Sexual dimorphism is the condition where sexes of the same

species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...

exhibit different morphological characteristics, particularly characteristics not directly involved in

reproduction Reproduction (or procreation or breeding) is the biological process by which new individual organisms – " offspring" – are produced from their "parent" or parents. Reproduction is a fundamental feature of all known life; each individual o ...

. The condition occurs in most

dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproducti ...

species, which consist of most animals and some plants. Differences may include

secondary sex characteristic Secondary sex characteristics are features that appear during puberty in humans, and at sexual maturity in other animals. These characteristics are particularly evident in the sexually dimorphic phenotypic traits that distinguish the sexes of ...

s, size, weight, color, markings, or behavioral or cognitive traits. Male-male reproductive competition has evolved a diverse array of sexually dimorphic traits. Aggressive utility traits such as "battle" teeth and blunt heads reinforced as battering rams are used as weapons in aggressive interactions between rivals. Passive displays such as ornamental feathering or song-calling have also evolved mainly through sexual selection. These differences may be subtle or exaggerated and may be subjected to

sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (in ...

and

natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...

. The opposite of dimorphism is ''monomorphism'', when both biological sexes are phenotypically indistinguishable from each other.

Overview

Ornamentation and coloration

Common and easily identified types of dimorphism consist of ornamentation and coloration, though not always apparent. A difference in the coloration of sexes within a given species is called sexual dichromatism, commonly seen in many species of birds and reptiles.

Sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (in ...

leads to 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. The peafowl constitute conspicuous illustrations of the principle. The ornate

plumage Plumage ( "feather") is a layer of feathers that covers a bird and the pattern, colour, and arrangement of those feathers. The pattern and colours of plumage differ between species and subspecies and may vary with age classes. Within species, ...

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 is of a subdued brown coloration. 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. Similar examples are manifold, such as in birds of paradise and argus pheasants. Another example of sexual dichromatism is that of 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

carotenoid Carotenoids (), also called tetraterpenoids, are yellow, orange, and red organic pigments that are produced by plants and algae, as well as several bacteria, and fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips, cor ...

lutein Lutein (;"Lutein"

zeaxanthin Zeaxanthin is one of the most common carotenoids in nature, and is used in the xanthophyll cycle. Synthesized in plants and some micro-organisms, it is the pigment that gives paprika (made from bell peppers), corn, saffron, goji (wolfberries), ...

. This diet also affects the sexually dimorphic colours in the human-invisible

ultraviolet Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30 PHz) to 400 nm (750 THz), shorter than that of visible light, but longer than X-rays. UV radiati ...

spectrum. 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. 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. Carotenoids play an important role in

immune function The immune system is a network of biological processes that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to parasitic worms, as well as cancer cells and objects such as wood splin ...

for many animals, so carotenoid dependent signals might indicate health. 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. ''Ranoidea lesueuri'' is an example of a dynamic frog with temporary color changes in males during the breeding season. ''

Hyperolius ocellatus ''Hyperolius ocellatus'' is a species of tropical West African frog in the family Hyperoliidae, that is split into the subspecies ''H. o. ocellatus'' and ''H. o. purpurescens''. It is found in Angola, Cameroon, Central African Republic, Republi ...

'' 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. In contrast, the females are rusty red to silver with small spots. The bright coloration in the male population attracts females and is an aposematic sign to potential predators. Females often show a preference for exaggerated male

secondary sexual characteristics Secondary sex characteristics are features that appear during puberty in humans, and at sexual maturity in other animals. These characteristics are particularly evident in the sexually dimorphic phenotypic traits that distinguish the sexes of a ...

in mate selection. The sexy son hypothesis explains that females prefer more elaborate males and select against males that are dull in color, independent of the species' vision. Similar sexual dimorphism and mating choice are also observed in many fish species. For example, male guppies have colorful spots and ornamentations, while females are generally grey. Female guppies prefer brightly colored males to duller males. 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.

