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Migratory patterns and behaviors also influence sexual dimorphisms. This aspect also stems back to the size dimorphism in species. It has been shown that the larger males are better at coping with the difficulties of migration and thusly are more successful in reproducing when reaching the breeding destination.<ref>{{cite journal | last1 = Rubolini | first1 = Diego | last2 = Spina | first2 = Fernando | last3 = Saino | first3 = Nicola | year = 2004 | title = Protandry and sexual dimorphism in trans-saharan migratory birds | url = | journal = Behavioral Ecology | volume = 15 | issue = 4| pages = 592–601 | doi=10.1093/beheco/arh048}}</ref> When viewing this in an evolutionary standpoint many theories and explanations come to consideration. If these are the result for every migration and breeding season the expected results should be a shift towards a larger male population through sexual selection. Sexual selection is strong when the factor of environmental selection is also introduced. The environmental selection may support a smaller chick size if those chicks were born in an area that allowed them to grow to a larger size, even though under normal conditions they would not be able to reach this optimal size for migration. When the environment gives advantages and disadvantages of this sort, the strength of selection is weakened and the environmental forces are given greater morphological weight. The sexual dimorphism could also produce a change in timing of migration leading to differences in mating success within the bird population.<ref>{{cite journal | last1 = Kissner | first1 = K. J. | last2 = Weatherhead | first2 = P. J. | last3 = Francis | first3 = C. M. | year = 2003 | title = Sexual size dimorphism and timing of spring migration in birds | url = | journal = Journal of Evolutionary Biology | volume = 16 | issue = 1| pages = 154–162 | doi=10.1046/j.1420-9101.2003.00479.x}}</ref> When the dimorphism produces that large of a variation between the sexes and between the members of the sexes multiple evolutionary effects can take place. This timing could even lead to a speciation phenomenon if the variation becomes strongly drastic and favorable towards two different outcomes.
Migratory patterns and behaviors also influence sexual dimorphisms. This aspect also stems back to the size dimorphism in species. It has been shown that the larger males are better at coping with the difficulties of migration and thusly are more successful in reproducing when reaching the breeding destination.<ref>{{cite journal | last1 = Rubolini | first1 = Diego | last2 = Spina | first2 = Fernando | last3 = Saino | first3 = Nicola | year = 2004 | title = Protandry and sexual dimorphism in trans-saharan migratory birds | url = | journal = Behavioral Ecology | volume = 15 | issue = 4| pages = 592–601 | doi=10.1093/beheco/arh048}}</ref> When viewing this in an evolutionary standpoint many theories and explanations come to consideration. If these are the result for every migration and breeding season the expected results should be a shift towards a larger male population through sexual selection. Sexual selection is strong when the factor of environmental selection is also introduced. The environmental selection may support a smaller chick size if those chicks were born in an area that allowed them to grow to a larger size, even though under normal conditions they would not be able to reach this optimal size for migration. When the environment gives advantages and disadvantages of this sort, the strength of selection is weakened and the environmental forces are given greater morphological weight. The sexual dimorphism could also produce a change in timing of migration leading to differences in mating success within the bird population.<ref>{{cite journal | last1 = Kissner | first1 = K. J. | last2 = Weatherhead | first2 = P. J. | last3 = Francis | first3 = C. M. | year = 2003 | title = Sexual size dimorphism and timing of spring migration in birds | url = | journal = Journal of Evolutionary Biology | volume = 16 | issue = 1| pages = 154–162 | doi=10.1046/j.1420-9101.2003.00479.x}}</ref> When the dimorphism produces that large of a variation between the sexes and between the members of the sexes multiple evolutionary effects can take place. This timing could even lead to a speciation phenomenon if the variation becomes strongly drastic and favorable towards two different outcomes.


Sexual dimorphism is maintained by the counteracting pressures of natural selection and sexual selection. For example, sexual dimorphism in coloration increases the vulnerability of bird species to predation by European sparrowhawks in Denmark.<ref name="Moller">{{cite journal |author=Anders Pape Møller & Jan Tøttrup Nielsen |year=2006 |title=Prey vulnerability in relation to sexual coloration of prey |url=http://www.springerlink.com/content/5838v731471n881w/fulltext.pdf |format=PDF | journal=[[Behavioral Ecology and Sociobiology]] |volume=60 |issue=2 |pages=227–233 |doi=10.1007/s00265-006-0160-x}}</ref> Presumably, increased sexual dimorphism means males are brighter and more conspicuous, leading to increased predation.<ref name="Moller"/> Moreover, the production of more exaggerated ornaments in males may come at the cost of suppressed immune function.<ref name="Lindsay"/> So long as the reproductive benefits of the trait due to sexual selection are greater than the costs imposed by natural selection, then the trait will propagate throughout the population. Reproductive benefits arise in the form of a larger number of offspring, while natural selection imposes costs in the form of reduced survival. This means that even if the trait causes males to die earlier, the trait is still beneficial so long as males with the trait produce more offspring than males lacking the trait. This balance keeps the dimorphism alive in these species and ensures that the next generation of successful males will also display these traits that are attractive to the females.
[[Grouse|Sexual dimorphism]] is maintained by the counteracting pressures of natural selection and sexual selection. For example, sexual dimorphism in coloration increases the vulnerability of bird species to predation by European sparrowhawks in Denmark.<ref name="Moller">{{cite journal |author=Anders Pape Møller & Jan Tøttrup Nielsen |year=2006 |title=Prey vulnerability in relation to sexual coloration of prey |url=http://www.springerlink.com/content/5838v731471n881w/fulltext.pdf |format=PDF | journal=[[Behavioral Ecology and Sociobiology]] |volume=60 |issue=2 |pages=227–233 |doi=10.1007/s00265-006-0160-x}}</ref> Presumably, increased sexual dimorphism means males are brighter and more conspicuous, leading to increased predation.<ref name="Moller"/> Moreover, the production of more exaggerated ornaments in males may come at the cost of suppressed immune function.<ref name="Lindsay"/> So long as the reproductive benefits of the trait due to sexual selection are greater than the costs imposed by natural selection, then the trait will propagate throughout the population. Reproductive benefits arise in the form of a larger number of offspring, while natural selection imposes costs in the form of reduced survival. This means that even if the trait causes males to die earlier, the trait is still beneficial so long as males with the trait produce more offspring than males lacking the trait. This balance keeps the dimorphism alive in these species and ensures that the next generation of successful males will also display these traits that are attractive to the females.


Such differences in form and reproductive roles often cause differences in behavior. As previously stated, males and females often have different roles in reproduction. The courtship and mating behavior of males and females are regulated largely by hormones throughout a bird's lifetime.<ref name="Adkins">{{cite journal |author=Elizabeth Adkins-Regan |year=2007 |title=Hormones and the development of sex differences in behavior |journal=[[Journal of Ornithology]] |volume=148 |issue=Supplement 1 |pages=S17–S26 |doi=10.1007/s10336-007-0188-3}}</ref> Activational hormones occur during puberty and adulthood and serve to 'activate' certain behaviors when appropriate, such as territoriality during breeding season.<ref name="Adkins"/> Organizational hormones occur only during a critical period early in development, either just before or just after hatching in most birds, and determine patterns of behavior for the rest of the bird's life.<ref name="Adkins"/> Such behavioral differences can cause disproportionate sensitivities to anthropogenic pressures.<ref name="Grubler">{{cite journal |author=Martin U. Grüebler, Heidi Schuler, Mathis Müller, Reto Spaar, Petra Horch & Beat Naef-Daenzer |year=2008 |title=Female biased mortality caused by anthropogenic nest loss contributes to population decline and adult sex ratio of a meadow bird |journal=[[Biological Conservation (journal)|Biological Conservation]] | volume = 141 | issue = 12| pages = 3040–3049 |doi=10.1016/j.biocon.2008.09.008}}</ref> Females of the whinchat in Switzerland breed in intensely managed grasslands.<ref name="Grubler"/> Earlier harvesting of the grasses during the breeding season lead to more female deaths.<ref name="Grubler"/> Populations of many birds are often male-skewed and when sexual differences in behavior increase this ratio, populations decline at a more rapid rate.<ref name="Grubler"/> Also not all male dimorphic traits are due to hormones like testosterone, instead they are a naturally occurring part of development, for example plumage.<ref>Owens, I. P. F., Short,R.V.,. (1995). Hormonal basis of sexual dimorphism in birds: Implications for new theories of sexual selection. Trends in Ecology & Evolution., 10(REF), 44.</ref>
Such differences in form and reproductive roles often cause differences in behavior. As previously stated, males and females often have different roles in reproduction. The courtship and mating behavior of males and females are regulated largely by hormones throughout a bird's lifetime.<ref name="Adkins">{{cite journal |author=Elizabeth Adkins-Regan |year=2007 |title=Hormones and the development of sex differences in behavior |journal=[[Journal of Ornithology]] |volume=148 |issue=Supplement 1 |pages=S17–S26 |doi=10.1007/s10336-007-0188-3}}</ref> Activational hormones occur during puberty and adulthood and serve to 'activate' certain behaviors when appropriate, such as territoriality during breeding season.<ref name="Adkins"/> Organizational hormones occur only during a critical period early in development, either just before or just after hatching in most birds, and determine patterns of behavior for the rest of the bird's life.<ref name="Adkins"/> Such behavioral differences can cause disproportionate sensitivities to anthropogenic pressures.<ref name="Grubler">{{cite journal |author=Martin U. Grüebler, Heidi Schuler, Mathis Müller, Reto Spaar, Petra Horch & Beat Naef-Daenzer |year=2008 |title=Female biased mortality caused by anthropogenic nest loss contributes to population decline and adult sex ratio of a meadow bird |journal=[[Biological Conservation (journal)|Biological Conservation]] | volume = 141 | issue = 12| pages = 3040–3049 |doi=10.1016/j.biocon.2008.09.008}}</ref> Females of the whinchat in Switzerland breed in intensely managed grasslands.<ref name="Grubler"/> Earlier harvesting of the grasses during the breeding season lead to more female deaths.<ref name="Grubler"/> Populations of many birds are often male-skewed and when sexual differences in behavior increase this ratio, populations decline at a more rapid rate.<ref name="Grubler"/> Also not all male dimorphic traits are due to hormones like testosterone, instead they are a naturally occurring part of development, for example plumage.<ref>Owens, I. P. F., Short,R.V.,. (1995). Hormonal basis of sexual dimorphism in birds: Implications for new theories of sexual selection. Trends in Ecology & Evolution., 10(REF), 44.</ref>

