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Extra-pair copulation

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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.
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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,
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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
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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
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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
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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
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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.
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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
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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
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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
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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
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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.
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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".
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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.
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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.
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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.
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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
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Another potential explanation for the occurrence of EPC in organisms where females solicit EPC is that the alleles controlling such behaviour are intersexually
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Forstmeier, Wolfgang; Nakagawa, Shinichi; Griffith, Simon C.; Kempenaers, Bart (2014). "Female extra-pair mating: adaptation or genetic constraint?".
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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
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Brekke, Patricia (2013). "Evolution of extreme-mating behaviour: Patterns of extrapair paternity in a species with forced extrapair copulation".
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Gowaty, P. A.; Bridges, W. C. (1991). "Behavioral, demographic, and environmental correlates of extrapair fertilizations in eastern bluebirds".
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Petrie, Marion; Kempenaers, Bart (1998-02-01). "Extra-pair paternity in birds: explaining variation between species and populations".
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CHRISTENSEN, HAROLD T. (1973-01-01). "Attitudes Toward Marital Infidelity: A Nine-Culture Sampling of University Student Opinion".
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Gangestad, Steven W; Thornhill, Randy (1997). "The evolutionary psychology of extrapair sex: The role of fluctuating asymmetry".
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For males, a number of theories are proposed to explain extra-pair copulations. One such hypothesis is that males maximise their
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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:
Birkhead, T. R.; Pellatt, J.; Hunter, F. M. (1988). "Extra-pair copulation and sperm competition in the zebra finch".
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Houtman, A. M. (1992-07-22). "Female Zebra Finches Choose Extra-Pair Copulations with Genetically Attractive Males".
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Birkhead, T.; Parker, G. (1996). "Sperm competition and mating systems". In Krebs, John; Davies, Nicholas (eds.).
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Buss, David M.; Shackelford, Todd K. (1997-06-01). "Susceptibility to Infidelity in the First Year of Marriage".
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Gray, E.M. (1997). "Do female red-winged blackbirds benefit genetically from seeking extra-pair copulations?".
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MΓΈller, A.P. (1988-04-14). "Female choice selects for male sexual tail ornaments in the monogamous swallow".
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Forstmeier, Wolfgang; Martin, Katrin; Bolund, Elisabeth; Schielzeth, Holger; Kempenaers, Bart (2011-06-28).
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Smith, S.M. (1988-01-01). "Extra-Pair Copulations in Black-Capped Chickadees: the Role of the Female".
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Sherman, P.W.; Morton, M.L. (1988). "Extra-pair fertilizations in mountain white-crowned sparrows".
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consider this finding to be support for the 'female choice' hypothesis of mating systems in birds.
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by copulating with as many females as possible outside of a pair bond relationship because their
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Various factors can increase the probability of EPC in males. Firstly, males with low levels of
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Gowaty, P.A. (2006). "Beyond extra-pair paternity". In Lucas, Jeffrey; Simmons, Leigh (eds.).
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Bollinger, E. K.; Gavin, T. A. (1991). "Patterns of extra-pair fertilizations in bobolinks".
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Reichard, U. (1995). "Extra-pair Copulations in a Monogamous Gibbon (Hylobates lar)".
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in males that promote extra-pair copulation as an evolutionary strategy to increase
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Crawford, J.C.; Liu, Z.; Nelson, T.A.; Nielsen, C.K.; Bloomquist, C.K. (2008).
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is shared between sexes leading to this behaviour being expressed in females.
