404:, two groups of small New World monkeys found in South America. Wild groups usually consist of three to ten individuals, with one reproductively active female, one or more reproductive males, and several nonreproductive helpers that can be either male or female. Cooperative polyandry is not the only mating system found in these primates. Polyandrous, monogamous, and polygynous groups can be found within the same population, and a group can even change mating systems, making it the most flexible mating system of any non-human primate. Unlike most primates who typically give birth to single young, twins are the average litter size for tamarins and marmosets. The entire group participates in raising the offspring, sharing the responsibilities of infant carrying, feeding, and grooming. The presence of nonreproductive helpers appears to be the most important factor in determining which mating system is used, as ecological and environmental variability have not been found to have a significant impact. Goldizen (1987) proposed the hypothesis that monogamy in Callitrichidae should develop only in groups with nonreproductive helpers to help raise the young, and in the absence of these helpers, both polyandrous males and females would have higher reproductive success than those in lone monogamous pairs. Indeed, in studies of
192:
more stable increasing male and female fitness. As opposed to polygyny, where it has been observed that polygynous females that lay a large number of eggs exceeding the optimal clutch size reduces the fitness of the group. Polygynous trios compared to polyandrous trios showed that having more eggs lowered the group fitness. Chao (1997) studies using acorn woodpeckers suggest that other mating systems are practiced at a lower rate when compared to polyandry because it is polyandry that can maximize the fitness of males and females when obtaining optimal clutch size. In polyandrous mating, optimal clutch size is obtained because there is only one female and it becomes more stable when all members remain together.
27:
315:), is one of the few that is polyandrous and exhibits parental care of eggs. The male frogs sit on the egg clutches, maintaining contact with the eggs. Parental care and males genetic contribution were found significantly correlated meaning that male frogs with a higher paternity attended their eggs far more significantly than those of partial paternity for the clutch. Females that mate with more than one male obtain the aid of both and as a result of all three parents providing food and care to the offspring there is an increase in the survival of their young. The amount of help is determined based on mating share and paternity share. (Mating Systems, p 275)
252:
has been seen in three different forms: forced copulation, sexual harassment, and intimidation. Forced copulation is seen in those males that cannot gain access to females for mating. As a result of the lack of access to mates, males are more likely to engage in sexually aggressive behaviors that could result in physical harm and death for females, as has been observed in elephant seals and ducks. Intimidation was seen in social primates, who coerce females into mating, therefore, pushing them to mate with their own aggressor. Males use intimidation as a method of punishment in females that do not mate with them.
169:
149:) store more sperm from gift-giving males suggesting that sperm storage is under female control through cryptic sperm choice. The increase in sperm storage from the gift giving males might allow females to produce "sexy sons" that also give gifts and increase the fitness of offspring. Sperm storage and fertilization success increased with copulation duration, suggesting an advantage in sperm competition.
131:) have shown indirect, genetic benefits, where females have increased offspring survival through multiple mating, showing that practicing polyandry mating results in an increase in offspring viability. In a meta analysis, including 10 different orders of insects, polyandry increased the production of eggs by females specifically in
384:
addition, they pose that there is no relationship between multiple paternity and reproductive success. Multiple male paternity is present in green turtles, but no significant fitness benefits have been found. Observations done on clutches showed that there was no correlation of reproductive success when mating with multiple males.
