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96-451: Toxotrypana is an obsolete genus of tephritid or fruit flies in the family Tephritidae , now Anastrepha This Trypetinae -related article is a stub . You can help Misplaced Pages by expanding it . Tephritidae The Tephritidae are one of two fly families referred to as fruit flies , the other family being the Drosophilidae . The family Tephritidae does not include

192-458: A brood parasite in that it attacks and enslaves other species within their subgenus, Alpinobombus to propagate their population. Various types of mating systems include monogamy , polygyny , polyandry , and promiscuity . Each is differentiated by the sexual behavior between mates, such as which males mate with certain females. An influential paper by Stephen Emlen and Lewis Oring (1977) argued that two main factors of animal behavior influence

288-457: A bumblebee that relies on host workers of various other Bombus species. Similarly, in Eulaema meriana , some Leucospidae wasps exploit the brood cells and nest for shelter and food from the bees. Vespula austriaca is another wasp in which the females force the host workers to feed and take care of the brood. In particular, Bombus hyperboreus , an Arctic bee species, is also classified as

384-425: A current of water that passed over the spermatophores and towards the female. Sperm packet uptake by the female would sometimes follow. Heather Proctor hypothesised that the vibrations trembling male legs made were done to mimic the vibrations that females detect from swimming prey - this would trigger the female prey-detection responses causing females to orient and then clutch at males, mediating courtship. If this

480-400: A dorsal preapical bristle. The female has an oviscape. The larva is amphipneustic (having only the anterior and posterior pairs of spiracle). The body varies from white to yellowish or brown. The posterior end of pale-coloured species is sometimes black. The body tapers at the anterior. The two mandibles sometimes have teeth along the ventral margin. The antennomaxillary lobes at each side of

576-434: A female to pass through. Big males are, therefore, more successful in mating because they claim territories near the female nesting sites that are more sought after. Smaller males, on the other hand, monopolize less competitive sites in foraging areas so that they may mate with reduced conflict. Another example of this is Sepsis cynipsea , where males of the species mount females to guard them from other males and remain on

672-450: A genetic level. Such 'choosiness' from the female individuals can be seen in wasp species too, especially among Polistes dominula wasps. The females tend to prefer males with smaller, more elliptically shaped spots than those with larger and more irregularly shaped spots. Those males would have reproductive superiority over males with irregular spots. In marbled newts , females show preference to mates with larger crests. This however,

768-472: A given sexual encounter, it benefits the male to mate, but benefits the female to be choosy and resist. For example, male small tortoiseshell butterfly compete to gain the best territory to mate. Another example of this conflict can be found in the Eastern carpenter bee, Xylocopa virginica . Males of this species are limited in reproduction primarily by access to mates, so they claim a territory and wait for

864-454: A humeral and a subcostal break. The apical part of the subcostal is usually indistinct or even transparent and at about a right angle with respect to the basal part. Crossvein BM-Cu is present; the cell cup (posterior cubital cell or anal cell) is closed and nearly always narrowing to an acute angle. It is closed by a geniculated vein (CuA2). The CuA2 vein is rarely straight or convex. The tibiae lack

960-413: A male finds a female, he slowly circles around the female whilst trembling his first and second leg near her. Male leg trembling causes females (who were in the 'net stance') to orient towards often clutch the male. This did not damage the male or deter further courtship; the male then deposited spermatophores and began to vigorously fan and jerk his fourth pair of legs over the spermatophore, generating

1056-524: A mating context, both sexes prefer animate orange objects, which suggests that preference originally evolved in another context, like foraging. 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

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1152-582: A pest of sunflowers and Rhagoletis mendax , a pest of blueberries. Another notorious agricultural pest is the Mediterranean fruit fly or Medfly, Ceratitis capitata , which is responsible for millions of dollars' worth in expenses by countries for control and eradication efforts, in addition to costs of damage to fruit crops. Similarly, the Queensland fruit fly ( Bactrocera tryoni ) is responsible for more than $ 28.5 million in damage to Australian fruit crops

1248-458: 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

1344-426: 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. 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

1440-445: 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. 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

1536-521: 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

1632-452: A waxy genital plug onto the tip of the female's abdomen that physically prevents the female from mating again. Males can also prevent future mating by transferring an anti-Aphrodiasic to the female during mating. This behavior is seen in butterfly species such as Heliconius melpomene , where males transfer a compound that causes the female to smell like a male butterfly and thus deter any future potential mates. Furthermore, males may control