Revision as of 05:25, 10 December 2015

For sex differences in humans, see Sex differences in humans.
Female (left) and male (right) common pheasant, illustrating the dramatic difference in both color and size between sexes

Sexual dimorphism is the condition where the two sexes of the same species exhibit different characteristics beyond the differences in their sexual organs themselves. The condition occurs in many animals, insects, birds and some plants. Differences may include secondary sex characteristics, size, color, markings, and may also include behavioral differences. These differences may be subtle or exaggerated, and may increase over time as a result of sexual selection.

Overview

The peacock, on the right, is courting the peahen, on the left.
Male (bottom) and female mallards. The male mallard has an unmistakable green head.
Orgyia antiqua male and female.

Ornamentation and coloration

Common and easily identified types of dimorphism are ornamentation and coloration. A difference in coloration of sexes within a given species is called sexual dichromatism, which is commonly seen in many species of birds and reptiles.[1] Sexual selection leads to the exaggerated dimorphic traits that are used predominantly in competition over mates. The increased fitness resulting from ornamentation offsets its cost to produce or maintain suggesting complex evolutionary implications, but the costs and evolutionary implications vary from species to species.[2]

Exaggerated dimorphic traits are used predominantly in the competition over mates, implying sexual selection.[3] Ornaments may be costly to produce or maintain, which has complex evolutionary implications[4] but the costs and implications differ depending on the nature of the ornamentation (such as the colour mechanism involved).

The peafowl constitute conspicuous illustrations of the principle. The ornate plumage of peacocks, as used in the courting display, attracts peahens. At first sight one might mistake peacocks and peahens for completely different species because of the vibrant colours and the sheer size of the male's plumage; the peahen being of a subdued brown coloration.[5] The plumage of the peacock increases its vulnerability to predators because it is a hindrance in flight, and it renders the bird conspicuous in general.[5] Similar examples are manifold, such as in birds of paradise and argus pheasants.

Another example of sexual dichromatism is that of the nestling blue tits. Males are chromatically more yellow than females. It is believed that this is obtained by the ingestion of green lepidopteran larvae, which contain large amounts of the carotenoids lutein and zeaxanthin.[6] This diet also affects the sexually dimorphic colours in the human-invisible UV spectrum.[7][8] Hence, the male birds, although appearing yellow to humans actually have a violet-tinted plumage that is seen by females. This plumage is thought to be an indicator of male parental abilities.[9] Perhaps this is a good indicator for females because it shows that they are good at obtaining a food supply from which the carotenoid is obtained. There is a positive correlation between the chromas of the tail and breast feathers and body condition.[10] Carotenoids play an important role in immune function for many animals, so carotenoid dependent signals might indicate health.[11]

Frogs constitute another conspicuous illustration of the principle. There are two types of dichromatism for frog species: ontogenetic and dynamic. Ontogenetic frogs are more common and have permanent color changes in males or females. Litoria lesueuri is an example of a dynamic frog that has temporarily color changes in males during breeding season.[12] Hyperolius ocellatus is an ontogenetic frog with dramatic differences in both color and pattern between the sexes. At sexual maturity, the males display a bright green with white dorsolateral lines.[13] In contrast, the females are rusty red to silver with small spots. The bright coloration in the male population serves to attract females and as an aposematic sign to potential predators.

Females often show a preference for exaggerated male secondary sexual characteristics in mate selection.[14] The sexy son hypothesis explains that females prefer more elaborate males and discriminate against males that are dull in color, independent of the species’ vision.[15]

Similar sexual dimorphism and mating choice are also observed in many fish species. For example, in guppies males have colorful spots and ornamentations while females are generally grey in color. Female guppies prefer brightly colored males to duller males.

Psychological and behavioral differentiation

Sex steroid-induced differentiation of adult reproductive and other behavior has been demonstrated experimentally in many animals. In some mammals, adult sex-dimorphic reproductive behavior (e.g., mounting or receptive lordosis) can be shifted to that of the other sex by supplementation or deprivation of androgens in fetal life or early infancy, even if adult levels are normal.

Physiological differentiation

In redlip blennies, only the male fish develops an organ at the anal-urogenital region that produces antimicrobial substances. During parental care, males rub their anal-urogenital regions over their nests' internal surfaces, thereby protecting their eggs from microbial infections, one of the most common causes for mortality in young fish.[16]

Plants

Most plants are hermaphroditic but approximately 6% have separate males and females (dioecy).[17] Males and females in insect-pollinated species generally look similar to one another because plants provide rewards (e.g. nectar) that encourage pollinators to visit another similar flower, completing pollination. Catasetum orchids are one interesting exception to this rule. Male Catasetum orchids violently attach pollinia to euglossine bee pollinators. The bees will then avoid other male flowers but may visit the female, which looks different from the males.[18]

Various other dioecious exceptions, such as Loxostylis alata have visibly different genders, with the effect of eliciting the most efficient behaviour from pollinators, who then use the most efficient strategy in visiting each gender of flower instead of searching say, for pollen in a nectar-bearing female flower.

Some plants, such as some species of Geranium have what amounts to serial sexual dimorphism. The flowers of such species might for example present their anthers on opening, then shed the exhausted anthers after a day or two and perhaps change their colours as well while the pistil matures; specialist pollinators are very much inclined to concentrate on the exact appearance of the flowers they serve, which saves their time and effort and serves the interests of the plant accordingly. Some such plants go even further and change their appearance again once they have been fertilised, thereby discouraging further visits from pollinators. This is advantageous to both parties because it avoids damage to the developing fruit and avoids wasting the pollinator's effort on unrewarding visits. In effect the strategy ensures that the pollinators can expect a reward every time they visit an appropriately advertising flower.

Females of the aquatic plant Vallisneria americana have floating flowers attached by a long flower stalk that are fertilized if they contact one of the thousands of free floating flowers released by a male.[19] Sexual dimorphism is most often associated with wind-pollination in plants due to selection for efficient pollen dispersal in males vs pollen capture in females, e.g. Leucadendron rubrum.[20]

Sexual dimorphism in plants can also be dependent on reproductive development. This can be seen in Cannabis sativa, a type of hemp, which have higher photosynthesis rates in males while growing but higher rates in females once the plants become sexually mature.[21]

It also should be borne in mind that every sexually reproducing extant species of vascular plant actually has an alternation of generations; the plants we see about us generally are diploid sporophytes, but their offspring really are not the seeds that people commonly recognise as the new generation. The seed actually is the offspring of the haploid generation of microgametophytes (pollen) and megagametophytes (the embryo sacs in the ovules). Each pollen grain accordingly may be seen as a male plant in its own right; it produces a sperm cell and is dramatically different from the female plant, the megagametophyte that produces the female gamete.

