. When researchers investigated the co-evolution of cooperation and choice in a choosy host and its symbiont (an organism that lives in a relationship that benefits all parties involved), their model indicated that although choice and cooperation may be initially selected for, this would often be unstable. In other words, one cooperative partner will choose another cooperative partner if given a choice. However, if this choice is made over and over, variation is removed and this selection can no longer be maintained. This situation is similar to the
482:
setting, the cleaner fish undergoes behavioral change in face of deterrents against eating their preferential food. In several trials, the plate of their preferential food source was immediately removed when they eat it, to mimic "client fleeing" in natural settings. In other trials, the plate of their preferential food source chased the cleaner fish when they eat it, mimicking "client chasing" in natural setting. After only six learning trials, the cleaners learned to choose against their preference, indicating that punishment is potentially a very effective countermeasure against cheating in mutualistic relationships.
486:
518:. The female yucca moths deposit their eggs one at a time to the yucca flower. At the same time, she also deposits a small amount of pollen from yucca flowers as nutrition for the yucca moths. Because most of the pollen is not consumed by the larva, yucca moths are therefore also the active pollinators for the yucca plant. Moreover, sometimes the female moths do not successfully deposit their eggs the first time, and may try again and again. The yucca plant receives scars from the multiple attempts, but they also receive more pollen, since the moths are depositing pollen with every try.
92:
exploitive individuals, otherwise known as cheaters, contribute enough genetic variation to maintain mutualism itself. Both this theory and the negative frequency-dependent theory suggest that that cheating exists as part of a stable mixed evolutionary strategy with mutualism. In other words, cheating is a stable strategy used by individuals in a population where many other individuals cooperate. Another study supports that cheating can exist as a mixed strategy with mutualism using a mathematical game model. Thus, cheating can arise and be maintained in mutualistic populations.
507:. Moreover, these plants cannot tell whether the bacteria are more or less parasitic until they are settled in the plant nodules. To prevent cheating, these plants seem to be able to punish the rhizobium bacteria. In a series of experiments, researchers forced non-cooperation between the bacteria and the plants by placing various nodules in nitrogen-free atmosphere. They saw a decrease in the rhizobium reproductive success by 50%. West et al. created a model for legume sanctioning the bacteria and hypothesizes that these behaviors exist to stabilize mutualistic interactions.
425:, Wenseleers et al. found that a combination of aggressive behavior and destruction of worker-laid eggs kept the number of worker-laid eggs low. In fact, 91% of the worker-laid eggs were policed within one day. They also found that about 20% of workers laying eggs were prevented from doing so through both the queen's and workers' aggressive behavior. The workers and the queen would grab the egg-laying worker and try to sting her or push her off the cell. This usually results in the worker removing her abdomen and not depositing her eggs.
400:, it is collectively beneficial for all the individuals in the community to have the queen to lay eggs rather than the workers lay eggs. This is because if the workers lay eggs, it benefits the egg-laying worker individually, but the rest of the workers are now twice removed from this worker's offspring. Therefore, though it is beneficial for one individual to have its own offspring, it is collectively beneficial to have the queen lay the eggs. Therefore, a system of worker and queen policing exists against worker-laid eggs.
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the experiment, they recruit coalition support against this individual by screaming. The formed coalition then chases this individual away. If higher ranked individuals find this discoverer, they either chase the discoverer away or became physically aggressive towards the individual. These results show that aggression as punishment is a way to encourage members to work together and share food when it is found.
445:, punishments by the queen are a way she motivates the lazier, less-related workers in their groups. The queen would shove the lazier workers, with the number of shoves increasing when there are fewer active workers. Reeve found that if the queen is removed when colonies are satiated, there is a significant drop in weight of the active workers because the lazier workers are taking advantage of the system.
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not producing the antibiotic agent themselves, they are more likely to be negatively impacted by the antibiotic substrate because there is less antibiotic agent to protect everyone. Thus, cheaters can persist in a population because their exploitative behavior gives them an advantage when they exist at low frequencies but these benefits are diminished when they are greater in number.
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the study. Researchers theorize that this system developed because the fish are usually not closely related (so kinship benefits have little impact), and because there is a high level of predation risk when the fish is outside the group (therefore a strong motivator for the helper fish to stay in the group).
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of a phenotype depends on its frequency relative to other phenotypes in a population. Cheater phenotypes often display negative frequency-dependent selection, where fitness increases as a phenotype becomes less common and vice versa. In other words, cheaters do best (in terms of evolutionary benefits
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in their cooperative breeding systems. It is a pay-to-stay system where helper fish are allowed to stay in certain territories in exchange for their help. Similar to the naked mole rats, the helpers that were prevented from helping, the "idle helpers", receive more aggression than control helpers in
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colonies, there are antibiotic-sensitive "cheaters" that persist at low numbers on antibiotic-laced mediums when in a cooperative colony. These cheaters enjoy the benefit of others producing antibiotic-resistant agents while producing none themselves. However, as numbers increase, if they persist in
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and number of kin: females call more often and females with more kin call more often. However, sometimes when food is found, the individual ("discoverer") does not call to attract its kin and, presumably, to share food. If lower ranked individuals find this discoverer to be in the food drop area of
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association, the study demonstrated that the non-producer was unable to do so in soil conditions, suggesting that the two strains could coexist. Further experiments suggested that this cheating prevention may be due to interactions with other microbes in the soil influencing the relationship or the
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Eusocial insects also serve as valuable tools in studying cheating. Eusocial insects behave cooperatively, where members of the community forgo reproduction to assist a few individuals to reproduce. Such model systems have potential for conflict of interest to arise among individuals, and thus also
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Others have proposed that cheating (exploitive behavior) can stabilize cooperation in mutualistic systems. In many mutualistic systems, there will be feedback benefits to those that cooperate. For instance, the fitness of both partners may be improved. If there is a high reward or many benefits for
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Cheaters, who do not cooperate but benefit from others who do cooperate gain a competitive edge. In an evolutionary context, this competitive edge refers to a greater ability to survive or to reproduce. If individuals who cheat are able to gain survivorship and reproductive benefits while incurring
37:
to describe behavior whereby organisms receive a benefit at the cost of other organisms. Cheating is common in many mutualistic and altruistic relationships. A cheater is an individual who does not cooperate (or cooperates less than their fair share) but can potentially gain the benefit from others
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from inside the roots of leguminous plants, providing this essential source of nitrogen to these plants while also receiving organic acids for themselves. However, some bacteria are more mutualistic, while others are more parasitic because they consume the plant's resources but fixes little to no
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susceptible to cheating individuals that take advantage of the reproductive behavior without paying the fair price. In other words, if certain individuals tend to become a part of the sorus more frequently, they can gain increased benefit from the fruiting body system without sacrificing their own
108:
Studies of cheating and dishonest communication in populations presupposes an organismal system that cooperates. Without a collective population that has signaling and interactions among individuals, behaviors such as cheating do not manifest. In other words, in order to study cheating behavior, a
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removed by the cleaners. In these situations, instead of picking off the parasites on the surface of the client fish, the cleaner can cheat by feeding on the client's tissue (mucus layer, scales, etc.), thereby gaining additional benefit from the symbiotic system. It has been well documented that
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populations are placed into growth conditions with a proportion of adenosine, the cheaters are suppressed because the bacteria that responds to the quorum signal now produces adenosine hydrolase that they privately use for themselves to digest adenosine food source. In wild populations where the
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One theory states that cheating maintains genetic variation in the face of selection for mutualism. One study shows that a small influx of immigrants with a tendency to cooperate less can generate enough genetic variability to stabilize selection for mutualism. This suggests that the presence of
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and reef fish. Bshary and
Grutter found that cleaner wrasse prefers the client tissue mucus over ectoparasites. This creates a conflict between the cleaner fish and reef fish, because the reef fish only benefit when the cleaner fish eats the ectoparasites. Further studies revealed that in a lab
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were studied in the soil, their natural environment. One strain, known as the producer, produced a higher level of siderophores, which meant that other strain, known as the non-producer, ceased siderophore production in favor of using the other's siderophores. Although one would expect that the
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that detect their population density and prompt the transcription of various resources when needed. In this case, the resources are publicly shared proteases that break down a food source like casein, and privately used adenosine hydrolase, which breaks down another food source, adenosine. The
234:
levels are associated with behavior changes. For smaller clients, increase cortisol levels in the water lead to more cooperative behavior, while for larger clients, the same treatment lead to more dishonest behavior. It has been suggested that "good behavior" toward smaller clients often allow
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By definition, individuals cheat to gain benefits that their non-cheating counterparts do not receive. This raises the question of how a cooperative system can exist in face of these cheaters. One answer is that the cheaters actually have a reduced fitness compared to the non-cheaters.
167:. The remaining 80% of cells become spores in the sorus of the fruiting body, which can germinate again once conditions are more favorable. In this case, 20% of the cells must give up reproduction so that the fruiting body forms successfully. This makes chimeric aggregates of
380:
Cheating is also commonly found in insects. The social and seemingly altruistic communities found in insects such as ants and bees provide ample opportunities for cheaters to take advantage of the system and accrue additional benefits at the expense of the community.