1728-400: A year. This species lays eggs in a wide variety of unripe fruit hosts, causing them to rot prior to ripening. Some fruit flies are used as agents of biological control , thereby reducing the populations of pest species. Several species of the genus Urophora are used as control agents against rangeland-destroying noxious weeds such as starthistles and knapweeds , but their effectiveness

1824-419: Is a type of behavioral negotiation between parents that leads to stabilized compensation. Sexual conflicts can give rise to antagonistic co-evolution between the sexes to try to get the other sex to care more for offspring. For example, in the waltzing fly Prochyliza xanthostoma , ejaculate feeding maximizes female reproductive success and minimizes the female's chance of mating multiply. Evidence suggests that

1920-475: Is asynchronous hatching in birds. A behavioral ecology hypothesis is known as Lack's brood reduction hypothesis (named after David Lack ). Lack's hypothesis posits an evolutionary and ecological explanation as to why birds lay a series of eggs with an asynchronous delay leading to nestlings of mixed age and weights. According to Lack, this brood behavior is an ecological insurance that allows the larger birds to survive in poor years and all birds to survive when food

2016-682: 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 game theoretic approaches to evolutionarily stable strategies (ESS) where the best strategy for one parent depends on the strategy adopted by the other parent. Recent research has found response matching in parents who determine how much care to invest in their offspring. Studies found that parent great tits match their partner's increased care-giving efforts with increased provisioning rates of their own. This cued parental response

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2112-412: Is divided into several subfamilies: The genera Oxyphora , Pseudorellia , and Stylia comprise 32 species, and are not included in any subfamily ( incertae sedis ). Behavioral ecology 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

2208-527: Is in the water mite Neumania papillator , an ambush predator that hunts copepods (small crustaceans) passing by in the water column. When hunting, N. papillator adopts a characteristic stance termed the 'net stance' - their first four legs are held out into the water column, with their four hind legs resting on aquatic vegetation; this allows them to detect vibrational stimuli produced by swimming prey and use this to orient towards and clutch at prey. During courtship, males actively search for females - if

2304-423: 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

2400-439: 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

2496-409: 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. 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

2592-419: Is of great interest to biologists. Some fruit flies have extensive mating rituals or territorial displays. Many are brightly colored and visually showy. Some fruit flies show Batesian mimicry , bearing the colors and markings of dangerous arthropods such as wasps or jumping spiders because it helps the fruit flies avoid predation, though the flies lack stingers . Adult tephritid fruit flies are often found on

2688-423: Is plentiful. We also see sex-ratio conflict between the queen and her workers in social hymenoptera . Because of haplodiploidy , the workers (offspring) prefer a 3:1 female to male sex allocation while the queen prefers a 1:1 sex ratio. Both the queen and the workers try to bias the sex ratio in their favor. In some species, the workers gain control of the sex ratio, while in other species, like B. terrestris ,

2784-686: Is questionable. Urophora sirunaseva produces larvae that pupate within a woody gall within the flower and disrupt seed production. Chaetorellia acrolophi is an effective biocontrol agent against knapweeds Chaetorellia australis and Chaetorellia succinea , deposit eggs into the starthistle seedheads, where their larvae consume the seeds and flower ovaries. Since economically important tephritid fruit flies exist worldwide, vast networks of researchers, several international symposia, and intensive activities on various subjects extend from ecology to molecular biology ( Tephritid Workers Database ). Pest management techniques applied to tephritid include

2880-473: Is required to reach a fuller understanding of the prevalence and mechanisms of sensory bias. Sexual conflict , in some form or another, may very well be inherent in the ways most animals reproduce. 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. This unequal investment leads, on one hand, to intense competition between males for mates and, on

2976-424: 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. 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. The sensory bias hypothesis states that

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3072-406: 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. Therefore, it seems that the willingness to invest in offspring is co-adapted to offspring demand. The lifetime parental investment

3168-455: 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

3264-755: 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. 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. 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. In other species, however, females suffer through

3360-404: 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

3456-466: 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. In some bird species, sibling rivalry is also abetted by the asynchronous hatching of eggs. In