Insects

Sexual dimorphism: Anthocharis cardamines male is brightly coloured.
Sexual dimorphism: Anthocharis cardamines female

Insects display a wide variety of sexual dimorphism between taxa including size, ornamentation and coloration.[22] The female-biased sexual size dimorphism observed in many taxa evolved despite intense male-male competition for mates.[23] In Osmia rufa, for example, the female is larger/broader than males, with males being 8–10 mm in size and females being 10–12 mm in size.[24] The reason for the sexual dimorphism is due to provision size mass, in which females consume more pollen than males.[25] In some species, there evidence of male dimorphism, but it appears to be for the purpose of distinctions of roles. This is seen in the bee species Perdita portalis in which there is a small headed morph, capable of flight, and large headed morph, incapable of flight, for males.[26] Anthidium manicatum also displays male-biased sexual dimorphism. The selection for larger size in males rather than females in this species may have resulted due to their aggressive territorial behavior and subsequent differential mating success.[27] Another example is the L. hemichalceum, which is a species of sweat bee that shows drastic physical dimorphisms between male offpsring.[28] Not all dimorphism has to have a drastic difference between the sexes. Andrena agilissima is a mining bee where the females only have a slightly larger head than the males.[29]

Weaponry leads to increased fitness by increasing success in male-male competition in many insect species.[30] The beetle horns in Onthophagus taurus are enlarged growths of the head or thorax expressed only in the males. Copris ochus also has distinct sexual and male dimorphism in head horns.[31] These structures are impressive because of the exaggerated sizes.[32] There is a direct correlation between male horn lengths and body size and higher access to mates and fitness.[32] In other beetle species, both males and females may have ornamentation such as horns.[31] Generally, insect sexual size dimorphism (SSD) within species increases with body size.[33]

Sexual dimorphism within insects is also displayed by dichromatism. In butterfly genera Bicyclus and Junonia, dimorphic wing patterns evolved due to sex-limited expression, which mediates the intralocus sexual conflict and leads to increased fitness in males.[34] The sexual dichromatic nature of Bicyclus anyana is reflected by female selection on the basis of dorsal UV-reflective eyespot pupils (Robertson & Monteiro, 2005).[35] Naturally selected deviation in protective female coloration is displayed in mimetic butterflies.[36]

Spiders and sexual cannibalism

Female (left) and Male (right) Argiope appensa, displaying typical sexual differences in spiders, with dramatically smaller males

Size dimorphism shows a correlation with sexual cannibalism, which is prominent in spiders (it is also found in insects such as praying mantises). In the size dimorphic wolf spider, food-limited females cannibalize more frequently.[37] Therefore, there is a high risk of low fitness for males due to pre-copulatory cannibalism, which led to male selection of larger females for two reasons: higher fecundity and lower rates of cannibalism.[37] In addition, female fecundity is positively correlated with female body size and large female body size is selected for, which is seen in the family Araneidae. All Argiope species, including Argiope bruennichi, use this method. Some males evolved ornamentation including binding the female with silk, having proportionally longer legs, modifying the females web, mating while the female is feeding, or providing a nuptial gift in response to sexual cannibalism.[37] Male body size is not under selection due to cannibalism in all spider species such as Nephila pilipes, but is more prominently selected for in less dimorphic species of spiders, which often selects for larger male size.[38]

Fish

There are also cases where males are substantially larger than females. An example is Lamprologus callipterus, a type of cichlid fish. In this fish, the males are characterized as being up to 60 times larger than the females. The male's increased size is believed to be advantageous because males collect and defend empty snail shells in each of which a female breeds.[39] Males must be larger and more powerful in order to collect the largest shells. The female's body size must remain small because in order for her to breed, she must lay her eggs inside the empty shells. If she grows too large, she will not fit in the shells and will be unable to breed. Another example is the dragonet, in which males are considerably larger than females and possess longer fins.

The female's small body size is also likely beneficial to her chances of finding an unoccupied shell. Larger shells, although preferred by females, are often limited in availability.[40] Hence, the female is limited to the growth of the size of the shell and may actually change her growth rate according to shell size availability.[41] In other words, the male's ability to collect large shells depends on his size. The larger the male, the larger the shells he is able to collect. This then allows for females to be larger in his brooding nest which makes the difference between the sizes of the sexes less substantial. Male-male competition in this fish species also selects for large size in males. There is aggressive competition by males over territory and access to larger shells. Large males win fights and steal shells from competitors. Sexual dimorphism also occurs in hermaphroditic fish. These species are known as sequential hermaphrodites. In fish, reproductive histories often include the sex-change from female to male where there is a strong connection between growth, the sex of an individual, and the mating system it operates within.[42] In protogynous mating systems where males dominate mating with many females, size plays a significant role in male reproductive success.[43] Males have a propensity to be larger than females of a comparable age but it is unclear whether the size increase is due to a growth spurt at the time of the sexual transition or due to the history of faster growth in sex changing individuals.[44] Larger males are able to stifle the growth of females and control environmental resources.

Social organization plays a large role in the changing of sex by the fish. It is often seen that a fish will change its sex when there is a lack of dominant male within the social hierarchy. The females that change sex are often those who attain and preserve an initial size advantage early in life. In either case, females which change sex to males are larger and often prove to be a good example of dimorphism.

In other cases with fish, males will go through noticeable changes in body size, and females will go through morphological changes that can only be seen inside of the body. For example, in sockeye salmon, males develop larger body size at maturity, including an increase in body depth, hump height, and snout length. Females experience minor changes in snout length, but the most noticeable difference is the huge increase in gonad size, which accounts for about 25% of body mass.[45]

Sexual selection was observed for female ornamentation in Gobiusculus flavescens, known as two-spotted gobies.[46] Traditional hypotheses suggest that male-male competition drives selection. However, selection for ornamentation within this species suggests that showy female traits can be selected through either female-female competition or male mate choice.[46] Since carotenoid-based ornamentation suggests mate quality, female two-spotted guppies that develop colorful orange bellies during the breeding season are considered favorable to males.[47] The males invest heavily in offspring during the incubation, which leads to the sexual preference in colorful females due to higher egg quality.[47]

Amphibians and reptiles

In amphibians and reptiles, the degree of sexual dimorphism varies widely among taxonomic groups. The sexual dimorphism in amphibians and reptiles may be reflected in any of the following: anatomy; relative length of tail; relative size of head; overall size as in many species of vipers and lizards; coloration as in many amphibians, snakes, and lizards, as well as in some chelonians; an ornament as in many newts and lizards; the presence of specific sex-related behaviour is common to many lizards; and vocal qualities which are frequently observed in frogs.

Anolis lizards show prominent size dimorphism with males typically being significantly larger than females. For instance, the average male Anolis sagrei was 53.4 mm vs. 40 mm in females.[48] Different sizes of the heads in anoles have been explained by differences in the estrogen pathway.[49] The sexual dimorphism in lizards is generally attributed to the effects of sexual selection, but other mechanisms including ecological divergence and fecundity selection provide alternative explanations.[50] The development of color dimorphism in lizards is induced by hormonal changes at the onset of sexual maturity, as seen in Psamodromus algirus, Sceloporus gadoviae, and S. undulates erythrocheilus.[50]

Birds

Mandarin ducks, male (left) and female (right)

Sexual dimorphism in birds can be manifested in size or plumage differences between the sexes. Sexual size dimorphism varies among taxa with males typically being larger, though this is not always the case i.e. birds of prey, hummingbirds, and some species of flightless birds.[51][52] Plumage dimorphism, in the form of ornamentation or coloration, also varies, though males are typically the more ornamented or brightly colored sex.[53] Such differences have been attributed to the unequal reproductive contributions of the sexes.[54] This difference produces a stronger female choice since they have more risk in producing offspring. In some species, the male's contribution to reproduction ends at copulation, while in other species the male becomes the main caregiver. Plumage polymorphisms have evolved to reflect these differences and other measures of reproductive fitness, such as body condition[55] or survival.[56] The male phenotype sends signals to females who then choose the 'fittest' available male.

Sexual dimorphism is a product of both genetics and environmental factors. An example of sexual polymorphism determined by environmental conditions exists in the red-backed fairywren. Red-backed fairywren males can be classified into three categories during breeding season: black breeders, brown breeders, and brown auxiliaries.[55] These differences arise in response to the bird's body condition: if they are healthy they will produce more androgens thus becoming black breeders, while less healthy birds produce less androgens and become brown auxiliaries.[55] The reproductive success of the male is thus determined by his success during each year's non-breeding season, causing reproductive success to vary with each year's environmental conditions.

Migratory patterns and behaviors also influence sexual dimorphisms. This aspect also stems back to the size dimorphism in species. It has been shown that the larger males are better at coping with the difficulties of migration and thusly are more successful in reproducing when reaching the breeding destination.[57] When viewing this in an evolutionary standpoint many theories and explanations come to consideration. If these are the result for every migration and breeding season the expected results should be a shift towards a larger male population through sexual selection. Sexual selection is strong when the factor of environmental selection is also introduced. The environmental selection may support a smaller chick size if those chicks were born in an area that allowed them to grow to a larger size, even though under normal conditions they would not be able to reach this optimal size for migration. When the environment gives advantages and disadvantages of this sort, the strength of selection is weakened and the environmental forces are given greater morphological weight. The sexual dimorphism could also produce a change in timing of migration leading to differences in mating success within the bird population.[58] When the dimorphism produces that large of a variation between the sexes and between the members of the sexes multiple evolutionary effects can take place. This timing could even lead to a speciation phenomenon if the variation becomes strongly drastic and favorable towards two different outcomes.