160:, recent studies have found evidence for the ability of female tortoises to produce clutches of eggs that demonstrate multiple paternity. Predictably, these hatchlings showed an increase in genetic variability compared to those sired by a single male. Potential for multiple paternity within a clutch is primarily a result of
285:
Males that compete for more mating opportunities are the ones that have shown less cooperation in breeding. However, there are other males that in order to gain access to females and mating opportunities practice parental care. Such parental care is mostly seen in polyandrous mating systems; but they
251:
has been observed to be one of the major reasons for why females begin to mate with multiple males to outweigh the cost under different circumstances. The sexual conflict hypothesis suggest that polyandry can occur due to sexual coercion preventing females from obtaining any benefits. Sexual coercion
199:
where females have a group of males close by for mating. Copulations are easily seen in jacanas, facilitating the observation of females copulating continuously with various males. The continuous copulations and the close proximity to mates have allowed the females to fly in within minutes leading to
270:
As a result of multiple male mating, multiple paternities were observed in species as a result of convenience polyandry. Convenience
Polyandry Hypothesis is the assumption that there is a greater cost for females when, refusing male mating attempts than in choice mating. It is the situation in which
164:
across reproductive cycles, since studies have confirmed the presence of multiple males' sperm in the female tortoise reproductive tract simultaneously. As a result of clutches with greater variation in paternal genes and increased sperm competition, females can maximize both the genetic quality and
191:
provide substantial information on breeding systems because they practice monogamy, polyandry, and polygyny. In polyandry the presence of more male breeders in acorn woodpeckers has shown that females reproduce the optimal clutch size and that with paternity sharing between males it is behaviorally
271:
females adjust their mating rate to balance the costs of male harassment. It has been suggested that convenience polyandry would increase when females are weaker than males, decreasing the costs of sexual aggression. Convenience polyandry is seen in several arthropod species, like water striders (
383:
that significantly shows that there are potential fitness gains through polyandry. Instead, it is suggested that the multiple mating is a result of male aggression. Both Wright and Lee and Hayes studies exhibit similar results that show a lack of support of the indirect benefits of polyandry. In
255:
Sexual coercion has many benefits to males allowing them to mate, but their strategies pose high cost for females. This has been observed in vertebrates like green turtles. The green turtles is an example of a species that does not receive any possible benefit from polyandry and only uses it to
240:
Polyandry may also impose costs on females, exposing them to diseases, increased predation risk, time and energy costs, and even physical harm due to sexual harassment. Polyandrous females encounter sexual harassment when courted by males resulting to be costly to females. In order to reduce the
318:
In polyandrous mating it has been suggested that male cooperation may outweigh the costs of sharing paternity in situations of scarcity, of foods and of high competition levels for land or females. Female size and a large breeding territory defended by more males may force male cooperation. For
266:
females can store sperm for at least two egg batches without their fertilization rate decreasing. Therefore, mating before each oviposition is not necessary. There is no significant overall benefit for females in multiple matings; large females are at an advantage due to their ability to resist
362:
Infanticide avoidance is one of the major reasons animals, like mammals, are inclined to choose polyandry. Infanticide is observed to be the major cause of mortality in various mammals. Polyandry is thought to evolve from their offspring's vulnerability to infanticide. There is evidence that
327:) has shown genetic evidence for paternity assurance. The comb-crested jacana females had one to three more mates available for copulation. In order to guarantee paternity to their male mates, the females would spend time in the territory of the male, laying her eggs.
247:, like in many other species, the high cost of resisting mating may exceed the cost of accepting numerous males in a breeding if copulating takes a short time, and therefore females in some species are inclined to being polyandrous. In the polyandrous system,
1696:
Huchard, E., Canale, C. I., Le Gros, C., Perret, M., Henry, P.-Y., & Kappeler, P. M. (2012). Convenience polyandry or convenience polygyny? Costly sex under female control in a promiscuous primate. Proceedings: Biological
Sciences, 279(1732),
124:
The adaptive significance of polyandry in animals is controversial. Polyandry has direct benefits for females allowing fertilization assurance, provision of resources, and parental care for their offspring. House mice
256:
reduce the cost of mating. As for house mice, multiple male mating was observed even when females had the opportunity to select their mate without sexual coercion, showing that it was due to female choice. In the fly
999:
Alcock, John; Eickwort, George C.; Eickwort, Kathleen R. (1977). "The reproductive behavior of
Anthidium maculosum (Hymenoptera: Megachilidae) and the evolutionary significance of multiple copulations by females".
214:) supported the hypothesis that sperm mixing indeed occurs in polyandrous social insects. It was further found that the eggs in the queen ants showed to be completely mixed and used randomly during egg laying.