3552-515: The biological model organisms of the genus Drosophila (in the family Drosophilidae), which is often called the "common fruit fly". Nearly 5,000 described species of tephritid fruit fly are categorized in almost 500 genera of the Tephritidae. Description , recategorization , and genetic analyses are constantly changing the taxonomy of this family. To distinguish them from the Drosophilidae,

3648-430: 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. Sibling relatedness in a brood also influences the level of sibling–sibling conflict. In a study on passerine birds, it

3744-500: The ghost moth males display in leks to attract a female mate. Additionally, it is difficult to classify them as direct competitors seeing as they put a great deal of effort into their defense of their territories before females arrive, and upon female arrival they put for the great mating displays to attract the females to their individual sites. These observations make it difficult to determine whether female or resource dispersion primarily influences male aggregation, especially in lieu of

3840-596: The larvae find their food upon emerging. The larvae develop in leaves , stems, flowers, seeds, fruits, and roots of the host plant, depending on the species. Some species are gall-forming . One exception to the phytophagous lifestyle is Euphranta toxoneura (Loew) whose larvae develop in galls formed by sawflies . The adults sometimes have a very short lifespan. Some live for less than a week. Some species are monophagous (feeding on only one plant species) others are polyphagous (feeding on several, usually related plant species). The behavioral ecology of tephritid fruit flies

3936-463: The Tephritidae are sometimes called peacock flies , in reference to their elaborate and colorful markings. The name comes from the Greek τεφρος, tephros , meaning "ash grey". They are found in all the biogeographic realms . For terms see Morphology of Diptera and Tephritidae glossary Tephritids are small to medium-sized (2.5–10 mm) flies that are often colourful, and usually with pictured wings,

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4032-430: The ants into believing that they are ant larvae, causing the ants to bring the butterfly larvae back to their own nests to feed them. Other examples of brood parasites are Polistes sulcifer , a paper wasp that has lost the ability to build its own nests so females lay their eggs in the nest of a host species, Polistes dominula , and rely on the host workers to take care of their brood, as well as Bombus bohemicus ,

4128-428: The apparent difficulty that males may have defending resources and females in such densely populated areas. Because the reason for male aggregation into leks is unclear, five hypotheses have been proposed. These postulates propose the following as reasons for male lekking: hotspot, predation reduction, increased female attraction, hotshot males, facilitation of female choice. With all of the mating behaviors discussed,

4224-445: 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. The value of a social behavior depends in part on the social behavior of an animal's neighbors. For example,

4320-424: The begging display. False gapes from brood parasite offspring cause host parents to collect more food. Another example of a brood parasite is Phengaris butterflies such as Phengaris rebeli and Phengaris arion , which differ from the cuckoo in that the butterflies do not oviposit directly in the nest of the host, an ant species Myrmica schencki . Rather, the butterfly larvae release chemicals that deceive

4416-424: 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

4512-458: 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. 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

4608-557: The cost of the parent's ability to invest in other offspring". 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. There

4704-438: 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. Resources usually include nest sites, food and protection. In some cases,

4800-440: The diversity of mating systems: the relative accessibility that each sex has to mates, and the parental desertion by either sex. In a system that does not have male parental care, resource dispersion , predation , and the effects of social living primarily influence female dispersion, which in turn influences male dispersion. Since males' primary concern is female acquisition, the males either indirectly or directly compete for

4896-416: The female is able to eject the subordinate male's sperm using cloacal contractions. 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. 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

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4992-403: 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 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

5088-403: 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,

5184-427: The female, and producing sterile parasperm to protect fertile eusperm in the female's reproductive tract. For example, the male spruce bud moth ( Zeiraphera canadensis ) secretes an accessory gland protein during mating that makes them unattractive to other males and thus prevents females from future copulation. The Rocky Mountain parnassian also exhibits this type of sexual conflict when the male butterflies deposit

5280-521: The female, attempting to copulate, until the female either shakes them off or consents to mating. Similarly the neriid fly Derocephalus angusticollis demonstrates mate guarding by using their long limbs to hold onto the female as well as push other males away during copulation. Extreme manifestations of this conflict are seen throughout nature. For example, the male Panorpa scorpionflies attempt to force copulation. Male scorpionflies usually acquire mates by presenting them with edible nuptial gifts in