Sexual dimorphism is maintained by the counteracting pressures of natural selection and sexual selection. For example, sexual dimorphism in coloration increases the vulnerability of bird species to predation by European sparrowhawks in Denmark.[59] Presumably, increased sexual dimorphism means males are brighter and more conspicuous, leading to increased predation.[59] Moreover, the production of more exaggerated ornaments in males may come at the cost of suppressed immune function.[55] So long as the reproductive benefits of the trait due to sexual selection are greater than the costs imposed by natural selection, then the trait will propagate throughout the population. Reproductive benefits arise in the form of a larger number of offspring, while natural selection imposes costs in the form of reduced survival. This means that even if the trait causes males to die earlier, the trait is still beneficial so long as males with the trait produce more offspring than males lacking the trait. This balance keeps the dimorphism alive in these species and ensures that the next generation of successful males will also display these traits that are attractive to the females.

Such differences in form and reproductive roles often cause differences in behavior. As previously stated, males and females often have different roles in reproduction. The courtship and mating behavior of males and females are regulated largely by hormones throughout a bird's lifetime.[60] Activational hormones occur during puberty and adulthood and serve to 'activate' certain behaviors when appropriate, such as territoriality during breeding season.[60] Organizational hormones occur only during a critical period early in development, either just before or just after hatching in most birds, and determine patterns of behavior for the rest of the bird's life.[60] Such behavioral differences can cause disproportionate sensitivities to anthropogenic pressures.[61] Females of the whinchat in Switzerland breed in intensely managed grasslands.[61] Earlier harvesting of the grasses during the breeding season lead to more female deaths.[61] Populations of many birds are often male-skewed and when sexual differences in behavior increase this ratio, populations decline at a more rapid rate.[61] Also not all male dimorphic traits are due to hormones like testosterone, instead they are a naturally occurring part of development, for example plumage.[62]

Sexual dimorphism may also influence differences in parental investment during times of food scarcity. For example, in the blue-footed booby, the female chicks grow faster than the males, resulting in booby parents producing the smaller sex, the males, during times of food shortage. This then results in the maximization of parental lifetime reproductive success.[63]

Sexual dimorphism may also only appear during mating season, some species of birds only show dimorphic traits in seasonal variation. The males of these species will molt into a less bright or less exaggerated color during the off breading season.[64] This occurs because the species is more focused on survival than reproduction, causing a shift into a less ornate state.

Consequently, sexual dimorphism has important ramifications for conservation. However, sexual dimorphism is not only found in birds and is thus important to the conservation of many animals. Such differences in form and behavior can lead to sexual segregation, defined as sex differences in space and resource use.[65] Most sexual segregation research has been done on ungulates,[65] but such research extends to bats,[66] kangaroos,[67] and birds.[68] Sex-specific conservation plans have even been suggested for species with pronounced sexual segregation.[66]

The term sesquimorphism (the Latin numeral prefix sesqui- means one-and-one-half, so halfway between mono- (one) and di- (two)) has been proposed for bird species in which "both sexes have basically the same plumage pattern, though the female is clearly distinguishable by reason of her paler or washed-out colour".[69]: 14  Examples include Cape sparrow (Passer melanurus),[69]: 67  rufous sparrow (subspecies P. motinensis motinensis),[69]: 80  and saxaul sparrow (P. ammodendri).[69]: 245 

Mammals

Just like in birds, the brains of many mammals, including humans, are significantly different for males and females of the species.[70] Both genes and hormones affect the formation of many animal brains before "birth" (or hatching), and also behaviour of adult individuals. Hormones significantly affect human brain formation, and also brain development at puberty. A 2004 review in Nature Reviews Neuroscience observed that "because it is easier to manipulate hormone levels than the expression of sex chromosome genes, the effects of hormones have been studied much more extensively, and are much better understood, than the direct actions in the brain of sex chromosome genes." It concluded that while "the differentiating effects of gonadal secretions seem to be dominant," the existing body of research "support the idea that sex differences in neural expression of X and Y genes significantly contribute to sex differences in brain functions and disease."[71]

Pinnipeds

Marine mammals show some of the greatest sexual size differences of mammals. The mating system of pinnipeds varies from polygyny to serial monogamy. Pinnipeds are known for early differential growth and maternal investment since the only nutrients for newborn pups is the milk provided by the mother.[72] For example, the males are significantly larger than the females at birth in sea lion pups. The pattern of differential investment can be varied principally prenatally and post-natally.[73] Mirounga leonina, the southern elephant seal, is one of the most dimorphic mammals.[74]

Sexual dimorphism in elephant seals is associated with the ability of a male to defend territories, which correlates with polygynic behavior. The large sexual size dimorphism is due to sexual selection, but also because females reach reproductive age much earlier than males. In addition the males do not provide parental care for the young and allocate more energy to growth.[75] This is supported by the secondary growth spurt in males during adolescent years.[75]

Primates

Main article: Sexual dimorphism in non-human primates

Humans

Main articles: Sex differences in humans and Sex differences in human psychology
Pioneer plaque
Pioneer plaque
Male pelvis
Male pelvis
Female pelvis
Female pelvis

Top: Stylised illustration of humans on the Pioneer plaque, showing both male (left) and female (right).
Bottom: Comparison between male (left) and female (right) pelvises.

File:Male female facial sexual dimorphism masculine feminine face.png
This is a remake of a facial geometric sexual dimorphism diagram from Valenzano, D. R. et al. (2006). The blue bell curve on the left represents the male faces, and the pink bell curve on the right represents the female faces. The purple area in the center represents the overlap of the two bell curves where the feminine male faces cannot be distinguished from the masculine female faces. The bell curves show that the proportion of female faces that are more feminine than the most feminine male faces is much greater than the proportion of male faces that are more masculine than the most masculine female faces.[76]

In humans, biological sex is determined by five factors present at birth: the presence or absence of a Y chromosome, the type of gonads, the sex hormones, the internal reproductive anatomy (such as the uterus in females), and the external genitalia.[77] Generally, the five factors are either all male or all female. Unexpected combinations cause intersex conditions.

The average differences between males and females include all the features related to reproduction. Sexual dimorphism among humans includes differentiation among gonads, internal genitals, external genitals, breasts, muscle mass, height, the endocrine (hormonal) systems and their physiological and behavioral effects. According to Clark Spencer Larsen, modern day Homo sapiens show a range of sexual dimorphism, with average body mass difference between the sexes being roughly equal to 15% . Ever since Charles Darwin's The Descent of Man and Selection in Relation to Sex in 1871 was published, there's been controversy regarding the social, cultural, and political significance of human sexual dimorphism.

The average basal metabolic rate is about 6 percent higher in adolescent males than females and increases to about 10 percent higher after puberty. Females tend to convert more food into fat, while males convert more into muscle and expendable circulating energy reserves. Aggregated data of absolute strength indicates that females have, on average, 40-60% the upper body strength of males, and 70-75% the lower body strength.[78] The difference in strength relative to body mass is less pronounced in trained individuals. In Olympic weightlifting, male records vary from 5.5× body mass in the lowest weight category to 4.2× in the highest weight category, while female records vary from 4.4× to 3.8×, a weight adjusted difference of only 10-20%, and an absolute difference of about 30% (i.e. 472 kg vs 333 kg for unlimited weight classes)(see Olympic weightlifting records). A study, carried about by analyzing annual world rankings from 1980–1996, found that males' running times were, on average, 11% faster than females'.[79]

Females are taller, on average, than males in early adolescence, but males, on average, surpass them in height in later adolescence and adulthood. In the United States, adult males are, on average, 4% taller[80] and 8% heavier[81] than adult females.

Males typically have larger tracheae and branching bronchi, with about 30 percent greater lung volume per body mass. On average, males have larger hearts, 10 percent higher red blood cell count, higher hemoglobin, hence greater oxygen-carrying capacity. They also have higher circulating clotting factors (vitamin K, prothrombin and platelets). These differences lead to faster healing of wounds and higher peripheral pain tolerance.[82]

Females typically have more white blood cells (stored and circulating), more granulocytes and B and T lymphocytes. Additionally, they produce more antibodies at a faster rate than males. Hence they develop fewer infectious diseases and succumb for shorter periods.[82] Ethologists argue that females, interacting with other females and multiple offspring in social groups, have experienced such traits as a selective advantage.[83][84][85][86][87]

Considerable discussion in academic literature concerns potential evolutionary advantages associated with sexual competition (both intrasexual and intersexual) and short- and long-term sexual strategies.[88]

According to Daly and Wilson, "The sexes differ more in human beings than in monogamous mammals, but much less than in extremely polygamous mammals."[89] One proposed explanation is that human sexuality has developed more in common with its close relative the bonobo, who have similar sexual dimorphism and which are polygynandrous and use recreational sex to reinforce social bonds and reduce aggression.[90]

In the human brain, a difference between sexes was observed in the transcription of the PCDH11X/Y gene pair unique to Homo sapiens.[91] Sexual differentiation in the human brain from the default female state is triggered by testosterone from the fetal testis. Testosterone is converted to estrogen in the brain through the action of the enzyme aromatase. Testosterone acts on many brain areas, including the SDN-POA, to create the masculinized brain pattern.[92] Female brains may be shielded from the masculinizing effects of estrogen through the action of sex-hormone binding globulin and a-fetoprotein.[93] The relationship between sex differences in the brain and human behavior is a subject of controversy in psychology and society at large.[94][95] Many females tend to have a higher ratio of gray matter in the left hemisphere of the brain in comparison to males.[96][97] However, on average, males have larger brains than females, and when adjusted for total brain volume the gray matter differences between sexes is almost nonexistent. Thus, the percentage of gray matter appears to be more related to brain size than it is to sex.[98][99] Differences in brain physiology between sexes do not necessarily relate to differences in intellect. Haier et al. found in a 2004 study that "men and women apparently achieve similar IQ results with different brain regions, suggesting that there is no singular underlying neuroanatomical structure to general intelligence and that different types of brain designs may manifest equivalent intellectual performance".[100] (See the sex and intelligence article for more on this subject.)