757:
Davy, C.; Edwards, T.; Lathrop, A.; Bratton, M.; Hagan, M.; Henen, B.; Nagy, K.; Stone, J.; Hillard, L.; Murphy, R. (2011). "Polyandry and multiple paternities in the threatened
Agassiz's desert tortoise,
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the expected sperm mixing and allowing the most viable sperm to fertilize most of the clutch or to increase genetic diversity for benefits in disease resistance. Snow and
Andrade (2005) concluded that the
379:
High levels of multiple paternity have widely been seen and reported in snakes, marine turtles, and lizards. However, Lee and Hayes (2004) have evidence that suggests that there is no set data in
298:). Observations that show male relatedness suggest the importance for male social relationships in taking care of offspring. These social relationships in males are seen in primates,
221:, as a form of adapting to multiple mating in females, human penile shape is indicative of an evolutionary history of polyandry. Male humans evolved to have a wedge- or spoon-shaped
951:
165:
number of offspring. Multiple paternities within a single clutch is therefore considered an effective strategy to increase the reproductive success and fitness of female tortoises.
112:
It is theorized that polyandry is more prevalent in organisms where incompatibility is more costly, and where this incompatibility is more likely. The former is especially true in
54:
where one female mates with several males in a breeding season. Polyandry is often compared to the polygyny system based on the cost and benefits incurred by members of each sex.
68:) of the invertebrate order Orthoptera (containing crickets, grasshoppers, and groundhoppers). Polyandrous behavior is also prominent in many other insect species, including the
208:) increases their ability to manipulate the paternity of their offspring by using the spermathecae to store multiple sperms. Additionally, a study on leaf-cutting ants (
656:
Simmons, L. W. (2005). THE EVOLUTION OF POLYANDRY: Sperm
Competition, Sperm Selection, and Offspring Viability. Annual Review of Ecology, Evolution, and Systematics.
471:
Colegrave, Nick; Kotiaho, Janne S.; Tomkins, Joseph L. (2002). "Mate choice or polyandry: reconciling genetic compatibility and good genes sexual selection".
1385:
Díaz-Muñoz, S. L. (2011). "Paternity and relatedness in a polyandrous nonhuman primate: testing adaptive hypotheses of male reproductive cooperation".
371:) has supported the infanticide avoidance hypothesis where polyandrous females' offspring have a higher survival than monandrous females' offspring.
1993:
872:"Emlen, S. T., Wrege, P. H., & Webster, M. S. (1998). Cuckoldry as a cost of polyandry in the sex-role-reversed wattled jacana, Jacana jacana"
721:
Johnston, E.; Rand, M.; Zweifel, S. (2006). "Detection of multiple paternity and sperm storage in a captive colony of the central Asian tortoise,
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1298:
Rowe, L (1992). "Convenience polyandry in a water strider: foraging conflicts and female control of copulation frequency and guarding duration".
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polyandry is an effective strategy that is adapted by many to increase their offspring survival through the avoidance of male infanticide.
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2284:
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males, such factors can promote an increase in reproductive success, leaving behind the cost of paternity share (Mating
Systems, p277).
58:
is where one male mates with several females in a breeding season (e.g., lions, deer, some primates, and many systems where there is an
1577:
1636:"'How Attractive Is the Girl Next Door? An Assessment of Spatial Mate Acquisition and Paternity in the Solitary Cape Dune Mole-Rat,
1425:"Genetic relationship between offspring and guardian adults of a rhacophorid frog and its care effort in response to paternal share"
2038:
950:
Gordon, G. Gallup Jr.; Burch, Rebecca L.; Zappieri, Mary L.; Parvez, Rizwan A.; Stockwell, Malinda L.; Davis, Jennifer A. (2003).
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organisms. Where the cost of having a low-quality father is significant, however, an organism is less likely to be polyandrous.
1986:
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must go through the cost of sharing paternity and parental care. This has been observed in avians like acorn woodpeckers (
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2518:
2630:
2824:
2028:
1979:
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2488:
2483:
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Arnqvist, G.; Nilsson, T. (2000). "The evolution of polyandry: multiple mating and female fitness in insects".
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248:
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2714:
139:. Indirect benefits of mating for females can be gained through sperm competition to attain "good genes",
2603:
2533:
2193:
1517:
Jarvis, Jennifer (May 1981). "Eusociality in a Mammal: Cooperative
Breeding in Naked Mole-Rat Colonies".
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Current research suggests that polyandry is the dominant social structure in the New World monkey family
26:
2793:
1219:"Repeated copulations as a strategy to maximize fertilization in the fly, Dryomyza anilis (Dryomyzidae)"
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Many reptile species also demonstrate polyandry, especially among members of the tortoise family (
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2010:
2002:
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Wright, L. I.; Fuller, W. J.; Godley, B. J.; McGowan, A.; Tregenza, T.; Broderick, A. C. (2013).