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birds, EPC is only half as common as in socially monogamous birds. Some
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have theorized that extra-pair copulation is an indirect result of
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The 97: 94: 68: 67:In human males 65: 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 2044: 2033: 2030: 2028: 2025: 2023: 2020: 2018: 2015: 2013: 2010: 2009: 2007: 1993: 1989: 1985: 1981: 1977: 1973: 1969: 1965: 1961: 1957: 1950: 1947: 1942: 1938: 1934: 1930: 1923: 1920: 1915: 1911: 1907: 1903: 1899: 1895: 1891: 1887: 1880: 1878: 1874: 1869: 1865: 1861: 1857: 1853: 1849: 1842: 1839: 1834: 1830: 1825: 1820: 1816: 1812: 1808: 1804: 1800: 1793: 1791: 1787: 1776:on 2018-03-05 1772: 1768: 1764: 1760: 1756: 1752: 1748: 1741: 1734: 1731: 1725: 1720: 1716: 1712: 1708: 1701: 1698: 1693: 1689: 1685: 1681: 1677: 1673: 1669: 1665: 1661: 1654: 1651: 1646: 1642: 1638: 1634: 1630: 1626: 1622: 1618: 1611: 1608: 1603: 1599: 1595: 1591: 1587: 1583: 1579: 1575: 1568: 1565: 1560: 1556: 1552: 1548: 1544: 1540: 1536: 1532: 1528: 1524: 1517: 1514: 1509: 1503: 1499: 1492: 1489: 1484: 1478: 1474: 1467: 1464: 1459: 1453: 1449: 1442: 1440: 1436: 1431: 1427: 1422: 1417: 1413: 1409: 1405: 1401: 1397: 1390: 1387: 1375: 1371: 1367: 1363: 1359: 1355: 1351: 1347: 1340: 1337: 1331: 1326: 1322: 1318: 1314: 1307: 1304: 1299: 1295: 1291: 1287: 1283: 1279: 1275: 1271: 1267: 1263: 1255: 1252: 1246: 1241: 1237: 1233: 1229: 1222: 1219: 1214: 1210: 1206: 1202: 1199:(2): 99–112. 1198: 1194: 1187: 1185: 1181: 1176: 1172: 1168: 1164: 1160: 1156: 1152: 1148: 1141: 1138: 1133: 1129: 1125: 1121: 1117: 1113: 1106: 1103: 1098: 1094: 1090: 1086: 1082: 1078: 1071: 1068: 1063: 1059: 1055: 1051: 1044: 1042: 1038: 1033: 1029: 1024: 1019: 1015: 1011: 1007: 1003: 999: 995: 991: 984: 981: 976: 972: 968: 964: 960: 956: 952: 948: 944: 940: 933: 931: 927: 922: 916: 912: 905: 902: 897: 891: 887: 882: 881: 872: 869: 864: 858: 854: 847: 844: 838: 833: 828: 823: 819: 815: 811: 804: 802: 800: 796: 791: 787: 783: 779: 775: 771: 764: 762: 758: 746: 742: 738: 734: 730: 726: 722: 718: 714: 710: 703: 700: 689:on 2016-03-04 685: 681: 675: 668: 667: 659: 656: 645:on 2016-03-04 641: 637: 631: 624: 623: 615: 612: 606: 601: 597: 593: 589: 582: 580: 576: 571: 567: 562: 557: 553: 549: 544: 539: 535: 531: 527: 520: 518: 514: 502: 498: 494: 490: 483: 481: 479: 477: 475: 471: 466: 462: 458: 454: 450: 446: 442: 438: 431: 429: 425: 420: 416: 412: 408: 401: 399: 397: 393: 388: 384: 380: 376: 373:(1324): 3–6. 