410:), Ratneiks & Visscher found that other workers effectively removed worker-laid eggs in all colonies, whether the eggs were from originated from the same colonies or not. An example of a combination of queen and worker policing is found in ants, in the genus
416:, in which worker-laid eggs are taken by other workers and fed to the "queen". In general, these signals that identify the eggs as queen-laid are likely incorruptible, since it must be an honest signal to be maintained and not be used by cheating workers.
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Santorelli, L.A.; Thompson, C. R.; Villegas, E.; Svetz, J.; Dinh, C.; Parikh, A.; Sucgang, R.; Kuspa, A.; Strassmann, J.E.; Queller, D.C.; Shaulsky, G. (2008). "Facultative cheater mutants reveal the genetic complexity of cooperation in social amoebae".
396:" for its ability to take on properties greater than those of the sum of individuals. A colony of insects in which different individuals are specialized for specific tasks means a greater colony production and greater efficiency. Moreover, based on the
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in female choice. For example, in lek paradox, if females consistently choose for a particular male trait, genetic variance for that trait should eventually be eliminated, removing the benefits of choice. However, that choice somehow still persists.
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no costs, natural selection should favor cheaters. However, mechanisms also exist that prevent cheaters from undermining mutualistic systems. One main factor is that the advantages of cheating are often frequency-dependent.
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Cheating and constraints of cheating are not limited to intraspecific interactions; it can also occur in a mutualistic relationship between two species. A common example is the mutualistic relationship between cleaner fish
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problem arises when some individuals ("cheaters") do not respond to these quorum sensing signals and therefore do not contribute to the costly protease production yet enjoy the benefits of the broken down resources.
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involved in cheating. Studies find that mating strategies are highly adaptable and depend on a variety of factors, such as competitiveness, energetic costs involved in defending territory or acquiring resources.
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Finally the countermeasures are not limited to organismal relationships. West et al. found a similar countermeasure against cheating in legume-rhizobium mutualism. In this relationship, nitrogen fixing bacteria
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One form of policing occurs by the oophagy of the worker-laid eggs, found in many ant and bee species. This could be done by both or either the queen or the workers. In a series of experiments with honeybees
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populations are placed into growth conditions where cooperation (and responding to the quorum signal) is costly, the number of cheaters increases, and the public resources are depleted, which can lead to a
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is well established, and many studies have attempted to elucidate the evolutionary and genetic mechanisms underlying the behavior. Having a 34Mb genome that is completely sequenced and well annotated makes
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presence of adenosine is common, this is an explanation for how individuals that cooperate could have higher fitness than those that cheat, thereby suppressing the cheaters and maintaining cooperation.
269:, iron-chelating particles that seek and bring back iron for the bacteria. Siderophores are not necessarily specific to its producer - sometimes another individual could take up the particles instead.
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states that facilitation, cooperation or mutualism should be more common in stressful environments, while cheating, competition or parasitism are common in benign environments (i.e nutrient excess).
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Like many other organisms, bacteria rely on iron intake for its biological processes. However, iron is sometimes difficult to access in certain environments, like soil. Some bacteria have evolved
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that develop between small and larger marine organisms often represent models useful for studying the evolution of stable social interactions and cheating. In the cleaning fish
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also use aggression as a punishment. These animals have five distinct calls that they can "decide" to produce upon finding food. Whether they call or not is related to their
283:, to retrieve the iron necessary for survival. However, when iron is readily available, either from freely diffusing in environment or another bacterium's siderophores,
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spatial structure of the soil preventing siderophore diffusion and therefore limiting the non-producer's ability to exploit the producer's siderophores.
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insects, such as ants, bees, and wasps; and inter-specific interactions found in cleaning mutualisms. Common examples of cleaning mutualisms include
815:(2009). "Frequency-Dependent Success of Cheaters During Foraging Bouts Might Limit Their Spread within Colonies of a Socially Polymorphic Spider".
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344:) under varying environmental conditions. These microorganisms, like many species of bacteria, use a cell-cell communication system called
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such as increased survival and reproduction) when there are relatively few of them, but as cheaters become more abundant, they do worse.
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cooperating. Cheaters are also those who selfishly use common resources to maximize their individual fitness at the expense of a group.
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Brepson, L.; TroĂŻanowski, M.; Voituron, Y.; Lengagne, T. (2004). "Cheating for sex: inherent disadvantage or energetic constraint?".
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Castellano, S.; Marconi, V.; Zanollo, V.; Berto, G. (2009). "Alternative mating tactics in the
Italian treefrog, Hyla intermedia".
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In a study by
Dandekar et al., the researchers examined the survival rates of cheating and non-cheating bacteria populations (
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Strassmann, J. E.; Y. Zhu; D.C.Queller (2000). "Altruism and social cheating in the social amoeba
Dictyostelium discoideum".
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ceased siderophore production in favor of taking up iron-bound coelichelin, an association also known as siderophore piracy.
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Schwartz, M. W.; Hoeksema, J. D (1998). "Specialization and resource trade: biological markets as a model of mutualisms".
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Kiers, E. T.; Rousseau, R. A; West, S. A.; Denison, R. F. (2003). "Host sanctions and the legumeârhizobium mutualism".
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Bshary, R.; Grutter, A.S. (2002). "Asymmetric cheating opportunities and partner control in a cleaner fish mutualism".
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Galet, Justine; Deveau, AurĂŠlie; HĂ´tel, Laurence; Frey-Klett, Pascale; Leblond, Pierre; Aigle, Bertrand (2015-05-01).
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MacLean, R. C.; Gudelj, I. (2006). "Resource competition and social conflict in experimental populations of yeast".
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1498:"Pseudomonas fluorescens Pirates both Ferrioxamine and Ferricoelichelin Siderophores from Streptomyces ambofaciens"
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West, S. A.; Griffin, A. S.; Gardner, A.; Diggle, S. P. (2006). "Social evolution theory for microorganisms".
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Kikuta, N.; Tsuji, K. (1999). "Queen and worker policing in the monogynous and monandrous ant, Diacamma sp".
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Li, S.I.; Purugganan, M.D. (2011). "The cooperative amoeba: Dictyostelium as a model for social evolution".
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The other form of policing occurs through aggression towards egg-laying workers. In a species of tree wasp
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a useful model in studying the genetic bases and molecular mechanisms of cheating, and in a broader sense,
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ceases production, allowing the bacterium to devote its energy towards growth. One study showed that when
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1928:"Group-size-dependent punishment of idle subordinates in a cooperative breeder where helpers pay to stay"
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Beekman, M.; Oldroyd, B. P. (2008). "When workers disunite: intraspecific parasitism by eusocial bees".
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Reeve, H.K. (1992). "Queen activation of lazy workers in colonies of the eusocial naked mole-rat".
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SzĂĄmadĂł, S. (2000). "Cheating as a mixed strategy in a simple model of aggressive communication".
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1297:
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1020:
938:
616:
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3797:
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2615:
2593:
2401:
2212:
2200:
2169:
2147:
2139:
2121:
2102:
2058:
2050:
2006:
1996:
1947:
1939:
1909:
1889:
1846:
1807:
1784:
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1188:
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1125:
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3939:
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1289:
4807:
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4290:
4163:
3879:
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3613:
3409:
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2695:
2475:
2279:
586:
375:
1192:
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4190:
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3121:
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2356:
2334:
1172:
Soares, M.C.; Cardoso, S.C.; Grutter, A.S.; Oliveira, R.F.; Bshary, R. (2014).
812:
458:
442:
345:
163:, roughly 20% of the cells in the aggregate become dead to make the stalk of a
128:
1977:"Costs of deception: cheaters are punished in rhesus monkeys (Macaca mulatta)"
1850:
1456:
1093:
4890:
4802:
4745:
4735:
4295:
4115:
4075:
4071:
4062:
3905:
3874:
3863:
3853:
3780:
3682:
3576:
3451:
3329:
3288:
3134:
2888:
2556:
2551:
2001:
1586:
1529:
1442:
537:
397:
393:
261:
Environmental conditions and social interactions affecting microbial cheating
113:
is needed. Models that provide insight on cheating include the social amoeba
1643:
876:
4592:
3893:
3870:
3446:
3394:
3339:
3172:
3167:
2761:
2519:
2161:
2072:
2054:
1961:
1943:
1741:
1703:
1661:
1604:
1578:
1547:
1403:
1385:
1351:
1333:
1240:
1200:
1153:
1101:
1051:
992:
930:
922:
895:
836:
723:
705:
659:
608:
573:
555:
538:
Ferriere, R.; Bronstein, J.L.; Rinaldi, S.; Law, R.; Gauduchon, M. (2002).
124:
3517:
2088:
2020:
1901:
1811:
1755:
Ratnieks, F.L.; Visscher, P.K. (1989). "Worker policing in the honeybee".