Cells

Phenotypic differences between sexes are evident even in cultured cells from tissues.[101] For example, female muscle-derived stem cells have a better muscle regeneration efficiency than male ones.[102] There are reports of several metabolic differences between male and female cells[103] and they also respond to stress differently.[104]

Species with females larger than males

In some species, such as many insects, many spiders, many fish, many reptiles, owls, birds of prey and certain mammals such as the spotted hyena, and blue whale, the female is larger than the male. As an example, in some species, females are sedentary and sparsely distributed, and so males must search for them. Fritz Vollrath and Geoff Parker argue that this difference in behaviour leads to radically different selection pressures on the two sexes, evidently favouring smaller males.[105] Cases where the male is larger than the female have been studied as well,[105] and require alternative explanations.

One example of sexual size dimorphism is the bat Myotis nigricans, where females are substantially larger than males in terms of body weight, skull measurement, and forearm length.[106] The interaction between the sexes and the energy needed to produce viable offspring make it favorable for females to be larger in this species. Females bear the energetic cost of producing eggs, which is much greater than the cost of making sperm by the males. The fecundity advantage hypothesis states that a larger female is able to produce more offspring and give them more favorable conditions to ensure their survival; this is true for most ectotherms. A larger female can provide parental care for a longer time while the offspring matures. The gestation and lactation periods are fairly long in M. nigricans, the females suckling their offspring until they reach nearly adult size.[107] They would not be able to fly and catch prey if they did not compensate for the additional mass of the offspring during this time. Smaller male size may be an adaptation to increase maneuverability and agility, allowing males to compete better with females for food and other resources.

Female triplewart seadevil, an anglerfish, with male attached near vent (arrow)

Some species of anglerfish also display extreme sexual dimorphism. Females are more typical in appearance to other fish, whereas the males are tiny rudimentary creatures with stunted digestive systems. A male must find a female and fuse with her: he then lives parasitically, becoming little more than a sperm-producing body. A similar situation is found in the Zeus water bug Phoreticovelia disparata where the female has a glandular area on her back that can serve to feed a male, which clings to her (note that although males can survive away from females, they generally are not free-living).[108]

Some plant species also exhibit dimorphism in which the females are significantly larger than the males, such as in the moss Dicranum[109] and the liverwort Sphaerocarpos.[110] There is some evidence that, in these genera, the dimorphism may be tied to a sex chromosome,[110][111] or to chemical signalling from females.[112]

Another complicated example of sexual dimorphism is in the Vespula squamosa, or southern yellowjacket. In this wasp species, the female workers are the smallest, the male workers are slightly larger, and the female queens are significantly larger than her female worker and male counterparts.

Evolution

See also: Sexual selection and Mate choice

Sexual size dimorphism has been known to present itself in extinct species capable of sexual reproduction, such as the velociraptor. In the case of velociraptors, the sexual size dimorphism can be caused by two factors: Males competing for hunting ground to attract mates, and/or females competing for a nesting location for their offspring, while also competing for mates, since males were considered a scarce breeding resource.[113]

In 1871 Charles Darwin advanced the theory of sexual selection, which related sexual dimorphism with sexual selection.

It has been proposed that the earliest sexual dimorphism is the size differentiation of sperm and eggs (anisogamy), but the evolutionary significance of sexual dimorphism is more complex than that would suggest.[114] Anisogamy and the usually large number of small male gametes relative to the larger female gametes usually lies in the development of strong sperm competition,[115][116] because small sperm enable organisms to produce a large number of sperm, and make males (or male function of hermaphrodites[117]) more redundant. This intensifies male competition for mates, and promotes the evolution of other sexual dimorphim in many species, especially in vertebrates, including mammals. However, in some species, the females can be larger than males, irrespective of gametes, and in some species females (usually of species in which males invest a lot in rearing offspring and thus no longer considered as so redundant) compete for mates in ways more usually associated with males.

In many non-monogamous species, the benefit to a male's reproductive fitness of mating with multiple females is large, whereas the benefit to a female's reproductive fitness of mating with multiple males is small or nonexistent.[118] In these species, there is a selection pressure for whatever traits enable a male to have more matings. The male may therefore come to have different traits from the female.

Male (left), offspring, and female (right) Sumatran orangutans.

These traits could be ones that allow him to fight off other males for control of territory or a harem, such as large size or weapons;[119] or they could be traits that females, for whatever reason, prefer in mates.[120] Male-male competition poses no deep theoretical questions[121] but mate choice does.

Females may choose males that appear strong and healthy, thus likely to possess "good alleles" and give rise to healthy offspring.[122] However, in some species females seem to choose males with traits that do not improve offspring survival rates, and even traits that reduce it (potentially leading to traits like the peacock's tail).[121] Two hypotheses for explaining this fact are the sexy son hypothesis and the handicap principle.

The sexy son hypothesis states that females may initially choose a trait because it improves the survival of their young, but once this preference has become widespread, females must continue to choose the trait, even if it becomes harmful. Those that do not will have sons that are unattractive to most females (since the preference is widespread) and so receive few matings.[123]

The handicap principle states that a male who survives despite possessing some sort of handicap thus proves that the rest of his genes are "good alleles". If males with "bad alleles" could not survive the handicap, females may evolve to choose males with this sort of handicap; the trait is acting as a hard-to-fake signal of fitness.[124]