Plants

Most

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. They include all forbs (flowering plants without a woody stem), grasses and grass-like plants, a vast majority of ...

s are hermaphroditic but approximately 6% of species have separate males and females (

dioecy Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...

). Sexual dimorphism is common in dioecious plants and dioicous species. 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 A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanism ...

, completing

pollination Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds ...

. '' Catasetum'' orchids are one interesting exception to this rule. Male ''Catasetum''

orchids Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Along with the Asteraceae, they are one of the two largest families of flowering ...

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. Various other dioecious exceptions, such as '' Loxostylis alata'' have visibly different sexes, with the effect of eliciting the most efficient behavior 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 ''Geranium'' is a genus of 422 species of annual, biennial, and perennial plants that are commonly known as geraniums or cranesbills. They are found throughout the temperate regions of the world and the mountains of the tropics, but mostly i ...

'' have what amounts to serial sexual dimorphism. The flowers of such species might, for example, present their

anthers The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...

on opening, then shed the exhausted anthers after a day or two and perhaps change their colours as well while the

pistil Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) '' pis ...

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 once fertilized, thereby discouraging further visits from pollinators. This is advantageous to both parties because it avoids damaging the developing fruit and wasting the pollinator's effort on unrewarding visits. In effect, the strategy ensures that pollinators can expect a reward every time they visit an appropriately advertising flower. Females of the aquatic plant ''

Vallisneria americana ''Vallisneria americana'', commonly called wild celery, water-celery, tape grass, or eelgrass, is a plant in the family Hydrocharitaceae, the "tape-grasses". ''V. americana'' is a fresh water species that can tolerate salt, living in saliniti ...

'' 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. Sexual dimorphism is most often associated with wind-pollination in plants due to selection for efficient

pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...

dispersal in males vs pollen capture in females, e.g. ''Leucadendron rubrum''. 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. p. 206 Every sexually reproducing extant species of the vascular plant has an alternation of generations; the plants we see about us generally are

diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respecti ...

sporophytes, but their offspring are not the seeds that people commonly recognize as the new generation. The seed actually is the offspring of the haploid generation of microgametophytes (

) and megagametophytes (the embryo sacs in the

ovules In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the '' integument'', forming its outer layer, the '' nucellus'' (or remnant of the megasporangium), and the ...

). 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

Insects display a wide variety of sexual dimorphism between taxa including size, ornamentation and coloration. The female-biased sexual size dimorphism observed in many taxa evolved despite intense male-male competition for mates. 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. In the

hackberry emperor ''Asterocampa celtis'', the hackberry emperor, is a North American butterfly that belongs to the brushfooted butterfly family, Nymphalidae. It gets its name from the hackberry tree (''Celtis occidentalis'' and others in the genus ''Celtis'') upon ...

females are similarly larger than males. The reason for the sexual dimorphism is due to provision size mass, in which females consume more pollen than males. In some species, there is evidence of male dimorphism, but it appears to be for distinctions of roles. This is seen in the bee species '' Macrotera portalis'' in which there is a small-headed morph, capable of flight, and large-headed morph, incapable of flight, for males. '' 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. Another example is '' Lasioglossum hemichalceum'', which is a species of sweat bee that shows drastic physical dimorphisms between male offspring. Not all dimorphism has to have a drastic difference between the sexes. ''

Andrena agilissima ''Andrena agilissima'' is a species of mining bee. They are present in most of Europe, the Near East and North Africa and can be found from April through July. ''Andrena agilissima'' is an ''oligolectic'' species, feeding only on the pollen of a ...