1563:"Siderophore cooperation of the bacterium Pseudomonas fluorescens in soil"
433:
54:
Organisms communicate and cooperate to perform a wide range of behaviors.
4731:
4643:
4587:
4425:
4210:
3909:
3728:
3473:
3344:
3073:
2608:
2438:
2344:
2302:
1832:
1521:
441:
Aggression and punishment are not just found in insects. For example, in
266:
248:
201:
120:
110:
2913:
2143:
1677:
1617:
1043:
651:
600:
510:
Another well-known example of plant-organism interaction occurs between
312:
cheating behavior could be suppressed. In another study, two strains of
4678:
4673:
4390:
4125:
3665:
3334:
3235:
3157:
3144:
2778:
2578:
2465:
2443:
2396:
2391:
2339:
2307:
2222:
1128:"Worker reproduction and policing in insect societies: an ESS analysis"
515:
280:
226:
84:
1835:"Queen and worker policing in the tree wasp Dolichovespula sylvestris"
275:
is a bacterium commonly found in the soil. Under low-iron conditions,
4340:
3468:
3319:
2928:
2546:
2294:
2204:
1893:
1776:
1318:"Mutualism or parasitism? The variable outcome of cleaning symbioses"
984:
2152:
4792:
4552:
4257:
3371:
2573:
495:
412:
231:
164:
34:
1925:
1126:
Wenseleers, T.; Helanterä, H.; Hart, A.; Ratnieks, F.L.W. (2004).
962:
4693:
2756:
2470:
1620:"Bacterial quorum sensing and metabolic incentives to cooperate"
329:
152:
1367:
384:
147:
is a widely used model for cooperation and the development of
42:
favors cheating, but there are mechanisms to regulate it. The
2677:
1561:
LujĂĄn, Adela M.; GĂłmez, Pedro; Buckling, Angus (2015-02-01).
780:
511:
155:, single-celled state that feeds independently and undergoes
1754:
810:
795:
10.1890/0012-9658(1998)079[1029:sartbm]2.0.co;2
1495:
849:
235:
wrasses to attract larger clients that are often cheated.
2089:
West, S. A.; Kiers, E.T.; Pen, I.; Denison, R.F. (2002).
1719:
462:
428:
301:, no pyoverdine was detected. This result suggested that
225:), as in many cleaner species, client fish seeks to have
205:
Fish cleaned by smaller cleaner wrasses on
Hawaiian reefs
1478:
Wells, K. (2001). "The energetics of calling in frogs".
1079:
852:"Parasites may help stabilize cooperative relationships"
629:
2036:
1932:
Proceedings of the Royal
Society B: Biological Sciences
1226:
694:
Proceedings of the Royal
Society B: Biological Sciences
540:"Cheating and the evolutionary stability of mutualisms"
2182:
1833:
Wenseleers, T.; Tofilski, A.; Ratnieks, F. L. (2005).
1315:
687:
1678:
Ratnieks, F.L.; Foster, K.R.; Wenseleers, T. (2006).
1618:
Dandekar, A.A.; Chugani, S.; Greenberg, E.P. (2012).
1268:
317:
non-producer would outcompete the producer, like the
151:. This species of amoeba is most commonly found in a
1797:
1560:
1363:
1361:
1121:
1119:
1117:
1115:
1113:
1111:
690:"Cheating can stabilize cooperation in mutualisms"
297:, another bacterium that produces the siderophore
255:
1016:
1014:
1012:
1010:
958:
956:
954:
952:
950:
948:
369:
172:opportunities to reproduce. Cheating behavior in
4888:
739:
469:
1981:Proceedings of the National Academy of Sciences
1926:Fischer, S.; ZĂśttl, M.; Groenewoud, F. (2014).
1673:
1671:
1358:
1108:
1921:
1919:
1826:
1715:
1713:
1419:
1417:
1415:
1413:
1264:
1262:
1260:
1258:
1007:
945:
806:
804:
134:
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2238:
2032:
2030:
1311:
1309:
1307:
1222:
1220:
1218:
1167:
1165:
1163:
330:Selection pressure in bacteria (intraspecies)
1974:
1748:
1668:
1611:
531:
448:Punishment is also a method used by cichlid
392:Sometimes, a colony of insects is called a "
3547:
2176:
2084:
2082:
1916:
1871:
1710:
1410:
1255:
908:
801:
774:
735:
733:
3540:
3526:
2245:
2231:
2027:
1968:
1791:
1426:Proceedings of the Royal Society of London
1374:Proceedings of the Royal Society of London
1304:
1215:
1160:
623:
2151:
2115:
2106:
2062:
2010:
2000:
1951:
1865:
1680:"Conflict resolution in insect societies"
1651:
1594:
1537:
1477:
1393:
1370:"Tragedy of the commons in Melipona bees"
1368:Wenseleers, T.; Ratnieks, F.L.W. (2004).
1341:
1143:
1075:
1073:
1071:
1069:
885:
875:
843:
765:
713:
563:
16:Exploitive behavior in behavioral ecology
4704:Transgenerational epigenetic inheritance
3479:Transgenerational epigenetic inheritance
2252:
2079:
730:
484:
432:
383:
209:
200:
99:
683:
681:
679:
677:
580:
4889:
1502:Applied and Environmental Microbiology
1066:
429:Policing/punishment in other organisms
308:More studies, however, suggested that
188:
4196:Psychological effects of Internet use
3521:
3242:Dialogues Concerning Natural Religion
2226:
1734:10.1146/annurev.ento.53.103106.093515
1696:10.1146/annurev.ento.51.110104.151003
1491:
1489:
850:Little, A. E.; Currie, C. R. (2009).
243:Other models of cheating include the
127:such as wrasses and gobies, and some
49:
674:
279:produces siderophores, specifically
4176:Digital media use and mental health
1839:Behavioral Ecology and Sociobiology
1800:Behavioral Ecology and Sociobiology
1445:Behavioral Ecology and Sociobiology
104:Stalked slime mould fruiting bodies
13:
3807:Automatic and controlled processes
2647:Evolutionary developmental biology
2037:Bshary, R.; Grutter, A.S. (2005).
1486:
489:Nitrogen-fixing nodules in legumes
14:
4913:
4216:Smartphones and pedestrian safety
1316:Cheney, K.L.; Cote, I.M. (2005).
688:Foster, K. R.; Kokko, H. (2006).
4867:
4854:
4842:
4841:
4241:Mobile phones and driving safety
3501:
3492:
3491:
2108:10.1046/j.1420-9101.2002.00441.x
1269:Chapuis, L.; Bshary, R. (2009).
1145:10.1111/j.1420-9101.2004.00751.x
829:10.1111/j.1558-5646.2009.00771.x
767:10.1046/j.1461-0248.2001.00218.x
742:"The exploitation of mutualisms"
4144:Computer-mediated communication
3304:Extended evolutionary synthesis
2493:Gene-centered view of evolution
2095:Journal of Evolutionary Biology
1554:
1471:
1436:
256:Constraints and countermeasures
4421:Empathisingâsystemising theory
3724:female intrasexual competition
3661:Evolutionarily stable strategy
3432:Hologenome theory of evolution
3299:History of molecular evolution
2525:Evolutionarily stable strategy
2414:Last universal common ancestor
902:
370:Policing/punishment in insects
1:
4781:Standard social science model
3834:Cognitive tradeoff hypothesis
3226:Renaissance and Enlightenment
1290:10.1016/j.anbehav.2009.06.001
524:
470:Interspecific countermeasures
109:model system that engages in
64:Frequency-dependent selection
4629:Missing heritability problem
4221:Social aspects of television
3844:Evolution of nervous systems
3812:Computational theory of mind
3437:Missing heritability problem
3064:Gamete differentiation/sexes
7:
4875:Evolutionary biology portal
1193:10.1016/j.yhbeh.2014.06.010
589:Nature Reviews Microbiology
388:Wasp nest, with some larvae
95:
10:
4918:
4836:Evolutionary psychologists
4709:TriversâWillard hypothesis
4624:Humanâanimal communication
4336:Ovulatory shift hypothesis
4186:Imprinted brain hypothesis
4154:Humanâcomputer interaction
3069:Life cycles/nuclear phases
2621:TriversâWillard hypothesis
373:
44:stress gradient hypothesis
18:
4830:
4756:Environmental determinism
4727:Cultural selection theory
4719:
4614:Evolutionary epistemology
4601:
4528:evolutionary neuroscience
4490:
4483:
4381:
4256:
4201:Rank theory of depression
4124:
4048:
3950:
3756:
3749:
3703:Parentâoffspring conflict
3612:
3555:
3487:
3387:
3312:
3216:
3143:
3099:
2954:
2858:
2675:
2634:
2567:Parentâoffspring conflict
2503:
2372:Earliest known life forms
2293:
2260:
1851:10.1007/s00265-004-0892-4
1457:10.1007/s00265-009-0756-z
1094:10.1016/j.tig.2010.11.003
740:Bronstein, J. L. (2001).
