Behavioral ecology
Behavioral ecology, also spelled behavioural ecology, is the study of the
If an organism has a trait that provides a selective advantage (i.e., has adaptive significance) in its environment, then natural selection favors it. Adaptive significance refers to the expression of a trait that affects fitness, measured by an individual's reproductive success. Adaptive traits are those that produce more copies of the individual's genes in future generations. Maladaptive traits are those that leave fewer. For example, if a bird that can call more loudly attracts more mates, then a loud call is an adaptive trait for that species because a louder bird mates more frequently than less loud birds—thus sending more loud-calling genes into future generations. Conversely, loud calling birds may attract the attention of predators more often, decreasing their presence in the gene pool.
Individuals are always in competition with others for limited resources, including food, territories, and mates. Conflict occurs between predators and prey, between rivals for mates, between siblings, mates, and even between parents and offspring.
Competing for resources
The value of a social behavior depends in part on the social behavior of an animal's neighbors. For example, the more likely a rival male is to back down from a threat, the more value a male gets out of making the threat. The more likely, however, that a rival will attack if threatened, the less useful it is to threaten other males. When a population exhibits a number of interacting social behaviors such as this, it can evolve a stable pattern of behaviors known as an evolutionarily stable strategy (or ESS). This term, derived from economic game theory, became prominent after John Maynard Smith (1982)[1] recognized the possible application of the concept of a Nash equilibrium to model the evolution of behavioral strategies.
Evolutionarily stable strategy
In short, evolutionary game theory asserts that only strategies that, when common in the population, cannot be "invaded" by any alternative (mutant) strategy is an ESS, and thus maintained in the population. In other words, at equilibrium every player should play the best strategic response to each other. When the game is two player and symmetric, each player should play the strategy that provides the response best for it.
Therefore, the ESS is considered the evolutionary end point subsequent to the interactions. As the fitness conveyed by a strategy is influenced by what other individuals are doing (the relative frequency of each strategy in the population), behavior can be governed not only by optimality but the frequencies of strategies adopted by others and are therefore frequency dependent (
Behavioral evolution is therefore influenced by both the physical environment and interactions between other individuals.
An example of how changes in geography can make a strategy susceptible to alternative strategies is the parasitization of the African honey bee,
Resource defense
The term economic defendability was first introduced by Jerram Brown in 1964. Economic defendability states that defense of a resource have costs, such as energy expenditure or risk of injury, as well as benefits of priority access to the resource.
Studies of the golden-winged sunbird have validated the concept of economic defendability. Comparing the energetic costs a sunbird expends in a day to the extra nectar gained by defending a territory, researchers showed that birds only became territorial when they were making a net energetic profit.[3] When resources are at low density, the gains from excluding others may not be sufficient to pay for the cost of territorial defense. In contrast, when resource availability is high, there may be so many intruders that the defender would have no time to make use of the resources made available by defense.
Sometimes the economics of resource competition favors shared defense. An example is the feeding territories of the white wagtail. The white wagtails feed on insects washed up by the river onto the bank, which acts as a renewing food supply. If any intruders harvested their territory then the prey would quickly become depleted, but sometimes territory owners tolerate a second bird, known as a satellite. The two sharers would then move out of phase with one another, resulting in decreased feeding rate but also increased defense, illustrating advantages of group living.[4]
Ideal free distribution
One of the major models used to predict the distribution of competing individuals amongst resource patches is the ideal free distribution model. Within this model, resource patches can be of variable quality, and there is no limit to the number of individuals that can occupy and extract resources from a particular patch. Competition within a particular patch means that the benefit each individual receives from exploiting a patch decreases logarithmically with increasing number of competitors sharing that resource patch. The model predicts that individuals will initially flock to higher-quality patches until the costs of crowding bring the benefits of exploiting them in line with the benefits of being the only individual on the lesser-quality resource patch. After this point has been reached, individuals will alternate between exploiting the higher-quality patches and the lower-quality patches in such a way that the average benefit for all individuals in both patches is the same. This model is ideal in that individuals have complete information about the quality of a resource patch and the number of individuals currently exploiting it, and free in that individuals are freely able to choose which resource patch to exploit.[5]
An experiment by Manfred Malinski in 1979 demonstrated that feeding behavior in three-spined sticklebacks follows an ideal free distribution. Six fish were placed in a tank, and food items were dropped into opposite ends of the tank at different rates. The rate of food deposition at one end was set at twice that of the other end, and the fish distributed themselves with four individuals at the faster-depositing end and two individuals at the slower-depositing end. In this way, the average feeding rate was the same for all of the fish in the tank.[6]
Mating strategies and tactics