'' is a mining bee where the females only have a slightly larger head than the males. Weaponry leads to increased fitness by increasing success in male–male competition in many insect species. 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. These structures are impressive because of the exaggerated sizes. There is a direct correlation between male horn lengths and body size and higher access to mates and fitness. In other beetle species, both males and females may have ornamentation such as horns. Generally, insect sexual size dimorphism (SSD) within species increases with body size. 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. The sexual dichromatic nature of '' Bicyclus anynana'' is reflected by female selection on the basis of dorsal UV-reflective eyespot pupils. The

common brimstone ''Gonepteryx rhamni'' (known as the common brimstone) is a butterfly of the family Pieridae. It lives throughout the Palearctic zone and is commonly found across Europe, Asia, and North Africa. Across much of its range, it is the only species ...

also displays sexual dichromatism; males have yellow and iridescent wings, while female wings are white and non-iridescent. Naturally selected deviation in protective female coloration is displayed in mimetic butterflies.

Spiders and sexual cannibalism

Many

arachnid Arachnida () is a class of joint-legged invertebrate animals ( arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and ...

groups exhibit sexual dimorphism, but it is most widely studied in the spiders. In the orb-weaving spider '' Zygiella x-notata'', for example, adult females have a larger body size than adult 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 Wolf spiders are members of the family Lycosidae (). They are robust and agile hunters with excellent eyesight. They live mostly in solitude, hunt alone, and do not spin webs. Some are opportunistic hunters, pouncing upon prey as they find it or ...

Tigrosa helluo ''Tigrosa helluo'' is a species of spider belonging to the family Lycosidae, also known as wolf spiders. ''T. helluo'' was formerly known as ''Hogna helluo'' before differences between dorsal color patterns, habitat preferences, body structu ...

'', food-limited females cannibalize more frequently. 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 Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the natural capability to pr ...

and lower rates of cannibalism. 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 Orb-weaver spiders are members of the spider family Araneidae. They are the most common group of builders of spiral wheel-shaped webs often found in gardens, fields, and forests. The English word "orb" can mean "circular", hence the English name ...

. 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 female's web, mating while the female is feeding, or providing a nuptial gift in response to sexual cannibalism. 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. In the species '' Maratus volans'', the males are known for their characteristic colorful fan which attracts the females during mating.

Fish

Ray-finned fish are an ancient and diverse class, with the widest degree of sexual dimorphism of any animal class. Fairbairn notes that "females are generally larger than males but males are often larger in species with male–male combat or male paternal care ... izes rangefrom dwarf males to males more than 12 times heavier than females." There are 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. 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. 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. 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. 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. Another example is the dragonet, in which males are considerably larger than females and possess longer fins. 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. In protogynous mating systems where males dominate mating with many females, size plays a significant role in male reproductive success. 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. 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 a 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 A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces ...

size, which accounts for about 25% of body mass. Sexual selection was observed for female ornamentation in '' Gobiusculus flavescens'', known as two-spotted gobies. 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. Since carotenoid-based ornamentation suggests mate quality, female two-spotted guppies that develop colorful orange bellies during breeding season are considered favorable to males. The males invest heavily in offspring during incubation, which leads to the sexual preference in colorful females due to higher egg quality.

Amphibians and non-avian 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 Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia a ...

; coloration as in many

amphibians Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arb ...

snakes Snakes are elongated, limbless, carnivorous reptiles of the suborder Serpentes . Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more j ...

, and lizards, as well as in some turtles; 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.

Anole Dactyloidae are a family of lizards commonly known as anoles () and native to warmer parts of the Americas, ranging from southeastern United States to Paraguay. Instead of treating it as a family, some authorities prefer to treat it as a subfami ...

lizards show prominent size dimorphism with males typically being significantly larger than females. For instance, the average male ''

Anolis sagrei The brown anole (''Anolis sagrei''), also known commonly as the Cuban brown anole, or De la Sagra's anole, is a species of lizard in the family Dactyloidae. The species is native to Cuba and the Bahamas. It has been widely introduced elsewhere, ...