437:A nest of naked mole rats
422:Dolichovespula sylvestris
291:grew in association with
223:Bluestreak cleaner wrasse
21:cheating (disambiguation)
4649:Cultural group selection
4533:Biocultural anthropology
4226:Societal impacts of cars
4159:Media naturalness theory
3849:Fight-or-flight response
3420:Cultural group selection
3284:The eclipse of Darwinism
3256:On the Origin of Species
3231:Transmutation of species
2002:10.1073/pnas.89.24.12137
856:BMC Evolutionary Biology
294:Streptomyces ambofaciens
238:
169:Dictyostelium discoideum
161:Dictyostelium discoideum
144:Dictyostelium discoideum
137:Dictyostelium discoideum
116:Dictyostelium discoideum
4902:Biological interactions
4849:Evolutionary psychology
4813:Sociocultural evolution
4654:Dual inheritance theory
4111:Personality development
3572:Theoretical foundations
3549:Evolutionary psychology
3425:Dual inheritance theory
3264:History of paleontology
1644:10.1126/science.1227289
877:10.1186/1471-2148-9-124
451:Neolamprologous pulcher
272:Pseudomonas fluorescens
4771:Social constructionism
4766:Psychological nativism
4741:Biological determinism
4689:Recent human evolution
4684:Punctuated equilibrium
4507:Behavioral epigenetics
4502:evolutionary economics
4471:Variability hypothesis
4416:Emotional intelligence
4149:Engineering psychology
3839:Evolution of the brain
3113:Punctuated equilibrium
2434:Non-adaptive radiation
2382:Evolutionary arms race
2055:10.1098/rsbl.2005.0344
1944:10.1098/rspb.2014.0184
1579:10.1098/rsbl.2014.0934
1386:10.1098/rsbl.2003.0159
1334:10.1098/rsbl.2004.0288
1241:10.1006/anbe.2001.1937
923:10.1006/anbe.1999.1293
706:10.1098/rspb.2006.3571
556:10.1098/rspb.2001.1900
490:
438:
389:
359:tragedy of the commons
341:Pseudomonas aeruginosa
206:
196:tragedy of the commons
105:
4798:Multilineal evolution
4761:Nature versus nurture
4720:Theoretical positions
4568:Functional psychology
4563:Evolutionary medicine
4538:Biological psychiatry
4246:Texting while driving
4236:Leadâcrime hypothesis
4096:Cognitive development
4081:Caregiver deprivation
3592:Gene selection theory
3405:Evolutionary medicine
3279:Mendelian inheritance
2987:Biological complexity
2975:Programmed cell death
2667:Phenotypic plasticity
2387:Evolutionary pressure
2377:Evidence of evolution
2275:Timeline of evolution
1975:Hauser, M.D. (1992).
1812:10.1007/s002650050608
1181:Hormones and Behavior
488:
436:
387:
374:Further information:
210:In cleaning symbiosis
204:
103:
4751:Cultural determinism
4558:Evolutionary biology
4543:Cognitive psychology
4491:Academic disciplines
4139:Cognitive ergonomics
4106:Language acquisition
4086:Childhood attachment
3899:Wason selection task
3793:Behavioral modernity
3582:Cognitive revolution
3565:Evolutionary thought
3379:Teleology in biology
3274:Blending inheritance
2652:Genetic assimilation
2515:Artificial selection
2254:Evolutionary biology
1522:10.1128/AEM.03520-14
1480:Anuran Communication
478:Labroides dimidiatus
398:kin-selection theory
219:Labroides dimidiatus
157:asexual reproduction
19:For other uses, see
4818:Unilineal evolution
4583:Population genetics
4368:Sexy son hypothesis
4306:Hormonal motivation
4286:Concealed ovulation
3827:Dual process theory
3698:Parental investment
3442:Molecular evolution
3400:Ecological genetics
3269:Transitional fossil
3059:Sexual reproduction
2899:endomembrane system
2828:pollinator-mediated
2784:dolphins and whales
2562:Parental investment
2197:1994Natur.372..257P
2144:10.1038/nature01931
2136:2003Natur.425...78K
1993:1992PNAS...8912137H
1987:(24): 12137â12139.
1886:1992Natur.358..147R
1769:1989Natur.342..796R
1636:2012Sci...338..264D
1514:2015ApEnM..81.3132G
1380:(Suppl 5): S310-2.
1044:10.1038/nature06558
1036:2008Natur.451.1107S
1030:(7182): 1107â1110.
977:2000Natur.408..965S
868:2009BMCEE...9..124L
758:2001EcolL...4..277B
700:(1598): 2233â2239.
652:10.1038/nature04624
644:2006Natur.441..498M
601:10.1038/nrmicro1461
498:fixes atmospheric N
189:In eusocial insects
4776:Social determinism
4659:Fisher's principle
4619:Great ape language
4609:Cultural evolution
4578:Philosophy of mind
4411:Division of labour
4373:Westermarck effect
4321:Mating preferences
4231:Distracted driving
3965:Literary criticism
3822:Domain specificity
3802:modularity of mind
3415:Cultural evolution
2530:Fisher's principle
2459:Handicap principle
2449:Parallel evolution
2313:Adaptive radiation
1938:(1789): 20140184.
1722:Annu. Rev. Entomol
1684:Annu. Rev. Entomol
544:Proc. R. Soc. Lond
491:
439:
390:
249:environmental cues
245:European tree frog
215:Cleaning symbiosis
207:
106:
50:Theoretical models
31:behavioral ecology
29:is a term used in
4884:
4883:
4862:Psychology portal
4826:
4825:
4669:Hologenome theory
4639:Unit of selection
4634:Primate cognition
4548:Cognitive science
4479:
4478:
4350:Sexual attraction
4326:Mating strategies
4091:Cinderella effect
4021:Moral foundations
3925:Visual perception
3817:Domain generality
3786:Facial expression
3734:Sexual dimorphism
3693:Natural selection
3639:Hamiltonian spite
3515:
3514:
3131:Uniformitarianism
3084:Sex-determination
2589:Sexual dimorphism
2584:Natural selection
2488:Unit of selection
2454:Signalling theory
2191:(6503): 257â260.
1763:(6251): 796â797.
1630:(6104): 264â266.
971:(6815): 965â967.
823:(11): 2966â2973.
638:(7092): 498â501.
550:(1493): 773â780.
40:Natural selection
4909:
4871:
4858:
4845:
4844:
4488:
4487:
4484:Related subjects
4271:Adult attachment
3798:Cognitive module
3754:
3753:
3741:Social selection
3715:Costly signaling
3710:Sexual selection
3597:Modern synthesis
3542:
3535:
3528:
3519:
3518:
3505:
3495:
3494:
3294:Modern synthesis
3054:Multicellularity
3049:Mosaic evolution
2934:auditory ossicle
2616:Social selection
2599:Flowering plants
2594:Sexual selection
2247:
2240:
2233:
2224:
2223:
2217:
2216:
2205:10.1038/372257a0
2180:
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2155:
2119:
2113:
2112:
2110:
2086:
2077:
2076:
2066:
2034:
2025:
2024:
2014:
2004:
1972:
1966:
1965:
1955:
1923:
1914:
1913:
1894:10.1038/358147a0
1869:
1863:
1862:
1830:
1824:
1823:
1795:
1789:
1788:
1777:10.1038/342796a0
1752:
1746:
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1717:
1708:
1707:
1675:
1666:
1665:
1655:
1615:
1609:
1608:
1598:
1558:
1552:
1551:
1541:
1508:(9): 3132â3141.
1493:
1484:
1483:
1475:
1469:
1468:
1451:(8): 1109â1118.
1440:
1434:
1433:
1421:
1408:
1407:
1397:
1365:
1356:
1355:
1345:
1313:
1302:
1301:
1278:Animal Behaviour
1275:
1266:
1253:
1252:
1229:Animal Behaviour
1224:
1213:
1212:
1178:
1169:
1158:
1157:
1147:
1123:
1106:
1105:
1077:
1064:
1063:
1018:
1005:
1004:
985:10.1038/35050087
960:
943:
942:
911:Animal Behaviour
906:
900:
899:
889:
879:
847:
841:
840:
808:
799:
798:
789:(3): 1029â1038.
778:
772:
771:
769:
737:
728:
727:
717:
685:
672:
671:
627:
621:
620:
584:
578:
577:
567:
535:
361:. However, when
183:social evolution
149:multicellularity
76:Escherichia coli
74:For example, in
66:occurs when the
4917:
4916:
4912:
4911:
4910:
4908:
4907:
4906:
4887:
4886:
4885:
4880:
4822:
4808:Neoevolutionism
4715:
4699:Species complex
4664:Group selection
4602:Research topics
4597:
4573:Neuropsychology
4475:
4461:Substance abuse
4383:Sex differences
4377:
4291:Coolidge effect
4252:
4164:Neuroergonomics
4129:
4120:
4044:
3946:
3880:Folk psychology
3761:
3745:
3615:
3608:
3551:
3546:
3516:
3511:
3483:
3410:Group selection
3383:
3308:
3212:
3139:
3101:Tempo and modes
3095:
2950:
2854:
2671:
2630:
2506:
2499:
2476:Species complex
2289:
2280:History of life
2256:
2251:
2221:
2220:
2181:
2177:
2130:(6953): 78â81.