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Zeh, Jeanne A.; Zeh, David W. (2001). "Reproductive mode and the genetic benefits of polyandry".
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Evolutionary anthropology of the human family; In C. A. Salmon and T. K. Shackelford (Eds.),
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184:, which often come from various hives. She mothers most or all offspring within a given hive.
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Kohda, M.; Heg, D.; Makino, Y.; Takeyama, T.; Shibata, J.; Watanabe, K.; Awata, S. (2009).
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costs to the females, females will take part in polyandry. For example, in the bee species
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8:
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1335:"Living on the wedge: female control of paternity in a cooperatively polyandrous cichlid"
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Otronen, Merja (1989-01-01). "Female Mating
Behaviour and Multiple Matings in the Fly,
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342:) or caste produces offspring while the other organisms (e.g., non-reproductive female
55:
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Wolff, J. O.; Macdonald, D. W. (2004). "Promiscuous females protect their offspring".
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1914:"Facultative polyandry and the role of infant-carrying in wild saddle-back tamarins (
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This article is about polyandry in non-human species. For polyandry in humans, see
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Proceedings of the
National Academy of Sciences of the United States of America
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346:) cooperate in caring for the young. Examples of mammalian eusociality include
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31:
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1451:
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Thonhauser, K. E.; Raveh, S.; Hettyey, A.; Beissmann, H.; Penn, D. J. (2013).
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306:. High proportions of related males at the half sibling or higher were found.
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2119:
2087:
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1281:
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484:
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62:). A common example of polyandrous mating can be found in the field cricket (
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1234:
1184:
827:
806:
599:
582:
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412:, no monogamous lone pairs have ever been seen to attempt a breeding cycle.
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2109:
1850:"On the mating system of the cooperatively breeding saddle-backed tamarin (
1826:
1777:
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1503:
1478:"Parentage and Relatedness in Polyandrous Comb-Crested Jacanas Using ISSRs"
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354:, among whom polyandry is the norm and polygyny has never been observed.
335:
222:
181:
132:
126:
2495:
1576:
Burda, H. Honeycutt; Begall, S.; Locker-Grutjen, O; Scharff, A. (2000).
856:"Chao, L. (1997). Evolution of polyandry in a communal breeding system.
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1944:
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343:
136:
113:
88:
36:
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143:, increased genetic quality, and genetic diversity. Females spiders (
20:
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2555:
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738:
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during copulation in order to draw foreign semen back away from the
671:"Sperm storage mediated by cryptic female choice for nuptial gifts"
397:
101:
83:
1744:"Polyandry enhances offspring survival in an infanticidal species"
889:
Stürup, M.; Nash, D. R.; Hughes, W. O. H.; Boomsma, J. J. (2014).
2677:
2415:
2208:
401:
291:
177:
97:
1575:
807:"Multiple paternity and breeding system in the gopher tortoise,
583:"Polyandry, sperm competition, and reproductive success in mice"
2235:
1091:
230:
1633:
374:
331:
119:
1578:"Are naked and common mole-rats eusocial and if so, why?"
1423:
Chen, Y.-H.; Cheng, W.-C.; Yu, H.-T.; Kam, Y.-C. (2011).
949:
756:
547:
1793:"Multiple paternity in reptiles: patterns and processes"
1634:
Bray, TC; Bloomer, P; O'Riain, MJ; Bennett, NC (2012).
888:
874:
Proceedings of the Royal Society B: Biological Sciences
675:
Proceedings of the Royal Society B: Biological Sciences
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92:
and bandicoots, around 1% of all bird species, such as
1332:
998:
804:
466:
464:
357:
805:
Moon, J.; McCoy, E.; Mushinsky, H.; Karl, S. (2006).
456:
The Oxford Handbook of Evolutionary Family Psychology
1047:
720:
716:
714:
461:
891:"Sperm mixing in the polyandrous leaf-cutting ant
550:"No benefits of polyandry to female green turtles"
1087:
1085:
1083:
82:. Polyandry also occurs in some primates such as
2806:
1847:
952:"The human penis as a semen displacement device"
711:
262:, females favor matings with quick oviposition.
752:
750:
748:
613:
1706:
1422:
1094:"Why do female mice mate with multiple males?"