'' was 53.4 mm vs. 40 mm in females. Different sizes of the heads in anoles have been explained by differences in the estrogen pathway. 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.Pinto, A., Wiederhecker, H., & Colli, G. (2005). Sexual dimorphism in the Neotropical lizard, Tropidurus torquatus (Squamata, Tropiduridae). Amphibia-Reptilia. 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''. Sexual dimorphism in size is also seen in frog species like '' P. bibronii''. Male painted dragon lizards, '' Ctenophorus pictus''. are brightly conspicuous in their breeding coloration, but male colour declines with

aging Ageing ( BE) or aging ( AE) is the process of becoming older. The term refers mainly to humans, many other animals, and fungi, whereas for example, bacteria, perennial plants and some simple animals are potentially biologically immortal. In ...

. Male coloration appears to reflect innate anti-oxidation capacity that protects against

oxidative DNA damage DNA oxidation is the process of oxidative damage of deoxyribonucleic acid. As described in detail by Burrows et al., 8-oxo-2'-deoxyguanosine (8-oxo-dG) is the most common oxidative lesion observed in duplex DNA because guanine has a lower one-e ...

. Male breeding coloration is likely an indicator to females of the underlying level of oxidative DNA damage (a significant component of aging) in potential mates.

Birds

Avian dinosaurs

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, e.g.

birds of prey Birds of prey or predatory birds, also known as raptors, are hypercarnivorous bird species that actively hunt and feed on other vertebrates (mainly mammals, reptiles and other smaller birds). In addition to speed and strength, these predato ...

hummingbird Hummingbirds are birds native to the Americas and comprise the biological family Trochilidae. With about 361 species and 113 genera, they occur from Alaska to Tierra del Fuego, but the vast majority of the species are found in the tropics ar ...

s, and some species of flightless birds. Plumage dimorphism, in the form of ornamentation or coloration, also varies, though males are typically the more ornamented or brightly colored sex. Such differences have been attributed to the unequal reproductive contributions of the sexes. 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 (or only) caregiver. Plumage polymorphisms have evolved to reflect these differences and other measures of reproductive fitness, such as body condition or survival. 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. 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. 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 size dimorphism in species. It has been shown that the larger males are better at coping with the difficulties of migration and thus are more successful in reproducing when reaching the breeding destination. When viewing this from an evolutionary standpoint, many theories and explanations come into 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. 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. 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. Presumably, increased sexual dimorphism means males are brighter and more conspicuous, leading to increased predation. Moreover, the production of more exaggerated ornaments in males may come at the cost of suppressed immune function. 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 dimorphism alive in these species and ensures that the next generation of successful males will also display these traits that are attractive to 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. Activational hormones occur during puberty and adulthood and serve to 'activate' certain behaviors when appropriate, such as territoriality during breeding season. 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. Such behavioral differences can cause disproportionate sensitivities to anthropogenic pressures. Females of the whinchat in Switzerland breed in intensely managed grasslands. Earlier harvesting of the grasses during the breeding season lead to more female deaths. Populations of many birds are often male-skewed and when sexual differences in behavior increase this ratio, populations decline at a more rapid rate. Also not all male dimorphic traits are due to hormones like testosterone, instead they are a naturally occurring part of development, for example plumage. In addition, the strong hormonal influence on phenotypic differences suggests that the genetic mechanism and genetic basis of these sexually dimorphic traits may involve transcription factors or cofactors rather than regulatory sequences. Sexual dimorphism may also influence differences in parental investment during times of food scarcity. For example, in the

blue-footed booby The blue-footed booby (''Sula nebouxii'') is a marine bird native to subtropical and tropical regions of the eastern Pacific Ocean. It is one of six species of the genus ''Sula'' – known as boobies. It is easily recognizable by its distincti ...