2120:
2116:
2087:
2080:
2043:Biology Letters
2035:
2028:
1973:
1969:
1924:
1917:
1880:(6382): 147â9.
1870:
1866:
1831:
1827:
1796:
1792:
1753:
1749:
1718:
1711:
1690:(51): 581â608.
1676:
1669:
1616:
1612:
1573:(2): 20140934.
1567:Biology Letters
1559:
1555:
1494:
1487:
1476:
1472:
1441:
1437:
1432:(5): 1253â1260.
1422:
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1273:
1267:
1256:
1225:
1216:
1176:
1170:
1161:
1124:
1109:
1078:
1067:
1019:
1008:
961:
946:
907:
903:
848:
844:
813:Riechert, S. E.
811:Pruitt, J. N.;
809:
802:
779:
775:
746:Ecology Letters
738:
731:
686:
675:
628:
624:
585:
581:
536:
532:
527:
506:
501:
472:
443:naked mole rats
431:
378:
376:Worker policing
372:
332:
310:P. fluorescens'
263:
258:
241:
212:
191:
140:
98:
52:
24:
17:
12:
11:
5:
4915:
4905:
4904:
4899:
4882:
4881:
4879:
4878:
4865:
4852:
4839:
4831:
4828:
4827:
4824:
4823:
4821:
4820:
4815:
4810:
4805:
4800:
4795:
4790:
4785:
4784:
4783:
4778:
4773:
4768:
4763:
4758:
4753:
4748:
4743:
4729:
4723:
4721:
4717:
4716:
4714:
4713:
4712:
4711:
4706:
4701:
4696:
4691:
4686:
4681:
4676:
4671:
4666:
4661:
4656:
4651:
4646:
4636:
4631:
4626:
4621:
4616:
4611:
4605:
4603:
4599:
4598:
4596:
4595:
4590:
4585:
4580:
4575:
4570:
4565:
4560:
4555:
4550:
4545:
4540:
4535:
4530:
4513:
4504:
4494:
4492:
4485:
4481:
4480:
4477:
4476:
4474:
4473:
4468:
4463:
4458:
4453:
4448:
4443:
4438:
4433:
4428:
4423:
4418:
4413:
4408:
4403:
4398:
4393:
4387:
4385:
4379:
4378:
4376:
4375:
4370:
4365:
4352:
4343:
4338:
4333:
4328:
4323:
4318:
4313:
4308:
4303:
4298:
4293:
4288:
4283:
4278:
4273:
4268:
4262:
4260:
4254:
4253:
4251:
4250:
4249:
4248:
4243:
4238:
4233:
4223:
4218:
4213:
4208:
4203:
4198:
4193:
4191:Mind-blindness
4188:
4183:
4178:
4173:
4168:
4167:
4166:
4161:
4156:
4151:
4146:
4135:
4133:
4122:
4121:
4119:
4118:
4113:
4108:
4103:
4098:
4093:
4088:
4083:
4078:
4065:
4060:
4054:
4052:
4046:
4045:
4043:
4042:
4037:
4036:
4035:
4025:
4024:
4023:
4013:
4012:
4011:
4006:
4001:
3991:
3986:
3985:
3984:
3974:
3973:
3972:
3967:
3956:
3954:
3948:
3947:
3945:
3944:
3943:
3942:
3937:
3932:
3922:
3917:
3912:
3903:
3902:
3901:
3896:
3886:
3884:theory of mind
3877:
3868:
3867:
3866:
3861:
3856:
3846:
3841:
3836:
3831:
3830:
3829:
3824:
3819:
3814:
3809:
3795:
3790:
3789:
3788:
3783:
3778:
3767:
3765:
3751:
3747:
3746:
3744:
3743:
3738:
3737:
3736:
3731:
3726:
3717:
3707:
3706:
3705:
3695:
3690:
3685:
3680:
3679:
3678:
3668:
3663:
3658:
3653:
3651:Baldwin effect
3648:
3647:
3646:
3641:
3636:
3626:
3620:
3618:
3610:
3609:
3607:
3606:
3601:
3600:
3599:
3594:
3589:
3584:
3579:
3569:
3568:
3567:
3556:
3553:
3552:
3545:
3544:
3537:
3530:
3522:
3513:
3512:
3510:
3509:
3499:
3488:
3485:
3484:
3482:
3481:
3476:
3471:
3466:
3461:
3460:
3459:
3449:
3444:
3439:
3434:
3429:
3428:
3427:
3422:
3417:
3407:
3402:
3397:
3391:
3389:
3385:
3384:
3382:
3381:
3376:
3375:
3374:
3369:
3364:
3363:
3362:
3352:
3347:
3342:
3337:
3332:
3322:
3316:
3314:
3310:
3309:
3307:
3306:
3301:
3296:
3291:
3286:
3281:
3276:
3271:
3266:
3261:
3260:
3259:
3250:Charles Darwin
3247:
3246:
3245:
3233:
3228:
3222:
3220:
3214:
3213:
3211:
3210:
3205:
3200:
3195:
3190:
3188:Non-ecological
3185:
3180:
3175:
3170:
3165:
3160:
3155:
3149:
3147:
3141:
3140:
3138:
3137:
3128:
3119:
3105:
3103:
3097:
3096:
3094:
3093:
3088:
3087:
3086:
3081:
3076:
3071:
3066:
3056:
3051:
3046:
3041:
3036:
3031:
3026:
3021:
3016:
3011:
3006:
3005:
3004:
2994:
2989:
2984:
2979:
2978:
2977:
2972:
2961:
2959:
2952:
2951:
2949:
2948:
2947:
2946:
2941:
2939:nervous system
2936:
2931:
2926:
2918:
2917:
2916:
2911:
2906:
2901:
2896:
2891:
2881:
2876:
2871:
2865:
2863:
2856:
2855:
2853:
2852:
2847:
2842:
2837:
2832:
2831:
2830:
2820:
2819:
2818:
2813:
2812:
2811:
2806:
2796:
2791:
2786:
2781:
2776:
2775:
2774:
2769:
2759:
2749:
2744:
2743:
2742:
2732:
2727:
2722:
2717:
2716:
2715:
2705:
2700:
2699:
2698:
2688:
2682:
2680:
2673:
2672:
2670:
2669:
2664:
2659:
2654:
2649:
2644:
2638:
2636:
2632:
2631:
2629:
2628:
2623:
2618:
2613:
2612:
2611:
2606:
2601:
2591:
2586:
2581:
2576:
2571:
2570:
2569:
2564:
2554:
2549:
2544:
2543:
2542:
2532:
2527:
2522:
2517:
2511:
2509:
2501:
2500:
2498:
2497:
2496:
2495:
2485:
2480:
2479:
2478:
2473:
2463:
2462:
2461:
2451:
2446:
2441:
2439:Origin of life
2436:
2431:
2426:
2424:Microevolution
2421:
2419:Macroevolution
2416:
2411:
2406:
2405:
2404:
2394:
2389:
2384:
2379:
2374:
2369:
2364:
2359:
2357:Common descent
2354:
2353:
2352:
2342:
2337:
2335:Baldwin effect
2332:
2331:
2330:
2325:
2315:
2310:
2305:
2299:
2297:
2291:
2290:
2288:
2287:
2282:
2277:
2272:
2267:
2261:
2258:
2257:
2250:
2249:
2242:
2235:
2227:
2219:
2218:
2175:
2114:
2101:(5): 830â837.
2078:
2049:(4): 396â399.
2026:
1967:
1915:
1864:
1825:
1806:(3): 180â189.
1790:
1747:
1709:
1667:
1610:
1553:
1485:
1470:
1435:
1409:
1357:
1303:
1284:(2): 455â459.
1254:
1235:(3): 547â555.
1214:
1187:(2): 346â350.
1159:
1138:(5): 1035â47.
1107:
1065:
1006:
944:
917:(1): 221â230.
901:
842:
800:
773:
752:(3): 277â287.
729:
673:
622:
595:(8): 597â607.