See also

Template:Wikipedia books

References

Notes

  1. ^ Armenta, J. K., Dunn, P. O., & Whittingham, L. A. (2008). Quantifying avian sexual dichromatism: a comparison of methods. The Journal of Experimental Biology, 211, 2423–2430.
  2. ^ Zahavi, A. (1975). Mate selection-a selection for a handicap. Journal of Theoretical Biology, 53, 205–214.
  3. ^ Andersson 1994
  4. ^ Amotz Zahavi (1975). "Mate selection – a selection for a handicap" (PDF). Journal of Theoretical Biology. 53 (1): 205–214. doi:10.1016/0022-5193(75)90111-3. PMID 1195756.
  5. ^ a b Zi, J., Yu, X., Li, Y., Hu, X., Xu, C., Wang, X., … Fu, R. (2003). Coloration strategies in peacock feathers. Proceedings of the National Academy of Sciences of the United States of America, 100(22), 12576–12578.
  6. ^ T. Slagsvold & J. T. Lifjeld (1985). "Variation in plumage colour of the Great tit Parus major in relation to habitat, season and food". Journal of Zoology. 206 (3): 321–328. doi:10.1111/j.1469-7998.1985.tb05661.x.
  7. ^ Stefan Andersson, Jonas Örnborg & Malte Andersson (1998). "Ultraviolet sexual dimorphism and assortative mating in blue tits". Proceedings of the Royal Society B: Biological Sciences. 265 (1395): 445–450. doi:10.1098/rspb.1998.0315. PMC 1688915.
  8. ^ Sarah Hunt, Andrew T. D. Bennett, Innes C. Cuthill & Richard Griffiths (1998). "Blue tits are ultraviolet tits". Proceedings of the Royal Society B: Biological Sciences. 265 (1395): 451–455. doi:10.1098/rspb.1998.0316. JSTOR 50814. PMC 1688906.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  9. ^ J. C. Senar, J. Figuerola & J. Pascual (2002). "Brighter yellow blue tits make better parents". Proceedings of the Royal Society B: Biological Sciences. 269 (1488): 257–261. doi:10.1098/rspb.2001.1882. PMC 1690890. PMID 11839194.
  10. ^ A. Johnsen, K. Delhey, S. Andersson & B. Kempenaers (2003). "Plumage colour in nestling blue tits: sexual dichromatism, condition dependence and genetic effects" (PDF). Proceedings of the Royal Society B. 270 (1521): 1263–1270. doi:10.1098/rspb.2003.2375. JSTOR 3558810. PMC 1691364. PMID 12816639.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  11. ^ George A. Lozano (1994). "Carotenoids, parasites, and sexual selection" (PDF). Oikos. 70 (2): 309–311. doi:10.2307/3545643. JSTOR 3545643.
  12. ^ Donnellan, S. C., & Mahony, M. J. (2004). Allozyme, chromosomal and morphological variability in the Litoria lesueuri species group (Anura : Hylidae), including a description of a new species. Australian Journal of Zoology
  13. ^ Bell, R. C., & Zamudio, K. R. (2012). Sexual dichromatism in frogs: natural selection, sexual selection and unexpected diversity. Proceedings of the Royal Society B: Biological Sciences.
  14. ^ Ryan, M. J., & Rand, S. (1993). Species Recognition and Sexual Selection as a Unitary Problem in Animal Communication. Evolution, 47, 647–657.
  15. ^ Rubolini, D., Spina, F., & Saino, N. (2004). Protandry and sexual dimorphism in trans-Saharan migratory birds. Behavioral Ecology, 15, 592–601.
  16. ^ Giacomello, Eva (2006). "A male sexually dimorphic trait provides antimicrobials to eggs in blenny fish". Biology Letters. 2: 330–333. doi:10.1098/rsbl.2006.0492.
  17. ^ Renner, Susanne; Ricklefs, Robert E (1995). "Dioecy and its correlates in the flowering plants". Americal Journal of Botany. 82 (5): 596–606. doi:10.2307/2445418. JSTOR 2445418.
  18. ^ Romero, Gustavo; Nelson, Craig E. (1986). "Sexual Dimorphism in Catasetum Orchids: Forcible Pollen Emplacement and Male Flower Competition". Science. 232 (4757): 1538–1540. Bibcode:1986Sci...232.1538R. doi:10.1126/science.232.4757.1538. JSTOR 1698050.
  19. ^ "Eel Grass (aka wild celery, tape grass)". University of Massachusetts.
  20. ^ Friedman, Jannice; Barrett, Spencer (2009). "Wind of change: new insights on the ecology and evolution of pollination and mating in wind-pollinated plants". Annals of Botany. 103 (9): 1515–1527. doi:10.1093/aob/mcp035.
  21. ^ Geber, Monica A. (1999). Gender and sexual dimorphism in flowering plants. Berlin: Springer. ISBN 3-540-64597-7. p. 206
  22. ^ Bonduriansky, R. (2007). The evolution of condition-dependent sexual dimorphism. The American Naturalist, 169, 9–19.
  23. ^ Barreto, F. S., & Avise, J. C. (2011). The genetic mating system of a sea spider with male-biased sexual size dimorphism: Evidence for paternity skew despite random mating success. Behavioral Ecology and Sociobiology, 65, 1595–1604.
  24. ^ Gruber, Bernd; Eckel, Katharina; Everaars, Jeroen; Dormann, Carsten F. (30 June 2011). "On managing the red mason bee (Osmia bicornis) in apple orchards". Apidologie. 42 (5): 564–576. doi:10.1007/s13592-011-0059-z. ISSN 0044-8435.
  25. ^ Rust, R.; Torchio, P.; Trostle, G. "Late embryogenesis and immature development of Osmia rufa cornigera (Rossi) (Hymenoptera : Megachilidae)". Apidologie. 20 (4): 359–367. doi:10.1051/apido:19890408.
  26. ^ Bryan Danforth (1991). "The morphology and behavior of dimorphic males in Perdita portalis (Hymenoptera : Andrenidae)". Behavioral Ecology and Sociobiology. 29 (4): pp 235-pp 247. {{cite journal}}: |pages= has extra text (help)
  27. ^ Jaycox, Elbert R. (October 1967). "Territorial Behavior Among Males of Anthidium Bamngense". Journal of the Kansas Entomological Society 40 (4): 565–570. Retrieved 8 October 2015.
  28. ^ Kukuk, Penelope F. (1 October 1996). "Male Dimorphism in Lasioglossum (Chilalictus) hemichalceum: The Role of Larval Nutrition". Journal of the Kansas Entomological Society. 69 (4): 147–157.
  29. ^ Paxton, R. J.; Giovanetti, M.; Andrietti, F.; Scamoni, E.; Scanni, B. (1 October 1999). "Mating in a communal bee, Andrena agilissima (Hymenoptera Andrenidae)". Ethology Ecology & Evolution. 11 (4): 371–382. doi:10.1080/08927014.1999.9522820. ISSN 0394-9370.
  30. ^ Wang, M.-Q., & Yang, D. (2005). Sexual dimorphism in insects. Chinese Bulletin of Entomology, 42, 721–725.
  31. ^ a b Sugiura, S., Yamaura, Y., & Makihara, H. (2007). Sexual and male horn dimorphism in Copris ochus (Coleoptera: Scarabaeidae). Zoological Science, 24, 1082–1085.
  32. ^ a b Emlen, D. J., Marangelo, J., Ball, B., & Cunningham, C. W. (2005). Diversity in the weapons of sexual selection: horn evolution in the beetle genus Onthophagus (Coleoptera: Scarabaeidae). Evolution; International Journal of Organic Evolution, 59, 1060–1084.
  33. ^ Teder, T., & Tammaru, T. (2005). Sexual size dimorphism within species increases with body size in insects. Oikos.
  34. ^ Oliver, J. C., & Monteiro, A. (2011). On the origins of sexual dimorphism in butterflies. Proceedings. Biological Sciences / The Royal Society, 278, 1981–1988.
  35. ^ Robertson, K. A., & Monteiro, A. (2005). Female Bicyclus anynana butterflies choose males on the basis of their dorsal UV-reflective eyespot pupils. Proceedings. Biological Sciences / The Royal Society, 272, 1541–1546.
  36. ^ Kunte, K. (2008). Mimetic butterflies support Wallace’s model of sexual dimorphism. Proceedings. Biological Sciences / The Royal Society, 275, 1617–1624.
  37. ^ a b c Wilder, S. M., & Rypstra, A. L. (2008). Sexual size dimorphism mediates the occurrence of state-dependent sexual cannibalism in a wolf spider. Animal Behaviour, 76, 447–454.
  38. ^ Foellmer, M. W., & Fairbairn, D. J. (2004). Males under attack: Sexual cannibalism and its consequences for male morphology and behaviour in an orb-weaving spider. Evolutionary Ecology Research, 6, 163–181.
  39. ^ Kazutaka Ota, Masanori Kohda & Tetsu Sato (2010). "Unusual allometry for sexual size dimorphism in a cichlid where males are extremely larger than females". Journal of Biosciences. 35 (2): 257–265. doi:10.1007/s12038-010-0030-6.
  40. ^ Tetsu Sato (1994). "Active accumulation of spawning substrate: a determinant of extreme polygyny in a shell-brooding cichlid fish". Animal Behaviour. 48 (3): 669–678. doi:10.1006/anbe.1994.1286.
  41. ^ Dolores Schütz & Michael Taborsky (2005). "Mate choice and sexual conflict in the size dimorphic water spider Argyroneta aquatica (Araneae: Argyronetidae)" (PDF). Journal of Arachnology. 33 (3): 767–775. doi:10.1636/S03-56.1.
  42. ^ Mark I. McCormick, Christopher A. Ryen, Philip L. Munday, Stefan P. W. Walker (2010). Briffa, Mark (ed.). "Differing mechanisms underlie sexual size-dimorphism in two populations of a sex-changing fish". PLoS One. 5 (5): e10616. Bibcode:2010PLoSO...510616M. doi:10.1371/journal.pone.0010616. PMC 2868897. PMID 20485547.{{cite journal}}: CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)
