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One of the most fascinating aspects of human life is how we choose our mates. Animals also choose their mates, sometimes with a great deal of care. Mating systems are important to understand because they reect the result of natural selection on mate choice, and ultimately on strategies for maximizing individual reproductive success.
A mating system describes how males and females pair when choosing a mate. Males and females di!er greatly in the investment each makes to reproduce, and may therefore approach mating with di!ering strategies. To study these di!erences, scientists observe mating systems and describe how males and females come together. When choosing mates, animals evolve species-typical strategies for maximizing their reproductive success this results in considerable diversity among animal species in their mating patterns. In this article we rst discuss why sexual reproduction exists, and how di!erences between males and females a!ect mating systems. We move on to consider the evolution of mate choice, and then we describe the types of mating systems found in animals.
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is stable. When the environment is going to turn cold, most species of aphids reproduce sexually, because sexual reproduction produces eggs that are freeze tolerant and can diapause during the winter (Simon et al. 2002). Genetic diversity may also lead to evolved defenses against parasites and disease. The mud snail, Potamopyrgus antipodarum, is host to several trematode parasites. Sexual individuals of this species are more common in areas where risk of trematode infection is high. In areas where the risk of infection is low, asexual individuals have displaced sexual ones (King et al. 2009). This suggests that the genetic diversity acquired from sexual reproduction is necessary for this species to defend against parasites, as asexual individuals may not easily survive in areas where parasites are high. Sexual reproduction often involves evolutionary di!erentiation of males and females. Females typically produce signicantly fewer gametes (eggs) than males and invest heavily in each one. On the other hand, males produce many gametes (sperm) and invest little into each one. These strong di!erences in gamete investment between the sexes leads to reproductive strategies between the sexes that, in some cases, conict. Females may spend more care than males selecting a mate due to the high cost of their gametes.
Figure 1: A male bighorn sheep. The large horns are used in combat between males during mating season, and likely evolved as a result of intrasexual selection. 2012
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Bateman's principle postulates that variance among females in mating success is low, whereas variance among males in mating success is high. This stems from the fact that one mating in females should be enough to fertilize all their eggs whereas in males reproductive success is based on the number of times they have mated. In other words, nearly all females in a population mate and have o!spring, but relatively few males mate successfully (Figure 2). Those males that do mate tend to mate with many females-thus a few males have very high reproductive output, but many males have little or no reproductive output (Bateman 1948). This leads to the prediction that sexual selection should act more strongly on males, leading to greater elaboration of behavior and structures used in attracting mates in males than in females.
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These gures illustrate Batemans principle after one mating, female mating frequency increases and relative tness remains constant, as the sperm from one mating is adequate to fertilize all the females eggs. In males, as mating frequency increases relative tness also increases proportionally. 2012
Nature Education Modied from Hauber & Lacey 2005. All rights reserved.
Criticisms of Bateman's theory focus on the generality of the predictions. Contrary to the predictions of Bateman's principle, there are several possible advantages to female multiple matings. The female cichlid sh Pseudotropheus spiliopterus mates with any male they meet because they have a high risk of getting predated and a small population. This often leads to multiple matings by a single female (Kellogg et al.1998). Mating with any male that is seen ensures that these cichlids have a chance at producing o!spring. The female Malawi blue cichlid has a high population but still participates in multiple matings. In this case multiple matings occur to avoid inbreeding and increase genetic diversity among the o!spring (Kellogg et al. 1998). Additionally, multiple matings by females may increase the likelihood that they will nd a compatible mate, one that is not sterile, or even help prevent infanticide.
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Figure 3: Eyespots from peacock tail feathers. Elaborate ornamentation usually evolves in intrasexual selection and is used in mate choice. 2012
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care and territorial resources, are maintained by having only one mate at a time, or by concealing extra-pair partnerships.
Figure 4: Blue-footed boobies. Many bird species, such as these blue-footed boobies are monogamous. 2012
Polygyny
Polygyny is the association of one male with multiple females. This mating system is found in a few birds and insects, but is most common in mammals. Polygyny is a strategy used by males to increase their reproductive tness.