579:
529:
528:
526:
523:
504:
499:
471:
468:
459:Rhesus monkeys
430:
427:
407:Apis mellifera
371:
368:
346:quorum sensing
331:
328:
323:S. ambofaciens
319:P. fluorescens
314:P. fluorescens
303:P. fluorescens
289:P. fluorescens
285:P. fluorescens
277:P. fluorescens
262:
259:
257:
254:
240:
237:
211:
208:
190:
187:
139:
133:
129:cleaner shrimp
97:
94:
51:
48:
15:
9:
6:
4:
3:
2:
4914:
4903:
4900:
4898:
4895:
4894:
4892:
4877:
4876:
4870:
4866:
4864:
4863:
4857:
4853:
4851:
4850:
4840:
4838:
4837:
4833:
4832:
4829:
4819:
4816:
4814:
4811:
4809:
4806:
4804:
4803:Neo-Darwinism
4801:
4799:
4796:
4794:
4791:
4789:
4788:Functionalism
4786:
4782:
4779:
4777:
4774:
4772:
4769:
4767:
4764:
4762:
4759:
4757:
4754:
4752:
4749:
4747:
4746:Connectionism
4744:
4742:
4739:
4738:
4737:
4736:indeterminism
4733:
4730:
4728:
4725:
4724:
4722:
4718:
4710:
4707:
4705:
4702:
4700:
4697:
4695:
4692:
4690:
4687:
4685:
4682:
4680:
4677:
4675:
4672:
4670:
4667:
4665:
4662:
4660:
4657:
4655:
4652:
4650:
4647:
4645:
4642:
4641:
4640:
4637:
4635:
4632:
4630:
4627:
4625:
4622:
4620:
4617:
4615:
4612:
4610:
4607:
4606:
4604:
4600:
4594:
4591:
4589:
4586:
4584:
4581:
4579:
4576:
4574:
4571:
4569:
4566:
4564:
4561:
4559:
4556:
4554:
4551:
4549:
4546:
4544:
4541:
4539:
4536:
4534:
4531:
4529:
4525:
4521:
4517:
4514:
4512:
4508:
4505:
4503:
4499:
4496:
4495:
4493:
4489:
4486:
4482:
4472:
4469:
4467:
4464:
4462:
4459:
4457:
4456:Schizophrenia
4454:
4452:
4449:
4447:
4444:
4442:
4441:Mental health
4439:
4437:
4434:
4432:
4429:
4427:
4424:
4422:
4419:
4417:
4414:
4412:
4409:
4407:
4404:
4402:
4399:
4397:
4394:
4392:
4389:
4388:
4386:
4384:
4380:
4374:
4371:
4369:
4366:
4364:
4360:
4356:
4353:
4351:
4347:
4344:
4342:
4339:
4337:
4334:
4332:
4329:
4327:
4324:
4322:
4319:
4317:
4316:Mate guarding
4314:
4312:
4309:
4307:
4304:
4302:
4299:
4297:
4294:
4292:
4289:
4287:
4284:
4282:
4279:
4277:
4276:Age disparity
4274:
4272:
4269:
4267:
4264:
4263:
4261:
4259:
4255:
4247:
4244:
4242:
4239:
4237:
4234:
4232:
4229:
4228:
4227:
4224:
4222:
4219:
4217:
4214:
4212:
4209:
4207:
4206:Schizophrenia
4204:
4202:
4199:
4197:
4194:
4192:
4189:
4187:
4184:
4182:
4179:
4177:
4174:
4172:
4169:
4165:
4162:
4160:
4157:
4155:
4152:
4150:
4147:
4145:
4142:
4141:
4140:
4137:
4136:
4134:
4132:
4131:Mental health
4127:
4126:Human factors
4123:
4117:
4116:Socialization
4114:
4112:
4109:
4107:
4104:
4102:
4099:
4097:
4094:
4092:
4089:
4087:
4084:
4082:
4079:
4077:
4076:paternal bond
4073:
4069:
4066:
4064:
4061:
4059:
4056:
4055:
4053:
4051:
4047:
4041:
4038:
4034:
4031:
4030:
4029:
4026:
4022:
4019:
4018:
4017:
4014:
4010:
4007:
4005:
4002:
4000:
3997:
3996:
3995:
3992:
3990:
3987:
3983:
3980:
3979:
3978:
3975:
3971:
3968:
3966:
3963:
3962:
3961:
3958:
3957:
3955:
3953:
3949:
3941:
3940:NaĂŻve physics
3938:
3936:
3933:
3931:
3928:
3927:
3926:
3923:
3921:
3918:
3916:
3913:
3911:
3907:
3906:Motor control
3904:
3900:
3897:
3895:
3892:
3891:
3890:
3887:
3885:
3881:
3878:
3876:
3872:
3869:
3865:
3864:Ophidiophobia
3862:
3860:
3857:
3855:
3854:Arachnophobia
3852:
3851:
3850:
3847:
3845:
3842:
3840:
3837:
3835:
3832:
3828:
3825:
3823:
3820:
3818:
3815:
3813:
3810:
3808:
3805:
3804:
3803:
3799:
3796:
3794:
3791:
3787:
3784:
3782:
3781:Display rules
3779:
3777:
3774:
3773:
3772:
3769:
3768:
3766:
3764:
3759:
3755:
3752:
3748:
3742:
3739:
3735:
3732:
3730:
3727:
3725:
3721:
3718:
3716:
3713:
3712:
3711:
3708:
3704:
3701:
3700:
3699:
3696:
3694:
3691:
3689:
3686:
3684:
3683:Kin selection
3681:
3677:
3674:
3673:
3672:
3669:
3667:
3664:
3662:
3659:
3657:
3654:
3652:
3649:
3645:
3642:
3640:
3637:
3635:
3632:
3631:
3630:
3627:
3625:
3622:
3621:
3619:
3617:
3611:
3605:
3602:
3598:
3595:
3593:
3590:
3588:
3585:
3583:
3580:
3578:
3577:Adaptationism
3575:
3574:
3573:
3570:
3566:
3563:
3562:
3561:
3558:
3557:
3554:
3550:
3543:
3538:
3536:
3531:
3529:
3524:
3523:
3520:
3508:
3504:
3500:
3498:
3490:
3489:
3486:
3480:
3477:
3475:
3472:
3470:
3467:
3465:
3462:
3458:
3455:
3454:
3453:
3452:Phylogenetics
3450:
3448:
3445:
3443:
3440:
3438:
3435:
3433:
3430:
3426:
3423:
3421:
3418:
3416:
3413:
3412:
3411:
3408:
3406:
3403:
3401:
3398:
3396:
3393:
3392:
3390:
3386:
3380:
3377:
3373:
3370:
3368:
3365:
3361:
3358:
3357:
3356:
3355:Structuralism
3353:
3351:
3348:
3346:
3343:
3341:
3338:
3336:
3333:
3331:
3330:Catastrophism
3328:
3327:
3326:
3323:
3321:
3318:
3317:
3315:
3311:
3305:
3302:
3300:
3297:
3295:
3292:
3290:
3289:Neo-Darwinism
3287:
3285:
3282:
3280:
3277:
3275:
3272:
3270:
3267:
3265:
3262:
3258:
3257:
3253:
3252:
3251:
3248:
3244:
3243:
3239:
3238:
3237:
3234:
3232:
3229:
3227:
3224:
3223:
3221:
3219:
3215:
3209:
3206:
3204:
3203:Reinforcement
3201:
3199:
3196:
3194:
3191:
3189:
3186:
3184:
3181:
3179:
3176:
3174:
3171:
3169:
3166:
3164:
3161:
3159:
3156:
3154:
3151:
3150:
3148:
3146:
3142:
3136:
3135:Catastrophism
3132:
3129:
3127:
3126:Macromutation
3123:
3122:Micromutation
3120:
3118:
3114:
3110:
3107:
3106:
3104:
3102:
3098:
3092:
3089:
3085:
3082:
3080:
3077:
3075:
3072:
3070:
3067:
3065:
3062:
3061:
3060:
3057:
3055:
3052:
3050:
3047:
3045:
3042:
3040:
3037:
3035:
3032:
3030:
3029:Immune system
3027:
3025:
3022:
3020:
3017:
3015:
3012:
3010:
3007:
3003:
3000:
2999:
2998:
2995:
2993:
2990:
2988:
2985:
2983:
2980:
2976:
2973:
2971:
2968:
2967:
2966:
2963:
2962:
2960:
2958:
2953:
2945:
2942:
2940:
2937:
2935:
2932:
2930:
2927:
2925:
2922:
2921:
2919:
2915:
2912:
2910:
2907:
2905:
2902:
2900:
2897:
2895:
2892:
2890:
2889:symbiogenesis
2887:
2886:
2885:
2882:
2880:
2877:
2875:
2872:
2870:
2867:
2866:
2864:
2862:
2857:
2851:
2848:
2846:
2843:
2841:
2838:
2836:
2833:
2829:
2826:
2825:
2824:
2821:
2817:
2814:
2810:
2807:
2805:
2802:
2801:
2800:
2797:
2795:
2792:
2790:
2787:
2785:
2782:
2780:
2777:
2773:
2770:
2768:
2765:
2764:
2763:
2760:
2758:
2755:
2754:
2753:
2750:
2748:
2745:
2741:
2738:
2737:
2736:
2733:
2731:
2728:
2726:
2723:
2721:
2718:
2714:
2711:
2710:
2709:
2706:
2704:
2701:
2697:
2694:
2693:
2692:
2689:
2687:
2684:
2683:
2681:
2679:
2674:
2668:
2665:
2663:
2660:
2658:
2655:
2653:
2650:
2648:
2645:
2643:
2640:
2639:
2637:
2633:
2627:
2624:
2622:
2619:
2617:
2614:
2610:
2607:
2605:
2602:
2600:
2597:
2596:
2595:
2592:
2590:
2587:
2585:
2582:
2580:
2577:
2575:
2572:
2568:
2565:
