88:
male. Psychosocial stress early on in life, including behaviours such as physical violence and substance abuse, can predict EPC in later life. This has been explained as being due to Life
History Theory, which argues that individuals who are reared in environments where resources are scarce and life expectancy is low, are more likely to engage in reproductive behaviours earlier in life in order to ensure the proliferation of their genes. Individuals reared in these environments are said to have short life histories. With respect to Life History Theory, these finding have been explained by suggesting that males who experienced psychosocial stress early in life have short life histories, making them more likely to try and reproduce as much as possible by engaging in EPC to avoid gene extinction.
154:
92:
genes surviving to the next generation. A second reason that EPCs may be avoided by a male is that it can be costly to them; their EPC may be discovered, leading to the dissolution of the long-term relationship with their partner and, in some cases, lead to their partner assaulting or even killing them. Men may also avoid EPCs to minimize the risk of putting themselves at increased opportunity for STD transmission which can be common in EPCs. The partners in the EPC may be promiscuous as well leading to a higher statistical chance and probability of contracting venereal diseases; this would counter the lower incidence of STD transmission among exclusively monogamous sexually active couples.
197:. In a laboratory study, female zebra finches copulated over several days, many times with one male and only once with another male. Results found that significantly more eggs were fertilised by the extra-pair male than expected proportionally from just one copulation versus many copulations with the other male. EPC proportion varies between different species of birds. For example, in eastern bluebirds, studies have shown that around 35% of offspring is due to EPC. Some of the highest levels of EPP are found in the New Zealand hihi/stitchbird (
229:
76:, and then further rearing of the offspring. Contrastingly, men are able to copulate and then abandon their mate as there is no risk of pregnancy for themselves, meaning there is a smaller risk of parental investment in any possible offspring. It has been suggested that, due to having such low parental investment, it is evolutionarily adaptive for men to copulate with as many women as possible. This will allow males to spread their genes with little risk of future investment but it does come with the increased risk of
283:. Under the hypothesis of intersexual antagonistic pleiotropy, the benefit males get from EPC cancels out the negative effects of EPC for females. Thus, the allele that controls EPC in both organisms would persist, even if it would be detrimental to the fitness of females. Similarly, according to the hypothesis of intrasexual antagonistic pleiotropy, the allele that controls EPC in females also controls a behaviour that is under positive selection, such as receptiveness towards within-pair copulation.
247:. A study of one group found 88% in-pair copulation and 12% extra-pair copulation. However, there is much variability in rates of EPC in mammals. One study found that this disparity in EPC is better predicted by the differing social structures of different mammals, rather than differing types of pair bonding. For example, EPC was lower in species who live in pairs compared to those who live in solitary or family structures.
113:
by engaging in extra-pair copulation with better quality males. A second theory is that a woman will engage in extra-pair copulation to seek additional resources for herself or her offspring. This is based on observations from the animal world in which females may copulate outside of their pair-bond relationship with neighbours to gain extra protection, food or nesting materials. Finally,
100:
From an evolutionary perspective, females have to invest a lot more in their offspring than males due to prolonged pregnancy and child rearing, and a child has a better chance of survival and development with two parents involved in child-rearing. Therefore, extra-pair copulations have a greater cost
112:
The most common theory is that women mate outside of the monogamous relationship to acquire better genetic material for their offspring. A female in a relationship with a male with 'poorer genetic quality' may try to enhance the fitness of her children and therefore the continuation of her own genes
62:
is lower, meaning they can copulate and leave the female with minimum risk to themselves. Females, on the other hand, have to invest a lot more in their offspring; extra-pair copulations produce a greater cost because they put the resources that their mate can offer at risk by copulating outside the
45:
species. Monogamy is the practice of having only one sexual partner at any one time, forming a long-term bond and combining efforts to raise offspring together; mating outside this pairing is extra-pair copulation. Across the animal kingdom, extra-pair copulation is common in monogamous species, and
132:
There are also social factors involved in extra-pair copulation. Both males and females have been found to engage in more sexual behaviour outside of the monogamous relationship when experiencing sexual dissatisfaction in the relationship, although how this links to evolutionary theory is unclear.