As with any competition of resources, species across the animal kingdom may also engage in competitions for mating. If one considers mates or potentials mates as a resource, these sexual partners can be randomly distributed amongst resource pools within a given environment. Following the ideal free distribution model, suitors distribute themselves amongst the potential mates in an effort to maximize their chances or the number of potential matings. For all competitors, males of a species in most cases, there are variations in both the strategies and tactics used to obtain matings. Strategies generally refer to the genetically determined behaviors that can be described as conditional. Tactics refer to the subset of behaviors within a given genetic strategy. Thus it is not difficult for a great many variations in mating strategies to exist in a given environment or species.[7]
An experiment conducted by Anthony Arak, where playback of synthetic calls from male natterjack toads was used to manipulate behavior of the males in a chorus, the difference between strategies and tactics is clear. While small and immature, male natterjack toads adopted a satellite tactic to parasitize larger males. Though large males on average still retained greater reproductive success, smaller males were able to intercept matings. When the large males of the chorus were removed, smaller males adopted a calling behavior, no longer competing against the loud calls of larger males. When smaller males got larger, and their calls more competitive, then they started calling and competing directly for mates.[8]
Sexual selection
Mate choice by resources
In many sexually reproducing species, such as mammals, birds, and amphibians, females are able to bear offspring for a certain time period, during which the males are free to mate with other available females, and therefore can father many more offspring to pass on their genes. The fundamental difference between male and female reproduction mechanisms determines the different strategies each sex employs to maximize their reproductive success. For males, their reproductive success is limited by access to females, while females are limited by their access to resources. In this sense, females can be much choosier than males because they have to bet on the resources provided by the males to ensure reproductive success.[9]
Resources usually include nest sites, food and protection. In some cases, the males provide all of them (e.g.
The female can evaluate the quality of the protection or food provided by the male so as to decide whether to mate or not or how long she is willing to copulate.Mate choice by genes
When males' only contribution to offspring is their sperm, females are particularly choosy. With this high level of female choice, sexual ornaments are seen in males, where the ornaments reflect the male's social status. Two hypotheses have been proposed to conceptualize the genetic benefits from female mate choice.[9]
First, the good genes hypothesis suggests that female choice is for higher genetic quality and that this preference is favored because it increases fitness of the offspring.[14] This includes Zahavi's handicap hypothesis and Hamilton and Zuk's host and parasite arms race. Zahavi's handicap hypothesis was proposed within the context of looking at elaborate male sexual displays. He suggested that females favor ornamented traits because they are handicaps and are indicators of the male's genetic quality. Since these ornamented traits are hazards, the male's survival must be indicative of his high genetic quality in other areas. In this way, the degree that a male expresses his sexual display indicates to the female his genetic quality.[9] Zuk and Hamilton proposed a hypothesis after observing disease as a powerful selective pressure on a rabbit population. They suggested that sexual displays were indicators of resistance of disease on a genetic level.[9]
Such 'choosiness' from the female individuals can be seen in wasp species too, especially among
In marbled newts, females show preference to mates with larger crests. This however, is not considered a handicap as it does not negatively affect males' chances of survival. It is simply a trait females show preference for when choosing their mate as it is an indication of health and fitness.[15]
Fisher's hypothesis of runaway sexual selection suggests that female preference is genetically correlated with male traits and that the preference co-evolves with the evolution of that trait, thus the preference is under indirect selection.[14] Fisher suggests that female preference began because the trait indicated the male's quality. The female preference spread, so that the females' offspring now benefited from the higher quality from specific trait but also greater attractiveness to mates. Eventually, the trait only represents attractiveness to mates, and no longer represents increased survival.[9]
An example of mate choice by genes is seen in the cichlid fish Tropheus moorii where males provide no parental care. An experiment found that a female T. moorii is more likely to choose a mate with the same color morph as her own.[16] In another experiment, females have been shown to share preferences for the same males when given two to choose from, meaning some males get to reproduce more often than others.[17]
Sensory bias
The sensory bias hypothesis states that the preference for a trait evolves in a non-mating context, and is then exploited by one sex to obtain more mating opportunities. The competitive sex evolves traits that exploit a pre-existing bias that the choosy sex already possesses. This mechanism is thought to explain remarkable trait differences in closely related species because it produces a divergence in signaling systems, which leads to reproductive isolation.[18]
Sensory bias has been demonstrated in guppies, freshwater fish from Trinidad and Tobago. In this mating system, female guppies prefer to mate with males with more orange body coloration. However, outside of a mating context, both sexes prefer animate orange objects, which suggests that preference originally evolved in another context, like foraging.[19] Orange fruits are a rare treat that fall into streams where the guppies live. The ability to find these fruits quickly is an adaptive quality that has evolved outside of a mating context. Sometime after the affinity for orange objects arose, male guppies exploited this preference by incorporating large orange spots to attract females.