2563:
2560:
2559:
2558:
2557:Kin selection
2555:
2553:
2552:Genetic drift
2550:
2548:
2545:
2541:
2538:
2537:
2536:
2533:
2531:
2528:
2526:
2523:
2521:
2518:
2516:
2513:
2512:
2510:
2508:
2502:
2494:
2491:
2490:
2489:
2486:
2484:
2481:
2477:
2474:
2472:
2469:
2468:
2467:
2464:
2460:
2457:
2456:
2455:
2452:
2450:
2447:
2445:
2442:
2440:
2437:
2435:
2432:
2430:
2427:
2425:
2422:
2420:
2417:
2415:
2412:
2410:
2407:
2403:
2400:
2399:
2398:
2395:
2393:
2390:
2388:
2385:
2383:
2380:
2378:
2375:
2373:
2370:
2368:
2365:
2363:
2360:
2358:
2355:
2351:
2348:
2347:
2346:
2343:
2341:
2338:
2336:
2333:
2329:
2326:
2324:
2321:
2320:
2319:
2316:
2314:
2311:
2309:
2306:
2304:
2301:
2300:
2298:
2296:
2292:
2286:
2283:
2281:
2278:
2276:
2273:
2271:
2268:
2266:
2263:
2262:
2259:
2255:
2248:
2243:
2241:
2236:
2234:
2229:
2228:
2225:
2214:
2210:
2206:
2202:
2198:
2194:
2190:
2186:
2179:
2171:
2167:
2163:
2159:
2154:
2149:
2145:
2141:
2137:
2133:
2129:
2125:
2118:
2109:
2104:
2100:
2096:
2092:
2085:
2083:
2074:
2070:
2065:
2060:
2056:
2052:
2048:
2044:
2040:
2033:
2031:
2022:
2018:
2013:
2008:
2003:
1998:
1994:
1990:
1986:
1982:
1978:
1971:
1963:
1959:
1954:
1949:
1945:
1941:
1937:
1933:
1929:
1922:
1920:
1911:
1907:
1903:
1899:
1895:
1891:
1887:
1883:
1879:
1875:
1868:
1860:
1856:
1852:
1848:
1844:
1840:
1836:
1829:
1821:
1817:
1813:
1809:
1805:
1801:
1794:
1786:
1782:
1778:
1774:
1770:
1766:
1762:
1758:
1751:
1743:
1739:
1735:
1731:
1728:(53): 19â37.
1727:
1723:
1716:
1714:
1705:
1701:
1697:
1693:
1689:
1685:
1681:
1674:
1672:
1663:
1659:
1654:
1649:
1645:
1641:
1637:
1633:
1629:
1625:
1621:
1614:
1606:
1602:
1597:
1592:
1588:
1584:
1580:
1576:
1572:
1568:
1564:
1557:
1549:
1545:
1540:
1535:
1531:
1527:
1523:
1519:
1515:
1511:
1507:
1503:
1499:
1492:
1490:
1481:
1474:
1466:
1462:
1458:
1454:
1450:
1446:
1439:
1431:
1427:
1420:
1418:
1416:
1414:
1405:
1401:
1396:
1391:
1387:
1383:
1379:
1375:
1371:
1364:
1362:
1353:
1349:
1344:
1339:
1335:
1331:
1327:
1323:
1319:
1312:
1310:
1308:
1299:
1295:
1291:
1287:
1283:
1279:
1272:
1265:
1263:
1261:
1259:
1250:
1246:
1242:
1238:
1234:
1230:
1223:
1221:
1219:
1210:
1206:
1202:
1198:
1194:
1190:
1186:
1182:
1175:
1168:
1166:
1164:
1155:
1151:
1146:
1141:
1137:
1133:
1129:
1122:
1120:
1118:
1116:
1114:
1112:
1103:
1099:
1095:
1091:
1087:
1083:
1076:
1074:
1072:
1070:
1061:
1057:
1053:
1049:
1045:
1041:
1037:
1033:
1029:
1025:
1017:
1015:
1013:
1011:
1002:
998:
994:
990:
986:
982:
978:
974:
970:
966:
959:
957:
955:
953:
951:
949:
940:
936:
932:
928:
924:
920:
916:
912:
905:
897:
893:
888:
883:
878:
873:
869:
865:
861:
857:
853:
846:
838:
834:
830:
826:
822:
818:
814:
807:
805:
796:
792:
788:
784:
777:
768:
763:
759:
755:
751:
747:
743:
736:
734:
725:
721:
716:
711:
707:
703:
699:
695:
691:
684:
682:
680:
678:
669:
665:
661:
657:
653:
649:
645:
641:
637:
633:
626:
618:
614:
610:
606:
602:
598:
594:
590:
583:
575:
571:
566:
561:
557:
553:
549:
545:
541:
534:
530:
522:
519:
517:
513:
508:
497:
487:
483:
480:
479:
467:
464:
460:
456:
453:
452:
446:
444:
435:
426:
424:
423:
417:
415:
414:
409:
408:
401:
399:
395:
394:superorganism
386:
382:
377:
367:
364:
363:P. aeruginosa
360:
355:
354:P. aeruginosa
350:
347:
343:
342:
336:
327:
324:
320:
315:
311:
306:
304:
300:
296:
295:
290:
286:
282:
278:
274:
273:
268:
253:
250:
246:
236:
233:
228:
227:ectoparasites
224:
220:
216:
203:
199:
197:
186:
184:
180:
179:D. discoideum
175:
174:D. discoideum
170:
166:
165:fruiting body
162:
158:
154:
150:
146:
145:
138:
132:
130:
126:
122:
118:
117:
112:
102:
93:
89:
86:
80:
77:
72:
69:
65:
59:
57:
47:
45:
41:
36:
32:
28:
22:
4873:
4860:
4847:
4834:
4593:Sociobiology
4451:Neuroscience
4431:Intelligence
3977:Anthropology
3930:Color vision
3915:Multitasking
3894:Flynn effect
3889:Intelligence
3871:Folk biology
3633:
3614:Evolutionary
3464:Polymorphism
3447:Astrobiology
3395:Biogeography
3350:Saltationism
3340:Orthogenesis
3325:Alternatives
3254:
3240:
3173:Cospeciation
3168:Cladogenesis
3117:Saltationism
3074:Mating types
2997:Color vision
2982:Avian flight
2904:mitochondria
2642:Canalisation
2520:Biodiversity
2322:
2265:Introduction
2188:
2184:
2178:
2127:
2123:
2117:
2098:
2094:
2046:
2042:
1984:
1980:
1970:
1935:
1931:
1877:
1873:
1867:
1845:(1): 80â86.
1842:
1838:
1828:
1803:
1799:
1793:
1760:
1756:
1750:
1725:
1721:
1687:
1683:
1627:
1623:
1613:
1570:
1566:
1556:
1505:
1501:
1479:
1473:
1448:
1444:
1438:
1429:
1425:
1377:
1373:
1328:(2): 162â5.
1325:
1321:
1281:
1277:
1232:
1228:
1184:
1180:
1135:
1131:
1088:(2): 48â54.
1085:
1082:Trends Genet
1081:
1027:
1023:
968:
964:
914:
910:
904:
859:
855:
845:
820:
816:
786:
782:
776:
749:
745:
697:
693:
635:
631:
625:
592:
588:
582:
547:
543:
533:
520:
509:
492:
476:
473:
457:
449:
447:
440:
420:
418:
411:
405:
402:
391:
379:
362:
353:
351:
339:
337:
333:
322:
318:
313:
309:
307:
302:
292:
288:
284:
276:
270:
267:siderophores
264:
242:
218:
213:
192:
178:
173:
168:
160:
142:
141:
136:
125:cleaner fish
114:
107:
90:
81:
75:
73:
60:
53:
26:
25:
4732:Determinism
4644:Coevolution
4588:Primatology
4426:Gender role
4331:Orientation
4211:Screen time
4068:Affectional
4050:Development
3729:Mate choice
3656:By-products
3624:Adaptations
3587:Cognitivism
3474:Systematics
3345:Mutationism
3163:Catagenesis
3091:Snake venom
3024:Eusociality
3002:in primates
2992:Cooperation
2920:In animals
2740:butterflies
2713:Cephalopods
2703:Brachiopods
2635:Development
2609:Mate choice
2362:Convergence
2345:Coevolution
2303:Abiogenesis
1132:J Evol Biol
516:yucca moths
299:coelichelin
111:cooperation
85:lek paradox
4891:Categories
4679:Population
4674:Lamarckism
4520:behavioral
4498:Behavioral
4446:Narcissism
4391:Aggression
4181:Hypophobia
4171:Depression
4058:Attachment
4040:Universals
4004:Psychology
3982:Biological
3970:Musicology
3960:Aesthetics
3859:Basophobia
3666:Exaptation
3644:Reciprocal
3335:Lamarckism
3313:Philosophy
3236:David Hume
3198:Peripatric
3193:Parapatric
3178:Ecological
3158:Anagenesis
3153:Allopatric
3145:Speciation
3109:Gradualism
3034:Metabolism
2894:chromosome
2884:Eukaryotes
2662:Modularity
2579:Population
2505:Population
2466:Speciation
2444:Panspermia
2397:Extinction
2392:Exaptation
2367:Divergence
2340:Cladistics
2328:Reciprocal
2308:Adaptation
862:(1): 124.