91:
However, men may also choose not to have EPCs for multiple reasons. One reason may be that long-term monogamous relationships can help form environments that will aid the successful rearing of offspring, as the male is present to help raise them, leading to an increased probability of the male's
87:
are more likely to have EPCs. This may be due to the fact that signals of low fluctuating asymmetry suggest that the males have "good genes", making females more likely to copulate with them as it will enhance the genes of their offspring, even if they do not expect long-term commitment from the
141:
As well as humans, EPC has been found in many other socially monogamous species. When EPC occurs in animals which show sustained female-male social bonding, this can lead to extra-pair paternity (EPP), in which the female reproduces with an extra-pair male, and hence produces EPO (extra-pair
71:
Extra-pair copulation in men has been explained as being partly due to parental investment. Research has suggested that copulation poses more of a risk to future investment for women, as they have the potential of becoming pregnant, and consequently require a large parental investment of the
275:
of genetic benefits of EPC in 55 bird species found that extra-pair offspring were not more likely to survive than within-pair offspring. Also, extra-pair males did not show significantly better 'good-genes' traits than within-pair males, except for being slightly larger overall.
267:, males with longer tails are involved in EPC more than those with shorter tails. Also female swallows with a shorter-tailed within-pair mates are more likely to conduct EPC than those whose mates have longer tails. A similar pattern has been found for
105:, which is suggested as a possible evolutionary reason for the transition from polygamous to monogamous relationships in humans. Despite this, females do seek out extra-pair copulation, with some research finding that women's levels of
50:
species are thought to be exclusively sexually monogamous. EPC in the animal kingdom has mostly been studied in birds and mammals. Possible benefits of EPC can be investigated within non-human species, such as birds.
1259:
Bishop, J.M.; Jarvis, J.U.M.; Spinks, A.C.; Bennett, N.C.; O'Ryan, C. (2004-03-31). "Molecular insight into patterns of colony composition and paternity in the common mole-rat
Cryptomys hottentotus hottentotus".
145:
Due to the obvious reproductive success benefits for males, it used to be thought that males exclusively controlled EPCs. However, it is now known that females also seek EPC in some situations.
744:
133:
Surveys have found cultural differences in attitudes towards infidelity, though it is relatively consistent that female attitudes are less favorable toward infidelity than male attitudes.
109:
are equal to that of men's, although this evidence is mixed. Due to the increased risk, there is more confusion about the evolutionary benefits of extra-pair copulation for females.
271:, in which all extra-pair males had higher rank than the within-pair males. But some argue that genetic benefits for offspring is not the reason females participate in EPC. A
1373:
63:
relationship. Despite this, females do seek out extra pair copulations, and, because of the risk, there is more debate about the evolutionary benefits for females.
101:
for women because they put the support and resources that their mate can offer at risk by copulating outside the relationship. There is also the increased risk of
279:
Another potential explanation for the occurrence of EPC in organisms where females solicit EPC is that the alleles controlling such behaviour are intersexually
153:
1954:
Forstmeier, Wolfgang; Nakagawa, Shinichi; Griffith, Simon C.; Kempenaers, Bart (2014). "Female extra-pair mating: adaptation or genetic constraint?".
263:
benefits which is why the extra-pair males involved in EPC seem to be a non-random subset. There is some evidence for this in birds. For example, in
708:
1615:
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664:
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54:
For males, a number of theories are proposed to explain extra-pair copulations. One such hypothesis is that males maximise their
2011:
102:
77:
1345:
500:
1660:"Effects of extra-pair paternity and maternity on the provisioning strategies of the Azure-winged Magpie Cyanopica cyanus"
1572:
Petrie, M.; Kempenaers, B. (1998). "Extra-pair paternity in birds: explaining variation between species and populations".
201:), in which up to 79% of offspring are sired by EPC. EPC can have significant consequences for parental care, as shown in
1396:"Subordinate male meerkats prospect for extra-group paternity: alternative reproductive tactics in a cooperative mammal"
1521:
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126:
55:
1145:
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244:
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consider this finding to be support for the 'female choice' hypothesis of mating systems in birds.
213:
202:
190:
186:
182:
59:
42:
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1909:
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1313:"Polygynandry in a red fox population: implications for the evolution of group living in canids?"