5376-613: The females. In direct competition , the males are directly focused on the females. Blue-headed wrasse demonstrate the behavior in which females follow resources—such as good nest sites—and males follow the females. Conversely, species with males that exemplify indirectly competitive behavior tend towards the males' anticipation of the resources desired by females and their subsequent effort to control or acquire these resources, which helps them to achieve success with females. Grey-sided voles demonstrate indirect male competition for females. The males were experimentally observed to home in on

5472-451: The forms of salivary secretions or dead insects. However, some males attempt to force copulation by grabbing females with a specialized abdominal organ without offering a gift. Forced copulation is costly to the female as she does not receive the food from the male and has to search for food herself (costing time and energy), while it is beneficial for the male as he does not need to find a nuptial gift. In other cases, however, it pays for

5568-488: The frequencies of strategies adopted by others and are therefore frequency dependent ( frequency dependence ). 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, A. m. scutellata . The term economic defendability

5664-405: The frontal bristles are inserted on a raised tubercle. Interfrontal setulae are usually absent or represented by one or two tiny setulae near the lunula. True vibrissae are absent, but several genera have strong bristles near the vibrissal angle. The wings usually have yellow, brown, or black markings or are dark-coloured with lighter markings. In a few species, the wings are clear. The costa has both

5760-455: 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

5856-531: The genetic benefits from female mate choice . 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. 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

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5952-412: 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. An experiment conducted by Anthony Arak, where playback of synthetic calls from male natterjack toads was used to manipulate behavior of

6048-426: The host eggs and young. Other examples of brood parasites include honeyguides , cowbirds , and the large blue butterfly . 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. Other cuckoos use visual deception with their wings to exaggerate

6144-532: The host plant and feeding on pollen, nectar, rotting plant debris, or honeydew. Natural enemies include parasitoid wasps of the genera Diapriidae and Braconidae . Tephritid fruit flies are of major economic importance in agriculture . Some have negative effects, some positive. Various species of fruit flies cause damage to fruit and other plant crops. The genus Bactrocera is of worldwide notoriety for its destructive impact on agriculture. The olive fruit fly ( B. oleae ), for example, feeds on only one plant:

6240-490: 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. 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. Situations that may lead to cooperation among males include when food

6336-531: 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. 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

6432-465: 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. 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

6528-421: 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

6624-464: The males provide all of them (e.g. sedge warblers ). The females dwell in their chosen males' territories for access to these resources. The males gain ownership to the territories through male–male competition that often involves physical aggression. Only the largest and strongest males manage to defend the best quality nest sites. Females choose males by inspecting the quality of different territories or by looking at some male traits that can indicate

6720-677: The mandibles have several transverse oral ridges or short laminae directed posteriorly. The anterior spiracles (prothoracic spiracles) end bluntly and are not elongated. Each has at least three openings or up to 50 arranged transversely in one to three groups or irregularly. Each posterior spiracle (anal spiracle) lacks a clearly defined peritreme and each has three spiracular openings (in mature larvae). These are usually more or less horizontal, parallel and usually bear branched spiracular hairs in four tufts. The larvae of almost all Tephritidae are phytophagous . Females deposit eggs in living, healthy plant tissue using their telescopic ovipositors . Here,

6816-411: The more habitable territories there are to inhabit, giving females of this species a large selection of males with whom to potentially mate. Leks and choruses have also been deemed another behavior among the phenomena of male competition for females. Due to the resource-poor nature of the territories that lekking males often defend, it is difficult to categorize them as indirect competitors. For example,

6912-517: 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) recognized

7008-814: 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 . Males' reproductive successes are often limited by access to mates, whereas females' reproductive successes are more often limited by access to resources. Thus, for

7104-474: 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. Thus, the cost of increased begging enforces offspring honesty. Another resolution for parent–offspring conflict

7200-511: 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. 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

7296-430: The possible application of the concept of a Nash equilibrium to model the evolution of behavioral strategies. 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

7392-639: 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. 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

7488-444: The preference is under indirect selection. 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. An example of mate choice by genes

7584-573: The primary factors influencing differences within and between species are ecology , social conflicts, and life history differences. 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. Monogamy

7680-405: 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. 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

7776-535: The quality of resources. One example of this is with the grayling butterfly ( Hipparchia semele ), where males engage in complex flight patterns to decide who defends a particular territory. The female grayling butterfly chooses a male based on the most optimal location for oviposition . Sometimes, males leave after mating. The only resource that a male provides is a nuptial gift , such as protection or food, as seen in Drosophila subobscura . The female can evaluate