1080:
1043:
1041:
1039:
100:, insects such as honeybees, and fish such as
1987:
1848:Terborgh, John; Goldizen, Ann Wilson (1985).
1048:Clutton- Brock, T. H.; Parker, G. A. (1995).
580:
1741:
868:
866:
745:
668:
543:
541:
539:
537:
35: – Palo Verde National Park,
2001:
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1036:
884:
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852:
850:
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846:
16:Class of mating system in non-human species
1994:
1980:
1384:
1149:
1147:
1145:
664:
662:
375:Polyandry benefit exception: green turtles
338:, meaning that a single female (e.g., the
107:
1952:
1888:
1816:
1767:
1673:
1663:
1493:
1358:
1193:
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1125:
926:
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826:
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565:
534:
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1911:
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1841:
1466:
1413:
1375:
1288:
879:
843:
669:Albo, M. J.; Bilde, T.; Uhl, G. (2013).
176:is usually the only female bee within a
167:
120:Benefits and costs of mating for females
25:
1255:
1216:
1153:
1142:
659:
309:As for frog species, rhacaphorid frog (
2807:
1516:
581:Firman, R. C.; Simmons, L. W. (2008).
497:
86:, mammal groups, the marsupial genus'
1975:
1050:"Sexual coercion in animal societies"
1475:
1297:
458:. New York: Oxford University Press.
280:
2519:Sexual selection in scaled reptiles
1925:Behavioral Ecology and Sociobiology
1861:Behavioral Ecology and Sociobiology
1582:Behavioral Ecology and Sociobiology
1432:Behavioral Ecology and Sociobiology
1098:Behavioral Ecology and Sociobiology
1002:Behavioral Ecology and Sociobiology
358:Polyandry and infanticide avoidance
13:
334:with high social organization are
14:
2846:
1709:Trends in Ecology & Evolution
387:
1818:10.1111/j.1365-294x.2008.03772.x
1339:Proceedings: Biological Sciences
195:Multiple mating is also seen in
1784:
1742:Klemme, I.; Ylönen, H. (2010).
1735:
1700:
1690:
1627:
1569:
1510:
1326:
1249:
1210:
992:
943:
798:
1912:Goldizen, Anne Wilson (1987).
1791:Uller, T.; Olsson, M. (2008).
1160:"Polyandry in a marine turtle"
650:
607:
574:
448:
396:. The Callitrichidae includes
1:
1399:10.1016/j.anbehav.2011.06.013
1217:Otronen, Meija (1994-03-01).
979:10.1016/S1090-5138(03)00016-3
473:Evolutionary Ecology Research
442:
1665:10.1371/journal.pone.0039866
1312:10.1016/0003-3472(92)90025-5
959:Evolution and Human Behavior
76:, and the species of spider
7:
2194:semelparity and iteroparity
727:Canadian Journal of Zoology
415:
10:
2851:
1721:10.1016/j.tree.2003.12.009
296:Julidochromis transcriptus
227:repeated thrusting motions
18:
2594:
2554:
2509:
2474:
2431:
2298:
2285:Non-reproductive behavior
2009:
1452:10.1007/s00265-011-1243-x
1118:10.1007/s00265-013-1604-8
784:10.1007/s10592-011-0232-y
156:). Through polyandry and
2825:Reproduction in animals
2003:Animal sexual behaviour
1539:10.1126/science.7209555
1274:10.1163/156853989X00592
1185:10.1073/pnas.0307982101
815:The Journal of Heredity
288:Melanerpes formicivorus
158:long-term sperm storage
108:Predictors of polyandry
2152:traumatic insemination
1760:10.1098/rsbl.2009.0500
1351:10.1098/rspb.2009.1175
1066:10.1006/anbe.1995.0166
687:10.1098/rspb.2013.1735
628:10.1006/anbe.2000.1446
512:10.1006/anbe.2000.1705
267:unnecessary matings.
185:
39:
2280:Interspecies breeding
1602:10.1007/s002650050669
1495:10.1093/jhered/esg072
1235:10.1093/beheco/5.1.51
899:Ecology and Evolution
893:Acromyrmex echinatior
876:265(1413), 2359–2364.