Another example of sensory exploitation is in the water mite
Other examples for the sensory bias mechanism include traits in auklets,[24] wolf spiders,[25] and manakins.[26] Further experimental work is required to reach a fuller understanding of the prevalence and mechanisms of sensory bias.[27]
Sexual conflict
Sexual conflict, in some form or another, may very well be inherent in the ways most animals reproduce.[28] Females invest more in offspring prior to mating, due to the differences in gametes in species that exhibit anisogamy, and often invest more in offspring after mating.[29] This unequal investment leads, on one hand, to intense competition between males for mates and, on the other hand, to females choosing among males for better access to resources and good genes. Because of differences in mating goals, males and females may have very different preferred outcomes to mating.
Sexual conflict occurs whenever the preferred outcome of mating is different for the male and female. This difference, in theory, should lead to each sex evolving adaptations that bias the outcome of reproduction towards its own interests. This sexual competition leads to sexually antagonistic coevolution between males and females, resulting in what has been described as an evolutionary arms race between males and females.[30][31]
Conflict over mating
Males' reproductive successes are often limited by access to mates, whereas females' reproductive successes are more often limited by access to resources. Thus, for a given sexual encounter, it benefits the male to mate, but benefits the female to be choosy and resist.
In other cases, however, it pays for the female to gain more matings and her social mate to prevent these so as to guard paternity. For example, in many socially monogamous birds, males follow females closely during their fertile periods and attempt to chase away any other males to prevent extra-pair matings. The female may attempt to sneak off to achieve these extra matings. In species where males are incapable of constant guarding, the social male may frequently copulate with the female so as to swamp rival males' sperm.[37]
Sexual conflict after mating has also been shown to occur in both males and females. Males employ a diverse array of tactics to increase their success in
Females also control the outcomes of matings, and there exists the possibility that females choose sperm (cryptic female choice).
Parental care and family conflicts
Parental care is the investment a parent puts into their offspring—which includes protecting and feeding the young, preparing burrows or nests, and providing eggs with yolk.[42] There is great variation in parental care in the animal kingdom. In some species, the parents may not care for their offspring at all, while in others the parents exhibit single-parental or even bi-parental care. As with other topics in behavioral ecology, interactions within a family involve conflicts. These conflicts can be broken down into three general types: sexual (male–female) conflict, parent–offspring conflict, and sibling conflict.
Types of parental care
There are many different patterns of parental care in the animal kingdom. The patterns can be explained by physiological constraints or ecological conditions, such as mating opportunities. In invertebrates, there is no parental care in most species because it is more favorable for parents to produce a large number of eggs whose fate is left to chance than to protect a few individual young. In other cases, parental care is indirect, manifested via actions taken before the offspring is produced, but nonetheless essential for their survival; for example, female Lasioglossum figueresi sweat bees excavate a nest, construct brood cells, and stock the cells with pollen and nectar before they lay their eggs, so when the larvae hatch they are sheltered and fed, but the females die without ever interacting with their brood.[43] In birds, biparental care is the most common, because reproductive success directly depends on the parents' ability to feed their chicks. Two parents can feed twice as many young, so it is more favorable for birds to have both parents delivering food. In mammals, female-only care is the most common. This is most likely because females are internally fertilized and so are holding the young inside for a prolonged period of gestation, which provides males with the opportunity to desert. Females also feed the young through lactation after birth, so males are not required for feeding. Male parental care is only observed in species where they contribute to feeding or carrying of the young, such as in marmosets.[44] In fish there is no parental care in 79% of bony fish.[45] In fish with parental care, it usually limited to selecting, preparing, and defending a nest, as seen in sockeye salmon, for example.[46] Also, parental care in fish, if any, is primarily done by males, as seen in gobies and redlip blennies.[47][42] The cichlid fish V. moorii exhibits biparental care.[48] In species with internal fertilization, the female is usually the one to take care of the young. In cases where fertilization is external the male becomes the main caretaker.