, 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. In Black-tailed Godwits ''Limosa limosa limosa'' females are also the larger sex, and the growth rates of female chicks are more susceptible to limited environmental conditions. 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-breeding season. This occurs because the species is more focused on survival than on 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. Most sexual segregation research has been done on ungulates, but such research extends to bats,

kangaroo Kangaroos are four marsupials from the family Macropodidae (macropods, meaning "large foot"). In common use the term is used to describe the largest species from this family, the red kangaroo, as well as the antilopine kangaroo, eastern ...

s, and birds. Sex-specific conservation plans have even been suggested for species with pronounced sexual segregation. The term sesquimorphism (the Latin

numeral prefix Numeral or number prefixes are prefixes derived from numerals or occasionally other numbers. In English and many other languages, they are used to coin numerous series of words. For example: * unicycle, bicycle, tricycle (1-cycle, 2-cycle, 3-cyc ...

''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 Examples include Cape sparrow (''Passer melanurus''), rufous sparrow (subspecies ''P. motinensis motinensis''), and saxaul sparrow (''P. ammodendri'').

Non-avian dinosaurs

Sexual dimorphism is thought to have been present in non-avian dinosaurs.

Mammals

In a large proportion of mammal species, males are larger than females. Both genes and

hormone A hormone (from the Greek participle , "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs by complex biological processes to regulate physiology and behavior. Hormones are required ...

s 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."

Pinnipeds

Male and female Mirounga angustirostris 2

Marine mammals show some of the greatest sexual size differences of mammals, because of sexual selection and environmental factors like breeding location. The mating system of pinnipeds varies from polygamy 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. For example, the males are significantly larger (about 10% heavier and 2% longer) than the females at birth in sea lion pups. The pattern of differential investment can be varied principally prenatally and post-natally. ''Mirounga leonina'', the southern elephant seal, is one of the most dimorphic mammals.

Primates

Humans

According to Clark Spencer Larsen, modern day ''

Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...

'' show a range of sexual dimorphism, with average body mass between the sexes differing by roughly 15%. Substantial discussion in academic literature considers potential evolutionary advantages associated with sexual competition (both intrasexual and intersexual), as well as short- and long-term sexual strategies. According to Daly and Wilson, "The sexes differ more in human beings than in monogamous mammals, but much less than in extremely polygamous mammals." The average

basal metabolic rate Basal metabolic rate (BMR) is the rate of energy expenditure per unit time by endothermic animals at rest. It is reported in energy units per unit time ranging from watt (joule/second) to ml O2/min or joule per hour per kg body mass J/(h·kg). P ...

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 Skeletal muscles (commonly referred to as muscles) are Organ (biology), organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other ...

and expendable circulating energy reserves. According to Tim Hewett, director of research in the department of sports medicine at

Ohio State University Wexner Medical Center The Ohio State University Wexner Medical Center is a multidisciplinary academic medical center located in Columbus, Ohio, United States, on the main campus of The Ohio State University. For 29 consecutive years, '' U.S. News & World Report'' has ...

, females have, on average, 50–60% of the upper body strength of males, and 80-90% of the lower body strength of males, relative to body size, but females have higher

endurance Endurance (also related to sufferance, resilience, constitution, fortitude, and hardiness) is the ability of an organism to exert itself and remain active for a long period of time, as well as its ability to resist, withstand, recover from an ...

than males. 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. 492 kg vs 348 kg for unlimited weight classes; see

Olympic weightlifting records Olympic or Olympics may refer to Sports Competitions * Olympic Games, international multi-sport event held since 1896 ** Summer Olympic Games ** Winter Olympic Games * Ancient Olympic Games, ancient multi-sport event held in Olympia, Greece b ...

). A study, carried out by analyzing annual world rankings from 1980 to 1996, found that males' running times were, on average, 11% faster than females'. In early adolescence, females are on average taller than males (as females tend to go through puberty earlier), but males, on average, surpass them in height in later adolescence and adulthood. In the United States, adult males are on average 9% taller and 16.5% heavier than adult females. Males typically have larger

tracheae The trachea, also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air- breathing animals with lungs. The trachea extends from the ...

and branching bronchi, with about 30 percent greater

lung volume Lung volumes and lung capacities refer to the volume of air in the lungs at different phases of the respiratory cycle. The average total lung capacity of an adult human male is about 6 litres of air. Tidal breathing is normal, resting breath ...

per body mass. On average, males have larger

heart The heart is a muscular organ found in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon diox ...

s, 10 percent higher

red blood cell Red blood cells (RBCs), also referred to as red cells, red blood corpuscles (in humans or other animals not having nucleus in red blood cells), haematids, erythroid cells or erythrocytes (from Greek ''erythros'' for "red" and ''kytos'' for "holl ...

count, higher

hemoglobin Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyte ...