7872-412: 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. 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

7968-493: The queen has a considerable amount of control over the colony sex ratio. Lastly, there has been recent evidence regarding genomic imprinting that is a result of parent–offspring conflict. Paternal genes in offspring demand more maternal resources than maternal genes in the same offspring and vice versa. This has been shown in imprinted genes like insulin-like growth factor-II . Parents need an honest signal from their offspring that indicates their level of hunger or need, so that

8064-414: 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

8160-422: The sites with the best food in anticipation of females settling in these areas. Males of Euglossa imperialis , a non-social bee species, also demonstrate indirect competitive behavior by forming aggregations of territories, which can be considered leks, to defend fragrant-rich primary territories. The purpose of these aggregations is largely only facultative, since the more suitable fragrant-rich sites there are,

8256-621: The social male may frequently copulate with the female so as to swamp rival males' sperm. 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 sperm competition . These can include removing other male's sperm from females, displacing other male's sperm by flushing out prior inseminations with large amounts of their own sperm, creating copulatory plugs in females' reproductive tracts to prevent future matings with other males, spraying females with anti-aphrodisiacs to discourage other males from mating with

8352-744: The sperm evolved to prevent female waltzing flies from mating multiply in order to ensure the male's paternity. According to Robert Trivers's theory on relatedness, 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

8448-550: The strategic allocation of sperm, producing more sperm when females are more promiscuous. All these methods are meant to ensure that females are more likely to produce offspring belonging to the males who uses the method. Females also control the outcomes of matings, and there exists the possibility that females choose sperm (cryptic female choice). A dramatic example of this is the feral fowl Gallus gallus . In this species, females prefer to copulate with dominant males, but subordinate males can force matings. In these cases,

8544-432: The subcostal vein curving forward at a right angle. The head is hemispherical and usually short. The face is vertical or retreating and the frons is broad. Ocelli and cellar bristles are present. The postvertical bristles are parallel to divergent. Two to eight pairs of frontal bristles are seen (at least one but usually several lower pairs curving inwards and at least one of the upper pairs curving backwards). In some species,

8640-496: 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. 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

8736-420: The use of cover sprays with conventional pesticides , however, due to deleterious impact of these pesticides, new, less impactful and more targeted pest control techniques have been used, such as toxic food baits , male annihilation technique using specific male attractant parapheromones in toxic baits or mass trapping , or even sterile insect technique as part of integrated pest management . Tephritidae

8832-504: The wild or commercially cultivated olive , Olea europaea . It has the capacity to ruin 100% of an olive crop by damaging the fruit. Bactrocera dorsalis is another highly invasive pest species that damages tropical fruit, vegetable, and nut crops. Euleia heraclei is a pest of celery and parsnips. The genus Anastrepha includes several important pests, notably A. grandis , A. ludens (Mexican fruit fly), A. obliqua , and A. suspensa . Other pests are Strauzia longipennis ,

8928-441: The young, such as in marmosets . In fish there is no parental care in 79% of bony fish . In fish with parental care, it usually limited to selecting, preparing, and defending a nest, as seen in sockeye salmon , for example. Also, parental care in fish, if any, is primarily done by males, as seen in gobies and redlip blennies . The cichlid fish V. moorii exhibits biparental care. In species with internal fertilization,

9024-414: 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. Territorial behavior arises when benefits are greater than the costs. Studies of the golden-winged sunbird have validated the concept of economic defendability. Comparing the energetic costs

9120-430: Was found that chicks begged more loudly in species with higher levels of extra-pair paternity . Some animals deceive other species into providing all parental care. These brood parasites selfishly exploit their hosts' parents and host offspring. The common cuckoo is a well known example of a brood parasite. Female cuckoos lay a single egg in the nest of the host species and when the cuckoo chick hatches, it ejects all

9216-440: Was true and males were exploiting female predation responses, then hungry females should be more receptive to male trembling – Proctor found that unfed captive females did orient and clutch at males significantly more than fed captive females did, consistent with the sensory exploitation hypothesis. Other examples for the sensory bias mechanism include traits in auklets , wolf spiders , and manakins . Further experimental work

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