Familial conflict
Familial conflict is a result of trade-offs as a function of lifetime parental investment. Parental investment was defined by Robert Trivers in 1972 as "any investment by the parent in an individual offspring that increases the offspring's chance of surviving at the cost of the parent's ability to invest in other offspring".[citation needed] Parental investment includes behaviors like guarding and feeding. Each parent has a limited amount of parental investment over the course of their lifetime. Investment trade-offs in offspring quality and quantity within a brood and trade offs between current and future broods leads to conflict over how much parental investment to provide and to whom parents should invest in. There are three major types of familial conflict: sexual, parent–offspring, and sibling–sibling conflict.[9]
Sexual conflict
There is conflict among parents as to who should provide the care as well as how much care to provide. Each parent must decide whether or not to stay and care for their offspring, or to desert their offspring. This decision is best modeled by
Parent–offspring conflict
According to Robert Trivers's theory on relatedness,[citation needed] each offspring is related to itself by 1, but is only 0.5 related to their parents and siblings. Genetically, offspring are predisposed to behave in their own self-interest while parents are predisposed to behave equally to all their offspring, including both current and future ones. Offspring selfishly try to take more than their fair shares of parental investment, while parents try to spread out their parental investment equally amongst their present young and future young. There are many examples of parent–offspring conflict in nature. One manifestation of this is asynchronous hatching in birds. A behavioral ecology hypothesis is known as Lack's brood reduction hypothesis (named after
Parent–offspring conflict resolution
Parents need an honest signal from their offspring that indicates their level of hunger or need, so that the parents can distribute resources accordingly. Offspring want more than their fair share of resources, so they exaggerate their signals to wheedle more parental investment. However, this conflict is countered by the cost of excessive begging. Not only does excessive begging attract predators, but it also retards chick growth if begging goes unrewarded.[56] Thus, the cost of increased begging enforces offspring honesty.
Another resolution for parent–offspring conflict is that parental provisioning and offspring demand have actually coevolved, so that there is no obvious underlying conflict. Cross-fostering experiments in great tits (Parus major) have shown that offspring beg more when their biological mothers are more generous.[57] Therefore, it seems that the willingness to invest in offspring is co-adapted to offspring demand.
Sibling–sibling conflict
The lifetime parental investment is the fixed amount of parental resources available for all of a parent's young, and an offspring wants as much of it as possible. Siblings in a brood often compete for parental resources by trying to gain more than their fair share of what their parents can offer. Nature provides numerous examples in which sibling rivalry escalates to such an extreme that one sibling tries to kill off broodmates to maximize parental investment (See Siblicide). In the Galápagos fur seal, the second pup of a female is usually born when the first pup is still suckling. This competition for the mother's milk is especially fierce during periods of food shortage such as an El Niño year, and this usually results in the older pup directly attacking and killing the younger one.[58]
In some bird species, sibling rivalry is also abetted by the asynchronous hatching of eggs. In the blue-footed booby, for example, the first egg in a nest is hatched four days before the second one, resulting in the elder chick having a four-day head start in growth. When the elder chick falls 20-25% below its expected weight threshold, it attacks its younger sibling and drives it from the nest.[59]
Sibling relatedness in a brood also influences the level of sibling–sibling conflict. In a study on passerine birds, it was found that chicks begged more loudly in species with higher levels of extra-pair paternity.[60]
Brood parasitism
Some animals
Brood parasite offspring have many strategies to induce their host parents to invest parental care. Studies show that the common cuckoo uses vocal mimicry to reproduce the sound of multiple hungry host young to solicit more food.Mating systems
Various types of mating systems include
Mating systems with no male parental care
In a system that does not have male parental care,
In some other instances, neither direct nor indirect competition is seen. Instead, in species like the Edith's checkerspot butterfly, males' efforts are directed at acquisition of females and they exhibit indiscriminate mate location behavior, where, given the low cost of mistakes, they blindly attempt to mate both correctly with females and incorrectly with other objects.[80]
Mating systems with male parental care
Monogamy
Monogamy is the mating system in 90% of birds, possibly because each male and female has a greater number of offspring if they share in raising a brood.[81] In obligate monogamy, males feed females on the nest, or share in incubation and chick-feeding. In some species, males and females form lifelong pair bonds. Monogamy may also arise from limited opportunities for polygamy, due to strong competition among males for mates, females suffering from loss of male help, and female–female aggression.[82]
Polygyny
In birds, polygyny occurs when males indirectly monopolize females by controlling resources. In species where males normally do not contribute much to parental care, females suffer relatively little or not at all.[83] In other species, however, females suffer through the loss of male contribution, and the cost of having to share resources that the male controls, such as nest sites or food. In some cases, a polygynous male may control a high-quality territory so for the female, the benefits of polygyny may outweigh the costs.[84]
Polyandry threshold
There also seems to be a "polyandry threshold" where males may do better by agreeing to share a female instead of maintaining a monogamous mating system.