828:10.1093/jhered/esj017
764:Conservation Genetics
600:10.1093/beheco/arm158
567:10.1093/beheco/art003
325:Irediparra gallinacea
290:), and fishes as the
211:Acromyrmex echinatior
171:
141:cryptic female choice
128:Mus musculus musculus
29:
2794:Short-beaked echidna
2534:side-blotched lizard
2051:sexual ornamentation
1916:Saguinus fuscicollis
1852:Saguinus fuscicollis
1476:Haig, S. M. (2003).
406:Saguinus fuscicollis
348:Damaraland mole-rats
312:Kurixalus eiffingeri
206:Latrodectus hasselti
79:Stegodyphus lineatus
2621:Homosexual behavior
2586:Homosexual behavior
2441:Spawning strategies
2241:Bateman's principle
2061:sexy son hypothesis
2039:hormonal motivation
2034:reproductive system
2024:Sexual reproduction
1937:1987BEcoS..20...99G
1873:1985BEcoS..16..293T
1809:2008MolEc..17.2566U
1656:2012PLoSO...739866B
1594:2000BEcoS..47..293B
1531:1981Sci...212..571J
1482:Journal of Heredity
1444:2011BEcoS..65.2329C
1176:2004PNAS..101.6530L
1110:2013BEcoS..67.1961T
1014:1977BEcoS...2..385A
971:2003EHumB..24..277G
911:2014EcoEv...4.3571S
809:Gopherus polyphemus
776:2011ConG...12.1313D
723:Testudo horsfieldii
410:saddle-back tamarin
321:Comb-crested jacana
300:Geoffroy's tamarins
244:Anthidium maculosum
65:Gryllus bimaculatus
2569:Breeding behaviour
2290:Fisher's principle
2115:sexual intercourse
2056:handicap principle
1945:10.1007/BF00572631
1881:10.1007/BF00295541
1638:Bathyergus suillus
1223:Behavioral Ecology
1022:10.1007/bf00299507
858:Behavioral Ecology
760:Gopherus agassizii
681:(1772): 20131735.
587:Behavioral Ecology
554:Behavioral Ecology
422:Green-veined white
304:Saguinus geoffroyi
235:compete with sperm
186:
74:adzuki bean weevil
44:behavioral ecology
40:
2802:
2801:
2616:Lordosis behavior
2496:Frog reproduction
2446:Polyandry in fish
2226:Sexual dimorphism
2147:sperm competition
2066:Fisherian runaway
2046:Courtship display
1797:Molecular Ecology
1525:(4494): 571–573.
1438:(12): 2329–2339.
1345:(1676): 4207–14.
1170:(17): 6530–6535.
1104:(12): 1961–1970.
919:10.1002/ece3.1176
427:Polyandry in fish
281:Paternity sharing
189:Acorn woodpeckers
180:reproducing with
146:Pisaura mirabilis
2842:
2815:Animal sexuality
2772:ringtailed lemur
2631:African wild dog
2604:Sexual selection
2564:Sexual selection
2484:Sexual selection
2125:pseudocopulation
2019:Sexual selection
1996:
1989:
1982:
1973:
1972:
1967:
1966:
1956:
1922:
1909:
1903:
1902:
1892:
1858:
1845:
1839:
1838:
1820:
1788:
1782:
1781:
1771:
1739:
1733:
1732:
1704:
1698:
1694:
1688:
1687:
1677:
1667:
1631:
1625:
1624:
1622:
1621:
1612:. Archived from
1573:
1567:
1566:
1514:
1508:
1507:
1497:
1473:
1464:
1463:
1429:
1420:
1411:
1410:
1387:Animal Behaviour
1382:
1373:
1372:
1362:
1330:
1324:
1323:
1300:Animal Behaviour
1295:
1286:
1285:
1253:
1247:
1246:
1214:
1208:
1207:
1197:
1187:
1151:
1140:
1139:
1129:
1089:
1078:
1077:
1060:(5): 1345–1365.
1054:Animal Behaviour
1045:
1034:
1033:
996:
990:
989:
987:
981:. Archived from
956:
947:
941:
940:
930:
886:
877:
870:
861:
860:, 8(6), 668–674.
854:
841:
840:
830:
802:
796:
795:
770:(5): 1313–1322.