  43. ^ Robert R. Warner (1998). "Sex change and the size-advantage model". Trends in Ecology and Evolution. 3 (6): 133–136. doi:10.1016/0169-5347(88)90176-0. PMID 21227182.
  44. ^ S. Adams & A. J. Williams (2001). "A preliminary test of the transitional growth spurt hypothesis using the protogynous coral trout Plectropomus maculatus". Journal of Fish Biology. 59 (1): 183–185. doi:10.1111/j.1095-8649.2001.tb02350.x.
  45. ^ Hendry, Andrew P.; Ole K. Berg (1999). "Secondary sexual characters, energy use, senescence, and the cost of reproduction in sockeye salmon". Canadian Journal of Zoology. 77: 1663–1675. doi:10.1139/cjz-77-11-1663.
  46. ^ a b Amundsen, T., & Forsgren, E. (2001). Male mate choice selects for female coloration in a fish. Proceedings of the National Academy of Sciences of the United States of America, 98, 13155–13160.
  47. ^ a b Svensson, P. A., Pélabon, C., Blount, J. D., Surai, P. F., & Amundsen, T. (2006). Does female nuptial coloration reflect egg carotenoids and clutch quality in the Two-Spotted Goby (Gobiusculus flavescens, Gobiidae)? Functional Ecology, 20, 689–698.
  48. ^ Butler M. A.; et al. (2000). "THE RELATIONSHIP BETWEEN SEXUAL SIZE DIMORPHISM AND HABITAT USE IN GREATER ANTILLEAN ANOLIS LIZARDS" (PDF). Evolution. 54 (1): 259–272. doi:10.1111/j.0014-3820.2000.tb00026.x.
  49. ^ Sanger TJ, Seav SM, Tokita M, Langerhans RB, Ross LM, Losos JB, Abzhanov A (2014). "The oestrogen pathway underlies the evolution of exaggerated male cranial shapes inAnolis lizards". Proceedings of the Royal Society B. 281 (1784): 20140329. doi:10.1098/rspb.2014.0329.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  50. ^ a b Pinto, A., Wiederhecker, H., & Colli, G. (2005). Sexual dimorphism in the Neotropical lizard, Tropidurus torquatus (Squamata, Tropiduridae). Amphibia-Reptilia.
  51. ^ Andersson 1994, p. 269
  52. ^ Berns, Chelsea M.; Adams, Dean C. (11 November 2012). "Becoming Different But Staying Alike: Patterns of Sexual Size and Shape Dimorphism in Bills of Hummingbirds". Evolutionary Biology. 40 (2): 246–260. doi:10.1007/s11692-012-9206-3. ISSN 0071-3260.
  53. ^ K. J. McGraw, G. E. Hill, R. Stradi & R. S. Parker (2002). "The effect of dietary carotenoid access on sexual dichromatism and plumage pigment composition in the American goldfinch" (PDF). Comparative Biochemistry and Physiology Part B-Biochemistry and Molecular Biology. 131 (2): 261–269. doi:10.1016/S1096-4959(01)00500-0. Archived from the original (PDF) on 28 August 2005.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  54. ^ I. P. F. Owens & I. R. Hartley (1998). "Sexual dimorphism in birds: why are there so many different forms of dimorphism?". Proceedings of the Royal Society B. 265 (1394): 397–407. doi:10.1098/rspb.1998.0308. JSTOR 50849. PMC 1688905.
  55. ^ a b c d Willow R. Lindsay, Michael S. Webster, Claire W. Varian & Hubert Schwabl (2009). "Plumage colour acquisition and behaviour are associated with androgens in a phenotypically plastic bird". Animal Behaviour. 77 (6): 1525–1532. doi:10.1016/j.anbehav.2009.02.027.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  56. ^ Marion Petrie (1994). "Improved growth and survival of offspring of peacocks with more elaborate trains" (PDF). Nature. 371 (6498): 598–599. Bibcode:1994Natur.371..598P. doi:10.1038/371598a0.
  57. ^ Rubolini, Diego; Spina, Fernando; Saino, Nicola (2004). "Protandry and sexual dimorphism in trans-saharan migratory birds". Behavioral Ecology. 15 (4): 592–601. doi:10.1093/beheco/arh048.
  58. ^ Kissner, K. J.; Weatherhead, P. J.; Francis, C. M. (2003). "Sexual size dimorphism and timing of spring migration in birds". Journal of Evolutionary Biology. 16 (1): 154–162. doi:10.1046/j.1420-9101.2003.00479.x.
  59. ^ a b Anders Pape Møller & Jan Tøttrup Nielsen (2006). "Prey vulnerability in relation to sexual coloration of prey" (PDF). Behavioral Ecology and Sociobiology. 60 (2): 227–233. doi:10.1007/s00265-006-0160-x.
  60. ^ a b c Elizabeth Adkins-Regan (2007). "Hormones and the development of sex differences in behavior". Journal of Ornithology. 148 (Supplement 1): S17 – S26. doi:10.1007/s10336-007-0188-3.
  61. ^ a b c d Martin U. Grüebler, Heidi Schuler, Mathis Müller, Reto Spaar, Petra Horch & Beat Naef-Daenzer (2008). "Female biased mortality caused by anthropogenic nest loss contributes to population decline and adult sex ratio of a meadow bird". Biological Conservation. 141 (12): 3040–3049. doi:10.1016/j.biocon.2008.09.008.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  62. ^ Owens, I. P. F., Short,R.V.,. (1995). Hormonal basis of sexual dimorphism in birds: Implications for new theories of sexual selection. Trends in Ecology & Evolution., 10(REF), 44.
  63. ^ Velando, Alberto (2002). "Experimental Manipulation of Maternal Effort Produces Differential Effects in Sons and Daughters: Implications for Adaptive Sex Ratios in the Blue-footed Booby". Behavioral Ecology. 13 (4): 443. doi:10.1093/beheco/13.4.443.
  64. ^ The Genetic Basis of Sexual Dimorphism in Birds Jerry A. Coyne, Emily H. Kay and Stephen Pruett-Jones Evolution, Vol. 62, No. 1 (Jan., 2008), pp. 214-219
  65. ^ a b Martin B. Main (2008). "Reconciling competing ecological explanations for sexual segregation in ungulates". Ecology. 89 (3): 693–704. doi:10.1890/07-0645.1. PMID 18459333.
  66. ^ a b Kamran Safi, Barbara König & Gerald Kerth (2007). "Sex differences in population genetics, home range size and habitat use of the parti-colored bat (Vespertilio murinus, Linnaeus 1758) in Switzerland and their consequences for conservation". Biological Conservation. 137 (1): 28–36. doi:10.1016/j.biocon.2007.01.011.
  67. ^ G. Coulson, A. M. MacFarlane, S. E. Parsons & J. Cutter (2006). "Evolution of sexual segregation in mammalian herbivores: kangaroos as marsupial models" (PDF). Australian Journal of Zoology. 54 (3): 217–224. doi:10.1071/ZO05062.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  68. ^ Jacob González-Solís, John P. Croxall & Andy G. Wood (2000). "Sexual dimorphism and sexual segregation in foraging strategies of northern giant petrels, Macronectes halli, during incubation". Oikos. 90 (2): 390–398. doi:10.1034/j.1600-0706.2000.900220.x.
  69. ^ a b c d Summers-Smith, J. Denis (1988). The Sparrows. Calton, Staffordshire, UK: T. & A. D. Poyser. ISBN 0-85661-048-8.
  70. ^ Robert W. Goy and Bruce S. McEwen (1980). Sexual Differentiation of the Brain: Based on a Work Session of the Neurosciences Research Program. Boston: MIT Press. ISBN 978-0-262-57207-1.
  71. ^ Arthur P. Arnold (2004). "Sex chromosomes and brain gender". Nature Reviews Neuroscience. 5 (9): 701–708. doi:10.1038/nrn1494. PMID 15322528.
  72. ^ Cappozzo, H. L., Campagna, C., & Monserrat, J. (1991). Sexual Dimorphism in Newborn Southern Sea Lions. Marine Mammal Science, 7, 385–394.
  73. ^ Ono, K. A., & Boness, D. J. (1996). Sexual dimorphism in sea lion pups: differential maternal investment, or sex-specific differences in energy allocation? Behavioral Ecology and Sociobiology.
  74. ^ Tarnawski, B. A., Cassini, G. H., & Flores, D. A. (2014). Skull allometry and sexual dimorphism in the ontogeny of the southern elephant seal (Mirounga leonina). Canadian Journal of Zoology, 31, 19–31.
  75. ^ a b Lindenfors, P., Tullberg, B. S., & Biuw, M. (2002). Phylogenetic analyses of sexual selection and sexual size dimorphism in pinnipeds. Behavioral Ecology and Sociobiology, 52, 188–193.
  76. ^ Valenzano, D. R. et al. (2006). Shape analysis of female facial attractiveness. Vision Research. 46, 1282.
  77. ^ Knox, David; Schacht, Caroline. Choices in Relationships: An Introduction to Marriage and the Family. 11 ed. Cengage Learning; 2011-10-10 [cited 17 June 2013]. ISBN 9781111833220. p. 64–66.
  78. ^ "Strength training for female athletes: A position paper: Part 1". NSCA. 11 (4). 1989.
  79. ^ Phillip B. Sparling, Elizabeth M. O'Donnell & Teresa K. Snow (1998). "The gender difference in distance running performance has plateaued: an analysis of world rankings from 1980 to 1996". Medicine & Science in Sports & Exercise. 30 (12): 1725–1729. doi:10.1097/00005768-199812000-00011. PMID 9861606.
  80. ^ "National Health Statistics Reports" (PDF). National Health Statistics Reports. 10. 22 October 2008. Retrieved 21 April 2012.