, hence greater oxygen-carrying capacity. They also have higher circulating clotting factors (

vitamin K Vitamin K refers to structurally similar, fat-soluble vitamers found in foods and marketed as dietary supplements. The human body requires vitamin K for post-synthesis modification of certain proteins that are required for blood coagulat ...

, pro thrombin and platelets). These differences lead to faster healing of

wound A wound is a rapid onset of injury that involves lacerated or punctured skin (an ''open'' wound), or a contusion (a ''closed'' wound) from blunt force trauma or compression. In pathology, a ''wound'' is an acute injury that damages the epider ...

s and lower sensitivity to nerve

pain Pain is a distressing feeling often caused by intense or damaging stimuli. The International Association for the Study of Pain defines pain as "an unpleasant sensory and emotional experience associated with, or resembling that associated with, ...

after injury. In males, pain-causing injury to the peripheral nerve occurs through the

microglia Microglia are a type of neuroglia (glial cell) located throughout the brain and spinal cord. Microglia account for about 7% of cells found within the brain. As the resident macrophage cells, they act as the first and main form of active immune ...

, while in females it occurs through the

T cell A T cell is a type of lymphocyte. T cells are one of the important white blood cells of the immune system and play a central role in the adaptive immune response. T cells can be distinguished from other lymphocytes by the presence of a T-cell ...

s (except in pregnant women, who follow a male pattern). Females typically have more white blood cells (stored and circulating), as well as more

granulocyte Granulocytes are cells in the innate immune system characterized by the presence of specific granules in their cytoplasm. Such granules distinguish them from the various agranulocytes. All myeloblastic granulocytes are polymorphonuclear. They ...

s 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.

Ethologists Ethology is the scientific study of animal behaviour, usually with a focus on behaviour under natural conditions, and viewing behaviour as an evolutionarily adaptive trait. Behaviourism as a term also describes the scientific and object ...

argue that females, interacting with other females and multiple offspring in social groups, have experienced such traits as a selective advantage. Females have a higher sensitivity to pain due to aforementioned nerve differences that increase the sensation, and females thus require higher levels of pain medication after injury. Hormonal changes in females affect pain sensitivity, and pregnant women have the same sensitivity as males. Acute pain tolerance is also more consistent over a lifetime in females than males, despite these hormonal changes. Despite differences in physical feeling, both sexes have similar psychological tolerance to (or ability to

cope The cope (known in Latin as ''pluviale'' 'rain coat' or ''cappa'' 'cape') is a liturgical vestment, more precisely a long mantle or cloak, open in front and fastened at the breast with a band or clasp. It may be of any liturgical colours, litu ...

with and ignore) pain. In the

human brain The human brain is the central organ (anatomy), organ of the human nervous system, and with the spinal cord makes up the central nervous system. The brain consists of the cerebrum, the brainstem and the cerebellum. It controls most of the act ...