Female desertion and sex role reversal
In birds, desertion often happens when food is abundant, so the remaining partner is better able to raise the young unaided. Desertion also occurs if there is a great chance of a parent to gain another mate, which depends on environmental and populational factors.[87] Some birds, such as the phalaropes, have reversed sex roles where the female is larger and more brightly colored, and compete for males to incubate their clutches.[88] In jacanas, the female is larger than the male and her territory could overlap the multiple territories of up to four males.[89] In the frog species P. bibronii, the female is fertilizes multiple nests, and the male is left to tend to each nest while the female moves on.
Social behaviors
Animals
Kin selection
Kin selection refers to evolutionary strategies where an individual acts to favor the
Kin selection predicts that individuals will harbor personal costs in favor of one or multiple individuals because this can maximize their genetic contribution to future generations. For example, an organism may be inclined to expend great time and energy in
Inclusive fitness
Inclusive fitness describes the component of reproductive success in both a focal individual and their relatives.[90] Importantly, the measure embodies the sum of direct and indirect fitness and the change in their reproductive success based on the actor's behavior.[98] That is, the effect an individual's behaviors have on: being personally better-suited to reproduce offspring, and aiding descendant and non-descendant relatives in their reproductive efforts.[90] Natural selection is predicted to push individuals to behave in ways that maximize their inclusive fitness. Studying inclusive fitness is often done using predictions from Hamilton's rule.
Kin recognition
Genetic cues
One possible method of kin selection is based on genetic cues that can be recognized phenotypically.
Kin can also be recognized a genetically determined odor, as studied in the primitively social sweat bee,
Environmental cues
There are two simple rules that animals follow to determine who is kin. These rules can be exploited, but exist because they are generally successful.
The first rule is 'treat anyone in my home as kin.' This rule is readily seen in the
The second rule, named by Konrad Lorenz as 'imprinting,' states that those who you grow up with are kin. Several species exhibit this behavior, including, but not limited to the Belding's ground squirrel.[9] Experimentation with these squirrels showed that regardless of true genetic relatedness, those that were reared together rarely fought. Further research suggests that there is partially some genetic recognition going on as well, as siblings that were raised apart were less aggressive toward one another compared to non-relatives reared apart.[106]
Another way animals may recognize their kin include the interchange of unique signals. While song singing is often considered a sexual trait between males and females, male–male song singing also occurs. For example, male vinegar flies Zaprionus tuberculatus can recognize each other by song.[107]
Cooperation
Cooperation is broadly defined as behavior that provides a benefit to another individual that specifically evolved for that benefit. This excludes behavior that has not been expressly selected for to provide a benefit for another individual, because there are many commensal and parasitic relationships where the behavior one individual (which has evolved to benefit that individual and no others) is taken advantage of by other organisms. Stable cooperative behavior requires that it provide a benefit to both the actor and recipient, though the benefit to the actor can take many different forms.[9]
Within species
Within species cooperation occurs among members of the same species. Examples of intraspecific cooperation include cooperative breeding (such as in weeper capuchins) and cooperative foraging (such as in wolves). There are also forms of cooperative defense mechanisms, such as the "fighting swarm" behavior used by the stingless bee Tetragonula carbonaria.[108] Much of this behavior occurs due to kin selection. Kin selection allows cooperative behavior to evolve where the actor receives no direct benefits from the cooperation.[9]
Cooperation (without kin selection) must evolve to provide benefits to both the actor and recipient of the behavior. This includes reciprocity, where the recipient of the cooperative behavior repays the actor at a later time. This may occur in vampire bats but it is uncommon in non-human animals.[109] Cooperation can occur willingly between individuals when both benefit directly as well. Cooperative breeding, where one individual cares for the offspring of another, occurs in several species, including wedge-capped capuchin monkeys.[110]
Cooperative behavior may also be enforced, where their failure to cooperate results in negative consequences. One of the best examples of this is worker policing, which occurs in social insect colonies.[111]