  81. ^ "United States National Health and Nutrition Examination Survey, 1999–2002" (PDF). Retrieved 1 May 2014.
  82. ^ a b Alfred Glucksman (1981). Sexual Dimorphism in Human and Mammalian Biology and Pathology. Academic Press. pp. 66–75. ISBN 978-0-12-286960-0. OCLC 7831448.
  83. ^ Jo Durden-Smith & Diane deSimone (1983). Sex and the Brain. New York: Arbor House. ISBN 978-0-87795-484-2.
  84. ^ Gersh, Eileen S.; Gersh, Isidore (1981). Biology of Women. Baltimore: University Park Press (original from the University of Michigan). ISBN 978-0-8391-1622-6.
  85. ^ Jay H. Stein (1987). Internal Medicine (2nd ed.). Boston: Little, Brown. ISBN 978-0-316-81236-8.
  86. ^ M. McLaughlin & T. Shryer (8 August 1988). "Men vs women: the new debate over sex differences". U.S. News & World Report: 50–58.
  87. ^ B. S. McEwen (1981). "Neural gonadal steroid actions". Science. 210 (4488): 1303–1311. Bibcode:1981Sci...211.1303M. doi:10.1126/science.6259728. PMID 6259728.
  88. ^ David M. Buss (2007). "The evolution of human mating" (PDF). Acta Psychologica Sinica. 39 (3): 502–512.
  89. ^ Martin Daly & Margo Wilson (1996). "Evolutionary psychology and marital conflict". In David M. Buss & Neil M. Malamuth (ed.). Sex, Power, Conflict: Evolutionary and Feminist Perspectives. Oxford University Press. p. 13. ISBN 978-0-19-510357-1.
  90. ^ Christopher Ryan & Cacilda Jethá (2010). Sex at Dawn: The Prehistoric Origins of Modern Sexuality. Harper. ISBN 978-0-06-170780-3.
  91. ^ Alexandra M. Lopes, Norman Ross, James Close, Adam Dagnall, António Amorim & Timothy J. Crow (2006). "Inactivation status of PCDH11X: sexual dimorphisms in gene expression levels in brain". Human Genetics. 119 (3): 1–9. doi:10.1007/s00439-006-0134-0. PMID 16425037.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  92. ^ "Fetal Testosterone Influences Sexually Dimorphic Gray Matter in the Human Brain". jneurosci.org.
  93. ^ "Diverse Roles for Sex Hormone-Binding Globulin in Reproduction". biolreprod.org.
  94. ^ Cordelia Fine (August 2010). Delusions of Gender: How Our Minds, Society, and Neurosexism Create Difference (1st ed.). W. W. Norton & Company. ISBN 978-0-393-06838-2.
  95. ^ Rebecca Jordan-Young (September 2010). Brain Storm: The Flaws in the Science of Sex Differences. Harvard University Press. ISBN 978-0-674-05730-2.
  96. ^ Lisbeth Marner, Jens R. Nyengaard, Yong Tang & Bente Pakkenberg (2003). "Marked loss of myelinated nerve fibers in the human brain with age". The Journal of Comparative Neurology. 462 (2): 144–152. doi:10.1002/cne.10714. PMID 12794739.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  97. ^ Ruben C. Gur, Bruce I. Turetsky, Mie Matsui, Michelle Yan, Warren Bilker, Paul Hughett & Raquel E. Gur (1999). "Sex differences in brain gray and white matter in healthy young adults: correlations with cognitive performance". The Journal of Neuroscience. 19 (10): 4065–4072. PMID 10234034.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  98. ^ Christiana M. Leonard, Stephen Towler, Suzanne Welcome, Laura K. Halderman, Ron Otto, Mark A. Eckert & Christine Chiarello (2008). "Size matters: cerebral volume influences sex differences in neuroanatomy". Cerebral Cortex. 18 (12): 2920–2931. doi:10.1093/cercor/bhn052. PMC 2583156. PMID 18440950.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  99. ^ Eileen Lüders, Helmuth Steinmetz & Lutz Jäncke (2002). "Brain size and grey matter volume in the healthy human brain". NeuroReport. 13 (17): 2371–2374. doi:10.1097/00001756-200212030-00040. PMID 12488829.
  100. ^ Richard J. Haier, Rex E. Jung, Ronald A. Yeo, Kevin Head & Michael T. Alkire (2005). "The neuroanatomy of general intelligence: sex matters" (PDF). NeuroImage. 25 (1): 320–327. doi:10.1016/j.neuroimage.2004.11.019. PMID 15734366. Archived from the original (PDF) on 24 May 2010.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  101. ^ Pollitzer, E. (2013). "Biology: Cell sex matters". Nature. 500 (7460): 23–24. doi:10.1038/500023a. PMID 23903733.
  102. ^ Deasy, B. M.; Lu, A.; Tebbets, J. C.; Feduska, J. M.; Schugar, R. C.; Pollett, J. B.; Sun, B.; Urish, K. L.; Gharaibeh, B. M.; Cao, B.; Rubin, R. T.; Huard, J. (2007). "A role for cell sex in stem cell-mediated skeletal muscle regeneration: Female cells have higher muscle regeneration efficiency". The Journal of Cell Biology. 177 (1): 73–86. doi:10.1083/jcb.200612094. PMC 2064113. PMID 17420291.
  103. ^ Mittelstrass, K.; Ried, J. S.; Yu, Z.; Krumsiek, J.; Gieger, C.; Prehn, C.; Roemisch-Margl, W.; Polonikov, A.; Peters, A.; Theis, F. J.; Meitinger, T.; Kronenberg, F.; Weidinger, S.; Wichmann, H. E.; Suhre, K.; Wang-Sattler, R.; Adamski, J.; Illig, T. (2011). McCarthy, Mark I (ed.). "Discovery of Sexual Dimorphisms in Metabolic and Genetic Biomarkers". PLoS Genetics. 7 (8): e1002215. doi:10.1371/journal.pgen.1002215. PMC 3154959. PMID 21852955.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  104. ^ Penaloza, C.; Estevez, B.; Orlanski, S.; Sikorska, M.; Walker, R.; Smith, C.; Smith, B.; Lockshin, R. A.; Zakeri, Z. (2009). "Sex of the cell dictates its response: Differential gene expression and sensitivity to cell death inducing stress in male and female cells". The FASEB Journal. 23 (6): 1869. doi:10.1096/fj.08-119388.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  105. ^ a b Fritz Vollrath & Geoff A. Parker (1992). "Sexual dimorphism and distorted sex ratios in spiders". Nature. 360 (6400): 156–159. Bibcode:1992Natur.360..156V. doi:10.1038/360156a0.
  106. ^ R. I. Bornholdt, L. R. Oliveira & M. E. Fabián (2008). "Sexual size dimorphism in Myotis nigricans (Schinz, 1821) (Chiroptera: Vespertilionidae) from south Brazil" (PDF). Brazilian Journal of Biology. 68 (4): 897–904. doi:10.1590/S1519-69842008000400028. PMID 19197511.
  107. ^ Virginia Hayssen & T. H. Kunz (1996). "Allometry of litter mass in bats: comparisons with maternal size, wing morphology, and phylogeny" (PDF). Journal of Mammalogy. 77 (2): 476–490. doi:10.2307/1382823. JSTOR 1382823.
  108. ^ Göran Arnqvist, Therésa M. Jones & Mark A. Elgar (2003). "Insect behaviour: reversal of sex roles in nuptial feeding" (PDF). Nature. 424 (6947): 387. Bibcode:2003Natur.424..387A. doi:10.1038/424387a. PMID 12879056. Archived from the original (PDF) on 15 September 2004.
  109. ^ A. Jonathan Shaw (2000). "Population ecology, population genetics, and microevolution". Bryophyte Biology. Cambridge: Cambridge University Press. pp. 379–380. ISBN 978-0-521-66097-6. {{cite book}}: Unknown parameter |editors= ignored (|editor= suggested) (help)
  110. ^ a b Schuster, Rudolf M. (1984). "Comparative Anatomy and Morphology of the Hepaticae". New Manual of Bryology. Vol. 2. Nichinan, Miyazaki, Japan: The Hattori botanical Laboratory. p. 891.
  111. ^ Howard A. Crum & Lewis E. Anderson (1980). Mosses of Eastern North America. Vol. 1. New York: Columbia University Press. p. 196. ISBN 978-0-231-04516-2.
  112. ^ D. Briggs (1965). "Experimental taxonomy of some British species of genus Dicranum". New Phytologist. 64 (3): 366–386. doi:10.1111/j.1469-8137.1965.tb07546.x. JSTOR 2430169.
  113. ^ Olsen, Jerry; Olsen, Penny (5 August 1986). "Sexual Size Dimorphism in Raptors: Intrasexual Competition in the Larger Sex for a Scarce Breeding Resource, the Smaller Se". EMU. 87. {{cite journal}}: |access-date= requires |url= (help)
  114. ^ Deborah Charlesworth & Judith E. Mank (2010). "The birds and the bees and the flowers and the rrees: lessons from genetic mapping of sex determination in plants and animals". Genetics. 186 (1): 9–31. doi:10.1534/genetics.110.117697.
  115. ^ G. A. Parker (1982). "Why are there so many tiny sperm? Sperm competition and the maintenance of two sexes". Journal of Theoretical Biology. 96 (2): 281–294. doi:10.1016/0022-5193(82)90225-9.
  116. ^ Jiang-Nan Yang (2010). "Cooperation and the evolution of anisogamy". Journal of Theoretical Biology. 264 (1): 24–36. doi:10.1016/j.jtbi.2010.01.019. PMID 20097207.
  117. ^ G. Bell (1985). "On the function of flowers". Proceedings of the Royal Society B: Biological Sciences. 224 (1235): 223–266. Bibcode:1985RSPSB.224..223B. doi:10.1098/rspb.1985.0031. JSTOR 36033.
  118. ^ Futuyma 2005, p. 330
  119. ^ Futuyma 2005, p. 331
  120. ^ Futuyma 2005, p. 332
  121. ^ a b Ridley 2004, p. 328
  122. ^ Futuyma 2005, p. 335
  123. ^ Ridley 2004, p. 330
  124. ^ Ridley 2004, p. 332

Bibliography

Further reading

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