372: 368: 361: 359: 357: 353: 348: 344: 340: 336: 332: 328: 324: 320: 316: 312: 305: 303: 301: 299: 297: 293: 286: 284: 282: 277: 274: 273:meta-analysis 270: 266: 262: 258: 250: 248: 246: 242: 235: 230: 223: 221: 219: 215: 210: 208: 204: 200: 196: 192: 188: 184: 180: 179:zebra finches 172: 168: 167:Niagara Falls 164: 160: 159:zebra finches 155: 148: 146: 143: 137:Other animals 136: 134: 130: 128: 124: 120: 116: 110: 108: 104: 95: 93: 89: 86: 81: 79: 75: 66: 64: 61: 57: 52: 49: 44: 41:behaviour in 40: 36: 32: 19: 1959: 1955: 1949: 1935:(1): 15–23. 1932: 1928: 1922: 1889: 1885: 1851: 1847: 1841: 1806: 1802: 1778:. Retrieved 1771:the original 1750: 1746: 1733: 1714: 1710: 1700: 1667: 1663: 1653: 1620: 1616: 1610: 1580:(2): 52–58. 1577: 1573: 1567: 1526: 1522: 1516: 1497: 1491: 1472: 1466: 1447: 1403: 1399: 1389: 1378:. Retrieved 1353: 1349: 1339: 1320: 1316: 1306: 1265: 1261: 1254: 1235: 1231: 1221: 1196: 1192: 1150: 1146: 1140: 1115: 1111: 1105: 1080: 1076: 1070: 1053: 1049: 997: 993: 983: 942: 938: 910: 904: 879: 871: 852: 846: 817: 813: 776:(2): 69–88. 773: 769: 749:. Retrieved 716: 712: 702: 691:. Retrieved 684:the original 665: 658: 647:. Retrieved 640:the original 621: 614: 595: 591: 533: 529: 505:. Retrieved 496: 492: 440: 436: 410: 406: 370: 366: 317:(2): 52–58. 314: 310: 278: 254: 238: 211: 206: 198: 176: 163:Bird Kingdom 144: 142:offspring). 140: 131: 111: 99: 90: 82: 70: 53: 34: 30: 29: 2032:Promiscuity 598:(1): 1–49. 281:pleiotropic 218:ethologists 181:, although 48:pair-bonded 2006:Categories 1780:2018-07-21 1380:2019-09-18 751:2016-03-02 693:2016-03-02 649:2016-03-02 507:2016-03-27 443:(1): 1–7. 287:References 224:In mammals 195:copulation 187:monogamous 107:infidelity 43:monogamous 2027:Sexuality 1984:0169-5347 1929:Behaviour 1692:209585885 1684:1474-919X 1000:: 11219. 959:0036-8075 552:0027-8424 339:0169-5347 191:courtship 119:selection 1992:24909948 1914:33212719 1868:53182918 1833:19324647 1767:53197373 1645:17937763 1602:21238200 1430:17456454 1374:Archived 1370:20780520 1298:23091402 1290:15078457 1193:Ethology 1175:44478142 1097:41600842 1032:27044573 975:31391458 911:Homicide 745:Archived 741:33245508 733:11301543 570:21670288 501:Archived 465:25208360 387:83985353 347:21238200 265:swallows 232:Pair of 173:, Canada 157:Pair of 149:In birds 1964:Bibcode 1894:Bibcode 1824:2679913 1625:Bibcode 1582:Bibcode 1559:4355745 1551:3386741 1531:Bibcode 1421:2169281 1270:Bibcode 1201:Bibcode 1155:Bibcode 1132:4087269 1112:The Auk 1023:4832056 1002:Bibcode 967:9767050 939:Science 778:Bibcode 561:3127899 445:Bibcode 319:Bibcode 261:genetic 241:mammals 171:Ontario 123:alleles 37:) is a 2017:Mating 1990:  1982:  1912:  1886:Nature 1866:  1831:  1821:  1765:  1690:  1682:  1643:  1600:  1557:  1549:  1523:Nature 1504:  1479:  1454:  1428:  1418:  1368:  1296:  1288:  1173:  1130:  1095:  1030:  1020:  973:  965:  957:  917:  892:  859:  739:  731:  676:  632:  568:  558:  550:  463:  385:  345:  337:  39:mating 1910:S2CID 1864:S2CID 1774:(PDF) 1763:S2CID 1743:(PDF) 1688:S2CID 1641:S2CID 1555:S2CID 1366:S2CID 1294:S2CID 1171:S2CID 1128:JSTOR 1093:JSTOR 971:S2CID 737:S2CID 687:(PDF) 670:(PDF) 643:(PDF) 626:(PDF) 499:(5). 