, a difference between sexes was observed in the transcription of the PCDH11X/Y gene pair unique to ''Homo sapiens''. Sexual differentiation in the human brain from the undifferentiated 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. The brains of pregnant females carrying male fetuses may be shielded from the masculinizing effects of androgen through the action of sex hormone-binding globulin. The relationship between sex differences in the brain and human behavior is a subject of controversy in psychology and society at large. Many females tend to have a higher ratio of gray matter in the left hemisphere of the brain in comparison to males. Males on average have larger brains than females; however, when adjusted for total brain volume, the gray matter differences between sexes are almost nonexistent. Thus, the percentage of gray matter appears to be more related to brain size than it is to sex. 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". (See the sex and intelligence article for more on this subject.) Strict graph-theoretical analysis of the human brain connections revealed that in numerous graph-theoretical parameters (e.g., minimum bipartition width, edge number, the expander graph property, minimum vertex cover), the structural connectome of women are significantly "better" connected than the connectome of men. It was shown that the graph-theoretical differences are due to the sex and not to the differences in the cerebral volume, by analyzing the data of 36 females and 36 males, where the brain volume of each man in the group was smaller than the brain volume of each woman in the group. Sexual dimorphism was also described in the gene level and shown to extend from the sex chromosomes. Overall, about 6500 genes have been found to have sex-differential expression in at least one tissue. Many of these genes are not directly associated with reproduction, but rather linked to more general biological features. In addition, it has been shown that genes with sex-specific expression undergo reduced selection efficiency, which leads to higher population frequencies of deleterious mutations and contributes to the prevalence of several human diseases.

Immune function

Sexual dimorphism in immune function is a common pattern in vertebrates and also in a number of invertebrates. Most often, females are more 'immunocompetent' than males. This trait is not consistent among all animals, but differs depending on taxonomy, with the most female-biased immune systems being found in insects. In mammals this results in more frequent and severe infections in males and higher rates of autoimmune disorders in females. One potential cause may be differences in gene expression of immune cells between the sexes. Another explanation is that endocrinological differences between the sexes impact the immune system – for example, testosterone acts as an immunosuppressive agent.

Cells

Phenotypic differences between sexes are evident even in

cultured cells Cell culture or tissue culture is the process by which cells are grown under controlled conditions, generally outside of their natural environment. The term "tissue culture" was coined by American pathologist Montrose Thomas Burrows. This te ...

from tissues. For example, female muscle-derived stem cells have a better muscle regeneration efficiency than male ones. There are reports of several metabolic differences between male and female cells and they also respond to stress differently.

Reproductively advantageous

In theory, larger females are favored by competition for mates, especially in polygamous species. Larger females offer an advantage in fertility, since the physiological demands of reproduction are limiting in females. Hence there is a theoretical expectation that females tend to be larger in species that are monogamous. Females are larger in many species of

insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs ...

s, many

spider Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species d ...

s, many

fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% ...

, many reptiles, owls, birds of prey and certain mammals such as the

spotted hyena The spotted hyena (''Crocuta crocuta''), also known as the laughing hyena, is a hyena species, currently classed as the sole extant member of the genus '' Crocuta'', native to sub-Saharan Africa. It is listed as being of least concern by the ...

, and baleen whales such as

blue whale The blue whale (''Balaenoptera musculus'') is a marine mammal and a baleen whale. Reaching a maximum confirmed length of and weighing up to , it is the largest animal known to have ever existed. The blue whale's long and slender body can b ...

. As an example, in some species, females are sedentary, and so males must search for them. Fritz Vollrath and

Geoff Parker Professor Geoffrey Alan Parker FRS (born 24 May 1944) is an emeritus professor of biology at the University of Liverpool and the 2008 recipient of the Darwin Medal. Parker has been called “the professional’s professional”. He has a ...

argue that this difference in behaviour leads to radically different selection pressures on the two sexes, evidently favouring smaller males. Cases where the male is larger than the female have been studied as well, and require alternative explanations. One example of this type of sexual size dimorphism is the bat ''

Myotis nigricans The black myotis (''Myotis nigricans'') is a vesper bat species from South and Central America. Its body is dark brown/black. The head to body length, not including the tail, is about 5 cm (2 in.). The black myotis is a tiny bat with a smal ...