The cooperative pulling paradigm is a popular experimental design used to assess if and under which conditions animals cooperate. It involves two or more animals pulling rewards towards themselves via an apparatus they can not successfully operate alone.[112]
Between species
Cooperation can occur between members of different species. For interspecific cooperation to be evolutionarily stable, it must benefit individuals in both species. Examples include pistol shrimp and goby fish, nitrogen fixing microbes and legumes,
Market economics often govern the details of the cooperation: e.g. the amount exchanged between individual animals follow the rules of supply and demand.[116]
Spite
An example of spite is the sterile soldiers of the
Another example is bacteria that release
Altruism and conflict in social insects
Many insect species of the order Hymenoptera (bees, ants, wasps) are eusocial. Within the nests or hives of social insects, individuals engage in specialized tasks to ensure the survival of the colony. Dramatic examples of these specializations include changes in body morphology or unique behaviors, such as the engorged bodies of the honeypot ant Myrmecocystus mexicanus or the waggle dance of honey bees and a wasp species, Vespula vulgaris.
In many, but not all social insects, reproduction is monopolized by the queen of the colony. Due to the effects of a
Cooperation in social organisms has numerous ecological factors that can determine the benefits and costs associated with this form of organization. One suggested benefit is a type of "life insurance" for individuals who participate in the care of the young. In this instance, individuals may have a greater likelihood of transmitting genes to the next generation when helping in a group compared to individual reproduction. Another suggested benefit is the possibility of "fortress defense", where soldier castes threaten or attack intruders, thus protecting related individuals inside the territory. Such behaviors are seen in the snapping shrimp
Conflicts in social insects
Although eusociality has been shown to offer many benefits to the colony, there is also potential for conflict. Examples include the sex-ratio conflict and
The sex-ratio conflict arises from a
According to Trivers and Hare's population-level sex-investment ratio theory, the ratio of relatedness between sexes determines the sex investment ratios.
Conflict can also arise between workers in colonies of social insects. In some species, worker females retain their ability to mate and lay eggs. The colony's queen is related to her sons by half of her genes and a quarter to the sons of her worker daughters. Workers, however, are related to their sons by half of their genes and to their brothers by a quarter. Thus, the queen and her worker daughters would compete for reproduction to maximize their own reproductive fitness. Worker reproduction is limited by other workers who are more related to the queen than their sisters, a situation occurring in many polyandrous hymenopteran species. Workers police the egg-laying females by engaging in oophagy or directed acts of aggression.[128][129]
The monogamy hypothesis
The monogamy hypothesis states that the presence of monogamy in insects is crucial for
This monogamous mating system has been observed in insects such as termites, ants, bees and wasps.[9]: 371–375 In termites the queen commits to a single male when founding a nest. In ants, bees and wasps the queens have a functional equivalent to lifetime monogamy. The male can even die before the founding of the colony. The queen can store and use the sperm from a single male throughout their lifetime, sometimes up to 30 years.[9]: 371–375
In an experiment looking at the mating of 267 hymenopteran species, the results were mapped onto a
Communication and signaling
Communication is varied at all scales of life, from interactions between microscopic organisms to those of large groups of people. Nevertheless, the signals used in communication abide by a fundamental property: they must be a quality of the receiver that can transfer information to a receiver that is capable of interpreting the signal and modifying its behavior accordingly. Signals are distinct from cues in that evolution has selected for signalling between both parties, whereas cues are merely informative to the observer and may not have originally been used for the intended purpose. The natural world is replete with examples of signals, from the luminescent flashes of light from
The nature of communication poses evolutionary concerns, such as the potential for deceit or manipulation on the part of the sender. In this situation, the receiver must be able to anticipate the interests of the sender and act appropriately to a given signal. Should any side gain advantage in the short term, evolution would select against the signal or the response. The conflict of interests between the sender and the receiver results in an evolutionarily stable state only if both sides can derive an overall benefit.