461:S2CID 383:S2CID 1988:PMID 1980:ISSN 1829:PMID 1680:ISSN 1664:Ibis 1598:PMID 1547:PMID 1502:ISBN 1477:ISBN 1452:ISBN 1426:PMID 1286:PMID 1028:PMID 963:PMID 955:ISSN 915:ISBN 890:ISBN 857:ISBN 729:PMID 674:ISBN 630:ISBN 566:PMID 548:ISSN 343:PMID 335:ISSN 1972:doi 1937:doi 1933:107 1902:doi 1890:332 1856:doi 1819:PMC 1811:doi 1755:doi 1719:doi 1672:doi 1668:162 1633:doi 1590:doi 1539:doi 1527:334 1416:PMC 1408:doi 1404:274 1358:doi 1325:doi 1278:doi 1240:doi 1209:doi 1197:100 1163:doi 1120:doi 1116:104 1085:doi 1058:doi 1018:PMC 1010:doi 947:doi 943:281 886:312 832:hdl 822:doi 786:doi 721:doi 600:doi 556:PMC 538:doi 534:108 453:doi 415:doi 375:doi 371:249 327:doi 212:In 209:). 35:EPC 2008:: 1986:. 1978:. 1970:. 1960:29 1958:. 1931:. 1908:. 1900:. 1888:. 1876:^ 1862:. 1852:53 1850:. 1827:. 1817:. 1805:. 1801:. 1789:^ 1761:. 1751:47 1749:. 1745:. 1715:12 1713:. 1709:. 1686:. 1678:. 1666:. 1662:. 1639:. 1631:. 1621:67 1619:. 1596:. 1588:. 1578:13 1576:. 1553:. 1545:. 1537:. 1525:. 1438:^ 1424:. 1414:. 1402:. 1398:. 1372:. 1364:. 1354:71 1352:. 1348:. 1321:15 1319:. 1315:. 1292:. 1284:. 1276:. 1266:13 1264:. 1236:89 1234:. 1230:. 1207:. 1195:. 1183:^ 1169:. 1161:. 1151:22 1149:. 1126:. 1114:. 1091:. 1079:. 1054:31 1052:. 1040:^ 1026:. 1016:. 1008:. 996:. 992:. 969:. 961:. 953:. 941:. 929:^ 888:. 830:. 816:. 812:. 798:^ 784:. 774:18 772:. 760:^ 743:. 735:. 727:. 717:23 715:. 711:. 596:12 594:. 590:. 578:^ 564:. 554:. 546:. 532:. 528:. 516:^ 495:. 491:. 473:^ 459:. 451:. 441:29 439:. 427:^ 409:. 395:^ 381:. 369:. 355:^ 341:. 333:. 325:. 315:13 313:. 295:^ 169:, 165:, 161:: 80:. 1994:. 1974:: 1966:: 1943:. 1939:: 1916:. 1904:: 1896:: 1870:. 1858:: 1835:. 1813:: 1807:5 1783:. 1757:: 1727:. 1721:: 1694:. 1674:: 1647:. 1635:: 1627:: 1604:. 1592:: 1584:: 1561:. 1541:: 1533:: 1510:. 1485:. 1460:. 1432:. 1410:: 1383:. 1360:: 1333:. 1327:: 1300:. 1280:: 1272:: 1248:. 1242:: 1215:. 1211:: 1203:: 1177:. 1165:: 1157:: 1134:. 1122:: 1099:. 1087:: 1081:4 1064:. 1060:: 1034:. 1012:: 1004:: 998:7 977:. 949:: 923:. 898:. 865:. 840:. 834:: 824:: 818:5 792:. 788:: 780:: 754:. 723:: 696:. 652:. 608:. 602:: 572:. 540:: 510:. 497:9 467:. 455:: 447:: 421:. 417:: 411:2 389:. 377:: 349:. 329:: 321:: 205:( 33:( 20:)

Index

Extra-pair mating
mating
monogamous
pair-bonded
reproductive success
parental investment
gestation period
sexually transmitted infections
fluctuating asymmetry
sexually transmitted infections
infidelity
evolutionary psychologists
selection
alleles
reproductive success

zebra finches
Bird Kingdom
Niagara Falls
Ontario
zebra finches
socially monogamous
monogamous
courtship
copulation
azure-winged magpie
socially polygynous
ethologists

white-handed gibbons

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