525:References
281:pyoverdine
4524:cognitive
4516:Affective
4401:Cognition
4355:Sexuality
4341:Pair bond
4101:Education
3758:Cognition
3676:Inclusive
3616:processes
3604:Criticism
3469:Protocell
3320:Darwinism
3208:Sympatric
2957:processes
2845:Tetrapods
2794:Kangaroos
2720:Dinosaurs
2657:Inversion
2626:Variation
2547:Gene flow
2540:Inclusive
2350:Mutualism
2295:Evolution
1587:1744-9561
1530:0099-2240
1322:Biol Lett
817:Evolution
496:rhizobium
56:Mutualism
4897:Behavior
4793:Memetics
4553:Ethology
4511:genetics
4346:Physical
4311:Jealousy
4266:Activity
4072:maternal
4028:Religion
4016:Morality
3994:Language
3875:taxonomy
3688:Mismatch
3634:Cheating
3629:Altruism
3497:Category
3372:Vitalism
3367:Theistic
3360:Spandrel
3044:Morality
3039:Monogamy
2914:plastids
2879:Flagella
2835:Reptiles
2816:sea cows
2799:primates
2708:Molluscs
2686:Bacteria
2574:Mutation
2507:genetics
2483:Taxonomy
2429:Mismatch
2409:Homology
2323:Cheating
2318:Altruism
2162:12955144
2153:1842/469
2073:17148216
1962:24990673
1859:16813247
1820:43077376
1742:17600462
1704:16332224
1662:23066081
1605:25694506
1548:25724953
1482:: 45â60.
1465:35783869
1404:15504003
1352:17148155
1298:53193030
1249:53181083
1201:24952103
1154:15312076
1102:21167620
1052:18272966
993:11140681
939:37319983
931:10640384
896:19486536
837:19619222
724:16901844
660:16724064
617:18451640
609:16845430
574:11958708
413:Diacamma
232:cortisol
121:eusocial
96:Examples
35:ethology
27:Cheating
4694:Species
4466:Suicide
4301:Fantasy
4281:Arousal
4063:Bonding
3952:Culture
3776:Display
3763:Emotion
3671:Fitness
3560:History
3388:Related
3218:History
3079:Meiosis
3014:Empathy
3009:Emotion
2909:nucleus
2850:Viruses
2840:Spiders
2752:Mammals
2735:Insects
2535:Fitness
2471:Species
2270:Outline
2213:4330563
2193:Bibcode
2170:4363192
2132:Bibcode
2064:1626376
2021:1465451
1989:Bibcode
1953:4100499
1910:4321146
1902:1614546
1882:Bibcode
1785:4366903
1765:Bibcode
1653:3587168
1632:Bibcode
1624:Science
1596:4360104
1539:4393426
1510:Bibcode
1395:1810046
1343:1626222
1209:3188654
1060:4341839
1032:Bibcode
1001:4307980
973:Bibcode
887:2701933
864:Bibcode
783:Ecology
754:Bibcode
715:1635526
668:4419943
640:Bibcode
565:1690960
153:haploid
68:fitness
4872:
4859:
4846:
4436:Memory
4396:Autism
4363:female
4296:Desire
4033:Origin
4009:Speech
3999:Origin
3771:Affect
3507:Portal
3183:Hybrid
3019:Ethics
2861:organs
2823:Plants
2809:lemurs
2804:humans
2789:horses
2779:hyenas
2767:wolves
2762:canids
2696:origin
2211:
2185:Nature
2168:
2160:
2124:Nature
2071:
2061:
2019:
2009:
1960:
1950:
1908:
1900:
1874:Nature
1857:
1818:
1783:
1757:Nature
1740:
1702:
1660:
1650:
1603:
1593:
1585:
1546:
1536:
1528:
1463:
1402:
1392:
1350:
1340:
1296:
1247:
1207:
1199:
1152:
1100:
1058:
1050:
1024:Nature
999:
991:
965:Nature
937:
929:
894:
884:
835:
722:
712:
666:
658:
632:Nature
615:
607:
572:
562:
512:yuccas
4406:Crime
3989:Crime
3920:Sleep
3910:skill
3750:Areas
2970:Death
2965:Aging
2944:brain
2730:Fungi
2691:Birds
2604:Fungi
2402:Event
2285:Index
2209:S2CID
2166:S2CID
2012:50713
1906:S2CID
1855:S2CID
1816:S2CID
1781:S2CID
1461:S2CID
1294:S2CID
1274:(PDF)
1245:S2CID
1205:S2CID
1177:(PDF)
1056:S2CID
997:S2CID
935:S2CID
664:S2CID
613:S2CID
352:When
239:Other
4359:male
3720:Male
3457:Tree
2929:hair
2869:Cell
2772:dogs
2757:cats
2747:Life
2725:Fish
2678:taxa
2158:PMID
2069:PMID
2017:PMID
1958:PMID
1898:PMID
1738:PMID
1700:PMID
1658:PMID
1601:PMID
1583:ISSN
1544:PMID
1526:ISSN
1400:PMID
1348:PMID
1197:PMID
1150:PMID
1098:PMID
1048:PMID
989:PMID
927:PMID
892:PMID
833:PMID
720:PMID
656:PMID
605:PMID
570:PMID
514:and
321:and
33:and
4258:Sex
3935:Eye
2955:Of
2924:eye
2874:DNA
2859:Of
2676:Of
2201:doi
2189:372
2148:hdl
2140:doi
2128:425
2103:doi
2059:PMC
2051:doi
2007:PMC
1997:doi
1948:PMC
1940:doi
1936:281
1890:doi
1878:358
1847:doi
1808:doi
1773:doi
1761:342
1730:doi
1692:doi
1648:PMC
1640:doi
1628:338
1591:PMC
1575:doi
1534:PMC
1518:doi
1453:doi
1390:PMC
1382:doi
1378:271
1338:PMC
1330:doi
1286:doi
1237:doi
1189:doi
1140:doi
1090:doi
1040:doi
1028:451
981:doi
969:408
919:doi
882:PMC
872:doi
825:doi
791:doi
762:doi
710:PMC
702:doi
698:273
648:doi
636:441
597:doi
560:PMC
552:doi
548:269
463:sex
135:In
4893::
2207:.
2199:.
2187:.
2164:.
2156:.
2146:.
2138:.
2126:.
2099:15
2097:.
2093:.
2081:^
2067:.
2057:.
2045:.
2041:.
2029:^
2015:.
2005:.
1995:.
1985:89
1983:.
1979:.
1956:.
1946:.
1934:.
1930:.
1918:^
1904:.
1896:.
1888:.
1876:.
1853:.
1843:58
1841:.
1837:.
1814:.
1804:46
1802:.
1779:.
1771:.
1759:.
1736:.
1726:53
1724:.
1712:^
1698:.
1688:51
1686:.
1682:.
1670:^
1656:.
1646:.
1638:.
1626:.
1622:.
1599:.
1589:.
1581:.
1571:11
1569:.
1565:.
1542:.
1532:.
1524:.
1516:.
1506:81
1504:.
1500:.
1488:^
1459:.
1449:63
1447:.
1430:84
1428:.
1412:^
1398:.
1388:.
1376:.
1372:.
1360:^
1346:.
1336:.
1324:.
1320:.
1306:^
1292:.
1282:78
1280:.
1276:.
1257:^
1243:.
1233:63
1231:.
1217:^
1203:.
1195:.
1185:66
1183:.
1179:.
1162:^
1148:.
1136:17
1134:.
1130:.
1110:^
1096:.
1086:27
1084:.
1068:^
1054:.
1046:.
1038:.
1026:.
1009:^
995:.
987:.
979:.
967:.
947:^
933:.
925:.
915:59
913:.
890:.
880:.
870:.
858:.
854:.
831:.
821:63
819:.
803:^
787:79
785:.
760:.
748:.
744:.
732:^
718:.
708:.
696:.
692:.
676:^
662:.
654:.
646:.
634:.
611:.
603:.
591:.
568:.
558:.
546:.
542:.
185:.
131:.
119:;
4734:/
4526:/
4522:/
4518:/
4509:/
4500:/
4361:/
4357:/
4348:/
4128:/
4074:/
4070:/
3908:/
3882:/
3873:/
3800:/
3760:/
3722:/
3541:e
3534:t
3527:v
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