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885:
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by copulating with as many females as possible outside of a pair bond relationship because their
83:
Various factors can increase the probability of EPC in males. Firstly, males with low levels of
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1979:
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374:
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73:
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1967:
1897:
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1204:
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1005:
781:
448:
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1212:
1022:
989:
560:
525:
1770:
1593:
1191:
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789:
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2005:
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1913:
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in males that promote extra-pair copulation as an evolutionary strategy to increase
1558:
162:
1361:
950:
1799:"Social structure influences extra-pair paternity in socially monogamous mammals"
17:
488:
178:
1975:
1226:
Crawford, J.C.; Liu, Z.; Nelson, T.A.; Nielsen, C.K.; Bloomquist, C.K. (2008).
826:
809:
1723:
1706:
1636:
1244:
1228:"Microsatellite analysis of mating and kinship in beavers (Castor canadensis)"
1227:
724:
604:
587:
280:
129:
is shared between sexes leading to this behaviour being expressed in females.
106:
1983:
1683:
1344:
Kitchen, A.M.; Gese, E.M.; Waits, L.P.; Karki, S.M.; Schauster, E.R. (2006).
990:"Disease dynamics and costly punishment can foster socially imposed monogamy"
958:
551:
526:"Female extrapair mating behavior can evolve via indirect selection on males"
418:
338:
1940:
1329:
1312:
1123:
542:
47:
1991:
1859:
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1601:
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732:
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378:
346:
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264:
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217:
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birds, EPC is only half as common as in socially monogamous birds. Some
1707:"Social mating systems and extrapair fertilizations in passerine birds"
1166:
1131:
456:
240:
170:
1675:
1905:
1542:
122:
38:
1448:
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709:"The evolution of human mating: Trade-offs and strategic pluralism"
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1450:. Burlington, MA: Elsevier Academic Press. pp. 221β254.
1346:"Multiple breeding strategies in the swift fox, Vulpes velox"
482:
480:
478:
476:
474:
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Extra-pair copulation is common in birds. For example,
913:. Aldine de Gruyter (Hawthorne, NY, US). p. 328.
988:Bauch, Chris T.; McElreath, Richard (2016-04-12).
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628:. University of Chicago Press. pp. 795β838.
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851:Kaplan, Hillard S; Gangestad, Steven W (2005).
530:Proceedings of the National Academy of Sciences
360:
358:
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1498:Behavioural ecology: an evolutionary approach
803:
801:
799:
763:
761:
8:
1740:"Extra-pair copulations in a monogamous ape"
581:
579:
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808:Koehler, Nicole; Chisholm, James S (2007).
707:Gangestad, Steven; Simpson, Jeffry (2000).
1822:
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1419:
1328:
1243:
1021:
835:
825:
603:
559:
541:
666:Parental investment and sexual selection
622:Parental investment and sexual selection
255:Some argue that EPC is one way in which
292:
1797:Cohas, A.; Allaine, D. (2009-03-04).
1705:Hasselquist, D.; Sherman, P. (2001).
1077:Journal of Comparative Family Studies
1043:
1041:
932:
930:
7:
1471:Welty, J.C.; Baptista, L.F. (1988).
909:Daly, Martin; Wilson, Margo (1988).
519:
517:
304:
302:
300:
298:
296:
1617:Behavioral Ecology and Sociobiology
1147:Behavioral Ecology and Sociobiology
487:Akçay, E.; Roughgarden, J. (2007).
437:Behavioral Ecology and Sociobiology
1213:10.1111/j.1439-0310.1995.tb00319.x
1050:Journal of Research in Personality
189:and hence do engage in extra-pair
27:Non-monogamy in monogamous species
25:
1956:Trends in Ecology & Evolution
1574:Trends in Ecology & Evolution
311:Trends in Ecology & Evolution
239:EPC has been shown in monogamous
1282:10.1111/j.1365-294X.2004.02131.x
1376:from the original on 2018-07-23
884:. Oxford: WH Freeman. pp.