754:
743:
742:
718:
709:
708:
698:
666:
657:
654:
648:
647:
616:Animal Behaviour
611:
605:
604:
602:
578:
572:
571:
569:
560:(4): 1022–1029.
545:
532:
531:
506:(6): 1051–1063.
500:Animal Behaviour
495:
489:
488:
468:
459:
452:
437:Bibron's toadlet
432:Superfecundation
369:Myodes glareolus
237:of other males.
219:Gordon G. Gallup
70:red flour beetle
2850:
2849:
2845:
2844:
2843:
2841:
2840:
2839:
2805:
2804:
2803:
2798:
2777:sexual swelling
2673:European badger
2590:
2550:
2505:
2470:
2427:
2294:
2263:Sexual conflict
2204:hermaphroditism
2005:
2000:
1970:
1920:
1910:
1906:
1856:
1846:
1842:
1803:(11): 2566–80.
1789:
1785:
1748:Biology Letters
1740:
1736:
1705:
1701:
1695:
1691:
1632:
1628:
1619:
1617:
1574:
1570:
1515:
1511:
1474:
1467:
1427:
1421:
1414:
1383:
1376:
1331:
1327:
1296:
1289:
1258:Dryomyza Anilis
1254:
1250:
1215:
1211:
1152:
1143:
1090:
1081:
1046:
1037:
997:
993:
985:
954:
948:
944:
905:(18): 3571–82.
887:
880:
871:
864:
855:
844:
803:
799:
755:
746:
739:10.1139/z06-023
719:
712:
667:
660:
655:
651:
612:
608:
579:
575:
546:
535:
496:
492:
469:
462:
453:
449:
445:
418:
390:
377:
360:
352:naked mole-rats
283:
259:Dryomyza anilis
249:sexual coercion
225:and to perform
197:wattled jacanas
122:
110:
24:
17:
12:
11:
5:
2848:
2838:
2837:
2832:
2830:Mating systems
2827:
2822:
2817:
2800:
2799:
2797:
2796:
2791:
2786:
2781:
2780:
2779:
2774:
2769:
2764:
2759:
2754:
2749:
2741:
2740:
2739:
2729:
2728:
2727:
2717:
2712:
2707:
2702:
2701:
2700:
2695:
2690:
2685:
2675:
2670:
2665:
2660:
2659:
2658:
2653:
2648:
2643:
2638:
2633:
2623:
2618:
2613:
2612:
2611:
2600:
2598:
2592:
2591:
2589:
2588:
2583:
2582:
2581:
2576:
2566:
2560:
2558:
2552:
2551:
2549:
2548:
2543:
2538:
2537:
2536:
2531:
2526:
2515:
2513:
2507:
2506:
2504:
2503:
2498:
2493:
2492:
2491:
2480:
2478:
2472:
2471:
2469:
2468:
2463:
2458:
2453:
2448:
2443:
2437:
2435:
2429:
2428:
2426:
2425:
2424:
2423:
2418:
2413:
2405:
2400:
2399:
2398:
2393:
2383:
2378:
2377:
2376:
2371:
2366:
2356:
2355:
2354:
2344:
2343:
2342:
2341:
2340:
2330:
2325:
2320:
2315:
2304:
2302:
2296:
2295:
2293:
2292:
2287:
2282:
2277:
2276:
2275:
2270:
2260:
2259:
2258:
2253:
2248:
2243:
2238:
2233:
2223:
2222:
2221:
2216:
2211:
2206:
2201:
2196:
2191:
2186:
2181:
2176:
2171:
2161:
2160:
2159:
2154:
2149:
2144:
2139:
2129:
2128:
2127:
2122:
2117:
2112:
2102:
2101:
2100:
2095:
2090:
2085:
2080:
2073:Mating systems
2070:
2069:
2068:
2063:
2058:
2053:
2043:
2042:
2041:
2036:
2031:
2021:
2015:
2013:
2007:
2006:
1999:
1998:
1991:
1984:
1976:
1969:
1968:
1904:
1867:(4): 293–299.
1840:
1783:
1734:
1699:
1689:
1626:
1588:(5): 293–303.
1568:
1509:
1488:(4): 302–309.
1465:
1412:
1393:(3): 563–571.
1374:
1325:
1306:(2): 189–202.
1287:
1268:(1–4): 77–97.