Although the potential benefits of deceit could be great in terms of mating success, there are several possibilities for how dishonesty is controlled, which include indices,
Signals are often honest, but there are exceptions. Prime examples of dishonest signals include the luminescent lure of the anglerfish, which is used to attract prey, or the mimicry of non-poisonous butterfly species, like the Batesian mimic Papilio polyxenes of the poisonous model Battus philenor.[136] Although evolution should normally favor selection against the dishonest signal, in these cases it appears that the receiver would benefit more on average by accepting the signal.
See also
- Autonomous foraging
- Behavioral plasticity
- Evolutionary models of food sharing
- Gene-centered view of evolution
- Human behavioral ecology
- Life history theory
- Marginal value theorem
- Optimization
- Mating effort
- Parental effort
- Phylogenetic comparative methods
- Selection
- Somatic effort
References
- ^ Maynard Smith, J. 1982. Evolution and the Theory of Games.
- JSTOR 4159278.
- JSTOR 1934964.
- JSTOR 4038.
- ^ Fretwell, Stephen D. (1972). Population in a Seasonal Environment. Princeton, NJ: Princeton University Press.
- .
- .
- S2CID 4363873.
- ^ ISBN 978-1-4051-1416-5.
- PMID 10722211.
- . Retrieved 13 November 2021.
- PMID 1924385.
- S2CID 53156432.
- ^ S2CID 86383459.
- ISSN 0067-8546.
- PMID 16543167.
- PMID 24293682.
- .
- PMID 11886639.
- ^ S2CID 53166756.
- ^ S2CID 54426553.
- JSTOR 2389961.
- ISBN 978-0-87893-966-4.
- .
- S2CID 24629559.
- S2CID 85039370.
- S2CID 4849390.
- PMID 16612884.
- ^ a b c d e Davies N, Krebs J, and West S. (2012). An Introduction to Behavioral Ecology, 4th Ed. Wiley-Blackwell; Oxford: pp. 209–220.[ISBN missing]
- ^ Parker, G. (1979). "Sexual selection and sexual conflict." In: Sexual Selection and Reproductive Competition in Insects (eds. M.S. Blum and N.A. Blum). Academic Press, New York: pp. 123–166.[ISBN missing]
- .
- ^ JSTOR 3480.
- S2CID 73520512.
- .
- S2CID 15548020.
- S2CID 53167007.
- ^ Birkhead, T. and Moller, A. (1992). Sperm Competition in Birds: Evolutionary Causes and Consequences. Academic Press, London.[ISBN missing][page needed]
- doi:10.1139/z94-284.
- ISBN 978-0-7748-4437-6. Retrieved 13 November 2017.
- S2CID 22090974.
- S2CID 4406792.
- ^ a b Clutton-Brock, T.H. (1991). The Evolution of Parental Care. Princeton NJ: Princeton University Press.
- S2CID 6867760.
- PMID 513786.
- .
- S2CID 53169458.
- S2CID 24806138.
- S2CID 23056207. Retrieved 30 September 2013.
- .
- ^ S2CID 16357692.
- JSTOR 3546359.
- S2CID 84884060.
- S2CID 29326420.
- PMID 11270430.
- S2CID 33682126.
- PMID 11572988.
- PMID 11416919.
- S2CID 41834534.
- ^ S2CID 53165189.
- S2CID 85105883.
- PMID 20421497.
- S2CID 30084518.
- S2CID 4379000.
- .
- S2CID 42439025.
- ^ PMC 1690087.
- ^ PMID 20585513.
- PMID 15019524.
- S2CID 7124725.
- JSTOR 25086012.
- .
- ^ Reed, H. C.; Akre, R. D.; Garnett, W. B. (1979). "A North American Host of the Yellowjacket Social Parasite Vespula austriaca (Panzer) (Hymenoptera: Vespidae)". Entomological News. 90 (2): 110–113.