747:from the original on 2016-03-07
503:from the original on 2016-04-19
103:sexually transmitted infections
78:sexually transmitted infections
1:
1594:10.1016/S0169-5347(97)01232-9
1362:10.1016/j.anbehav.2005.06.015
951:10.1126/science.281.5385.1982
790:10.1016/S1090-5138(97)00003-2
713:Behavioral and Brain Sciences
592:Behavioral and Brain Sciences
493:Evolutionary Ecology Research
331:10.1016/S0169-5347(97)01232-9
770:Evolution and Human Behavior
2048:
1976:10.1016/j.tree.2014.05.005
1738:Palombit, Ryne A. (1994).
827:10.1177/147470490700500111
115:evolutionary psychologists
1637:10.1007/s00265-013-1522-9
1245:10.1644/07-MAMM-A-251R1.1
725:10.1017/S0140525X0000337X
605:10.1017/S0140525X00023992
663:Trivers, Robert (1996).
619:Trivers, Robert (1996).
2022:Reproduction in animals
1941:10.1163/156853988X00160
1724:10.1093/beheco/12.4.457
876:Mellen, Sydney (1981).
814:Evolutionary Psychology
543:10.1073/pnas.1103195108
269:black-capped chickadees
1860:10.1006/anbe.1996.0337
1815:10.1098/rsbl.2008.0760
1759:10.1006/anbe.1994.1097
1412:10.1098/rspb.2007.0316
1062:10.1006/jrpe.1997.2175
586:Buss, David M (1989).
419:10.1093/beheco/2.4.339
379:10.1098/rspb.1992.0075
236:
174:
2012:Developmental biology
1330:10.1093/beheco/arh077
1124:10.1093/auk/104.4.597
994:Nature Communications
880:The Evolution of Love
837:10536/DRO/DU:30062082
251:Reasons for evolution
231:
156:
85:fluctuating asymmetry
31:Extra-pair copulation
1232:Journal of Mammalogy
1089:10.3138/jcfs.4.2.197
234:white-handed gibbons
127:reproductive success
56:reproductive success
1968:2014TEcoE..29..456F
1898:1988Natur.332..640M
1629:2013BEcoS..67..963B
1586:1998TEcoE..13...52P
1535:1988Natur.334...60B
1406:(1618): 1603β1609.
1274:2004MolEc..13.1217B
1205:1995Ethol.100...99R
1159:1988BEcoS..22..413S
1014:10.1038/ncomms11219
1006:2016NatCo...711219B
945:(5385): 1982β1983.
782:1997EHumB..18...69G
536:(26): 10608β10613.
449:1991BEcoS..29....1B
323:1998TEcoE..13...52P
245:white-handed gibbon
214:socially polygynous
203:azure-winged magpie
185:, are not sexually
183:socially monogamous
60:parental investment
1711:Behavioral Ecology
1317:Behavioral Ecology
1167:10.1007/BF00294979
457:10.1007/BF00164288
407:Behavioral Ecology
237:
199:Notiomystis cincta
175:
1892:(6165): 640β642.
1676:10.1111/ibi.12800
1507:978-0-86542-731-0
1482:978-0-03-068923-9
1473:The life of birds
1457:978-0-12-369499-7
1262:Molecular Ecology
679:978-0-226-35456-9
635:978-0-226-35456-9
259:is operating for
18:Extra-pair mating
16:(Redirected from
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1995:
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1906:10.1038/332640a0
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1848:Animal Behaviour
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1769:. Archived from
1747:Animal Behaviour
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1356:(5): 1029β1038.
1350:Animal Behaviour
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682:. Archived from
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257:sexual selection
207:Cyanopica cyanus
193:and attempts at
96:In human females
74:gestation period
46:only a very few
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1529:(6177): 60β62.
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179:zebra finches
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159:zebra finches
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137:Other animals
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218:ethologists
181:, although
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2006:Categories
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751:2016-03-02
693:2016-03-02
649:2016-03-02
507:2016-03-27
443:(1): 1β7.
287:References
224:In mammals
195:copulation
187:monogamous
107:infidelity
43:monogamous
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1984:0169-5347
1929:Behaviour
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1193:Ethology
1175:44478142
1097:41600842
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911:Homicide
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265:swallows
232:Pair of
173:, Canada
157:Pair of
149:In birds
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319:Bibcode
261:genetic
241:mammals
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