'', (black myotis bat) where females are substantially larger than males in terms of body weight, skull measurement, and forearm length. The interaction between the sexes and the energy needed to produce viable offspring makes 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. 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. Cryptopsaras couesii (triplewart seadevil)

Cryptopsaras couesii (triplewart seadevil)

Some species of

anglerfish The anglerfish are fish of the teleost order Lophiiformes (). They are bony fish named for their characteristic mode of predation, in which a modified luminescent fin ray (the esca or illicium) acts as a lure for other fish. The luminescence ...

also display extreme sexual dimorphism. Females are more typical in appearance to other fish, whereas 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 in what amounts to an effectively hermaphrodite composite organism. 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 (although males can survive away from females, they generally are not free-living). This is taken to the logical extreme in the Rhizocephala crustaceans, like the Sacculina, where the male injects itself into the female's body and becomes nothing more than sperm producing cells, to the point that the superorder used to be mistaken for hermaphroditic. Some plant species also exhibit dimorphism in which the females are significantly larger than the males, such as in the moss '' Dicranum'' and the liverwort '' Sphaerocarpos''. There is some evidence that, in these genera, the dimorphism may be tied to a sex chromosome, or to chemical signalling from females. Another complicated example of sexual dimorphism is in '' Vespula squamosa'', the 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 workers and male counterparts.

Evolution

Eccaparadoxides mediterraneus - Murero, Zaragoza - Museo Ciencias Naturales Universidad Zaragoza

In 1871,

Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...

advanced the theory of sexual selection, which related sexual dimorphism to

. The first step towards sexual dimorphism is the size differentiation of sperm and eggs ( anisogamy). 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 Sperm competition is the competitive process between spermatozoa of two or more different males to fertilize the same egg during sexual reproduction. Competition can occur when females have multiple potential mating partners. Greater choice and ...

, because small sperm enable organisms to produce a large number of sperm, and make males (or male function of hermaphrodites) more redundant. This intensifies male competition for mates and promotes the evolution of other sexual dimorphism in many species, especially in vertebrates including

mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fu ...

. However, in some species females compete for mates in ways more usually associated with males (usually species in which males invest a lot in rearing offspring and thus are no longer considered as so redundant). Sexual dimorphism by size is evident in some extinct species such as the velociraptor. In the case of velociraptors the sexual size dimorphism may have been caused by two factors: male competition for hunting grounds to attract mates, and/or female competition for nesting locations and mates, males being a scarce breeding resource. Volvocine algae have been useful in understanding the evolution of sexual dimorphism and species like the beetle '' C. maculatus'', where the females are larger than the males, are used to study its underlying genetic mechanisms. 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. In these species, there is a

selection pressure Any cause that reduces or increases reproductive success in a portion of a population potentially exerts evolutionary pressure, selective pressure or selection pressure, driving natural selection. It is a quantitative description of the amount of ...

for whatever traits enable a male to have more matings. The male may therefore come to have different traits from the female. Sumatran orangutan family in Toronto Zoo

Sumatran orangutan family in Toronto Zoo

These traits could be ones that allow him to fight off other males for control of territory or a

harem Harem ( Persian: حرمسرا ''haramsarā'', ar, حَرِيمٌ ''ḥarīm'', "a sacred inviolable place; harem; female members of the family") refers to domestic spaces that are reserved for the women of the house in a Muslim family. A har ...

, such as large size or weapons; or they could be traits that females, for whatever reason, prefer in mates. Male–male competition poses no deep theoretical questions but mate choice does. Females may choose males that appear strong and healthy, thus likely to possess "good

allele An allele (, ; ; modern formation from Greek ἄλλος ''állos'', "other") is a variation of the same sequence of nucleotides at the same place on a long DNA molecule, as described in leading textbooks on genetics and evolution. ::"The chro ...

s" and give rise to healthy offspring. In some species, however, 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). 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. 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.

References

Sources

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External links

* {{DEFAULTSORT:Sexual Dimorphism Animal anatomy Sexual selection Polymorphism (biology) Asymmetry