- ^ Gjershaug, Jan Ove (2009). "The social parasite bumblebee Bombus hyperboreus Schönherr, 1809 usurp nest of Bombus balteatus Dahlbom, 1832 (Hymenoptera, Apidae) in Norway". Norwegian Journal of Entomology. 56 (1): 28–31.
- S2CID 16786432.
- ^ a b c d e f g Davies, N.B., Krebs, J.R. and West., S.A., (2012). An Introduction to Behavioural Ecology. 4th ed. John Wiley & Sons, pp. 254–263[ISBN missing]
- S2CID 53177154.
- JSTOR 5070.
- S2CID 53161684.
- ^ Bradbury, J. E. and Gibson, R. M. (1983) Leks and mate choice. In: Mate Choice (ed. P. Bateson). pp. 109–138. Cambridge University Press. Cambridge[ISBN missing]
- S2CID 84989304.
- ^ Lack, D. (1968) Ecological Adaptations for Breeding in Birds. Methuen, London.
- ISBN 978-1-4051-1416-5.
- S2CID 84147769.
- JSTOR 1935753.
- S2CID 83991131.
- S2CID 3334592.
- S2CID 53192930.
- ^ (Reynolds)
- S2CID 24253395.
- ^ ISBN 978-1-4051-1416-5.
- S2CID 125841.
- S2CID 4177102.
- ^ Fisher, R. A. (1930). The Genetical Theory of Natural Selection. Oxford: Clarendon Press.
- ^ Haldane, J.B.S. (1932). The Causes of Evolution. London: Longmans, Green & Co.
- ^ Haldane, J. B. S. (1955). "Population Genetics". New Biology. 18: 34–51.
- ^ S2CID 84216415.
- ^ S2CID 5310280.
- S2CID 1792464.
- S2CID 4335666.
- PMID 19285397.
- S2CID 44298800.
- PMID 19551647.
- S2CID 53147658.
- ISSN 1045-2249.
- S2CID 53191651.
- .
- S2CID 53194769.
- S2CID 44720135.
- S2CID 4354558.
- .
- PMID 19805425.
- ISBN 978-1-78378-305-2, p. 276
- ^ Postgate, J (1998). Nitrogen Fixation, 3rd Edition. Cambridge University Press, Cambridge UK.
- ^ a b Dawkins, Richard (1976). The Selfish Gene. Oxford University Press.
- .
- ^ Crair, Ben (1 August 2017). "The Secret Economic Lives of Animals". Bloomberg News. Retrieved 1 August 2017.
- ^ Foster, Kevin; Tom Wenseleers; Francis L. W. Ratnieks (10 September 2001). "Spite: Hamilton's unproven theory" (PDF). Annales Zoologici Fennici: 229–238.
- S2CID 25384748.
- S2CID 37559224.
- S2CID 2186614.
- PMID 15312076.
- S2CID 12009153.
- ^ Vespula vulgaris#Defensive behaviors
- .
- .
- S2CID 4366903.
- S2CID 16428974.
- S2CID 886746.
- S2CID 40949867.
- ^ O'Donnell, Sean (1997). "Gaster-Flagging during Colony Defense in Neotropical Swarm-Founding Wasps (Hymenoptera: Vespidae, Epiponini)". Journal of the Kansas Entomological Society.
- S2CID 15709135.
- PMID 15749110.
- ^ Tarsitano, Michael; Wolfgang, Kirchner (2001). "Vibrational courtship signals of Zygiella x-notata" (PDF). Bulletin of the British Arachnological Society. 12: 26–32.
- PMID 28568560.
Further reading
- Alcock, J. (2009). Animal Behavior: An Evolutionary Approach (9th edition). Sinauer Associates Inc. Sunderland, MA.[ISBN missing]
- Bateson, P. (2017) Behaviour, Development and Evolution. Open Book Publishers,
- Danchin, É., Girladeau, L.-A. and Cézilly, F. (2008). Behavioural Ecology: An Evolutionary Perspective on Behaviour. Oxford University Press, Oxford.[ISBN missing]
- ISBN 0-632-03546-3
- ISBN 0-86542-731-3
- Wajnberg, E., Bernstein E. and van Alphen, E. (2008). Behavioral Ecology of Insect Parasitoids – From Theoretical Approaches to Field Applications, Blackwell Publishing.[ISBN missing]
External links
- Media related to Behavioral ecology at Wikimedia Commons