Harems
Another common type of polygyny is membership in a harem, a defended group of females associated with one male. Females may initially associate in a harem for group defense, or they may be herded together by a male. Males compete for control of the groups. Harems typically exhibit a dominance hierarchy among the females in the group.
Leks
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A lek is an aggregation of males that are each seeking to attract a mate. Within a lek, males typically perform sexual displays. Unlike most other mating systems, leks are not associated with resources. Aggregations of males may be near particularly attractive females or in areas where females are likely to travel (Lank et al. 1995, Aspbury & Gibson 2004). It is thought that males form leks because they attract more females than do isolated males. Attracting more females is a strategy used by males to help increase their reproductive success.
Polyandry
Polyandry is a group with one female and many males. Polyandry is a reproductive strategy that helps a female ensure reproductive success by providing her with multiple mating options.
Cooperative Polyandry
The Galapagos hawk exhibits cooperative polyandry. In this case all males in the group copulate with the female and all participate in brood provisioning (Fabborg et al. 1995).
Polygynandry
Some mating systems have looser male-female bonds within groups. In polygynandrous groups, multiple females and males mate with each other, and males may care for the broods of several females. Chimpanzees and bonobos rely on this strategy it allows groups of males and females to live together and spend less time being concerned with mate competition. Polygynandry may be advantageous from the female's perspective because it causes paternity confusion, which decreases infanticide and allows her to have multiple males care for her brood (Hrdy 1981, 2000).
Promiscuity
In promiscuity there are no pair bonds, and males and females, although sometimes choosy, often seem to mate randomly. As it is typically more advantageous for one or both sexes to pick their mate, promiscuity may occur in species for which the environment is unpredictable (Birkhead 2000, Burton 2002).
Sperm Competition
Although sperm competition is not a type of mating system per se, it is a form of male-male competition that plays an important role in mating systems. If more than one male mates with a female in a short time period, competition can occur after the males have released their sperm (Fisher & Hoekstra 2010). In other words, once a male has released sperm, its sperm must be the rst to reach an egg. This is often apparent in animals that use external fertilization. In aquatic animals that release their gametes into the water, animals that release the largest amount of sperm, and sperm that are highly capable of swimming, are likely to produce the most o!spring (Stoltz & Ne! 2006). Animals with internal fertilization also experience sperm competition. Several mechanisms have evolved to facilitate a male's reproductive success with females that have multiple mates. For example, in one species of damsely, males physically remove any sperm present from the female before it mates (Waage 1979). Sperm competition adds to the di"culty of obtaining a successful reproductive event by males.
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Conclusions
To transfer their genes to the next generation successfully, animals need to choose a suitable mate. Failure to do so leads to low or no reproductive success that is, poor tness. But reproductive success can also hinge on the number of mates, and on social interactions that extend beyond mating. By classifying social interactions, scientists have been able to identify di!erent types of mating systems, such as monogamy and polygyny. The mating systems described in this article represent a variety of strategies to achieve reproductive success. The diversity of mating systems in animals is a fascinating example of the incredible variety of solutions that a complex evolutionary problem can yield.
Glossary
Bateman's Principle: The theory that females almost always invest more energy into producing o!spring than males, and therefore, in most species, females are a limiting resource over which the other sex will compete. tness: The relative measure of the reproductive success of an individual passing its genes to the next generation. genetic recombination: The process of creating allelic variation in o!spring by exchanging DNA; typically happens during sexual reproduction. harem: A group of females associated with a single male. Typically the male in the harem defends his group of females. lek: A mating system which consists of an aggregation of males where each is seeking to attract a mate. Leks are not associated with resources; however it is thought that leks attract more females than a single male would attract. resource defense polygyny: A mating systems in which males establish a territory around resources needed for mating success. In this system multiple females will join the male in his territory. polyandry: A mating system where one female pairs with many males. polygyny: A mating system where one male is associated with many females. promiscuity: A mating system where there are no pair bonds. In this case is seems that males and females mate randomly. serial monogamy: A mating system in animals where they pair with a mate for one mating season but change mates over the course of a lifetime. social monogamy: The behavioral pairing of a single female with a single male.
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Science
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