1248:
1209:
1158:(April 2004).
1141:
1079:
1035:
1008:(4): 385–396.
991:
988:on 2011-01-24.
965:(4): 277–289.
942:
878:
862:
842:
821:(2): 150–157.
797:
744:
733:(4): 520–526.
710:
658:
649:
622:(2): 145–164.
606:
593:(4): 695–702.
573:
533:
490:
479:(6): 911–917.
460:
446:
444:
441:
440:
439:
434:
429:
424:
417:
414:
408:, common name
394:Callitrichidae
389:
388:Callitrichidae
386:
376:
373:
359:
356:
282:
279:
202:redback spider
121:
118:
109:
106:
50:is a class of
32:Jacana spinosa
15:
9:
6:
4:
3:
2:
2847:
2836:
2833:
2831:
2828:
2826:
2823:
2821:
2818:
2816:
2813:
2812:
2810:
2795:
2792:
2790:
2787:
2785:
2782:
2778:
2775:
2773:
2770:
2768:
2765:
2763:
2760:
2758:
2755:
2753:
2750:
2748:
2745:
2744:
2742:
2738:
2735:
2734:
2733:
2730:
2726:
2723:
2722:
2721:
2718:
2716:
2715:Spotted hyena
2713:
2711:
2708:
2706:
2703:
2699:
2696:
2694:
2691:
2689:
2686:
2684:
2681:
2680:
2679:
2676:
2674:
2671:
2669:
2666:
2664:
2661:
2657:
2654:
2652:
2649:
2647:
2644:
2642:
2639:
2637:
2634:
2632:
2629:
2628:
2627:
2624:
2622:
2619:
2617:
2614:
2610:
2607:
2606:
2605:
2602:
2601:
2599:
2597:
2593:
2587:
2584:
2580:
2577:
2575:
2572:
2571:
2570:
2567:
2565:
2562:
2561:
2559:
2557:
2553:
2547:
2544:
2542:
2539:
2535:
2532:
2530:
2527:
2525:
2522:
2521:
2520:
2517:
2516:
2514:
2512:
2508:
2502:
2499:
2497:
2494:
2490:
2487:
2486:
2485:
2482:
2481:
2479:
2477:
2473:
2467:
2464:
2462:
2459:
2457:
2454:
2452:
2449:
2447:
2444:
2442:
2439:
2438:
2436:
2434:
2430:
2422:
2421:penis fencing
2419:
2417:
2414:
2412:
2409:
2408:
2406:
2404:
2401:
2397:
2394:
2392:
2391:apophallation
2389:
2388:
2387:
2384:
2382:
2379:
2375:
2372:
2370:
2367:
2365:
2362:
2361:
2360:
2357:
2353:
2350:
2349:
2348:
2345:
2339:
2336:
2335:
2334:
2331:
2329:
2326:
2324:
2321:
2319:
2316:
2314:
2311:
2310:
2309:
2306:
2305:
2303:
2301:
2300:Invertebrates
2297:
2291:
2288:
2286:
2283:
2281:
2278:
2274:
2271:
2269:
2266:
2265:
2264:
2261:
2257:
2254:
2252:
2249:
2247:
2244:
2242:
2239:
2237:
2234:
2232:
2229:
2228:
2227:
2224:
2220:
2217:
2215:
2212:
2210:
2207:
2205:
2202:
2200:
2199:opportunistic
2197:
2195:
2192:
2190:
2187:
2185:
2182:
2180:
2177:
2175:
2172:
2170:
2167:
2166:
2165:
2162:
2158:
2157:penile spines
2155:
2153:
2150:
2148:
2145:
2143:
2140:
2138:
2135:
2134:
2133:
2132:Fertilisation
2130:
2126:
2123:
2121:
2120:Pelvic thrust
2118:
2116:
2113:
2111:
2108:
2107:
2106:
2103:
2099:
2096:
2094:
2091:
2089:
2088:mate guarding
2086:
2084:
2081:
2079:
2076:
2075:
2074:
2071:
2067:
2064:
2062:
2059:
2057:
2054:
2052:
2049:
2048:
2047:
2044:
2040:
2037:
2035:
2032:
2030:
2027:
2026:
2025:
2022:
2020:
2017:
2016:
2014:
2012:
2008:
2004:
1997:
1992:
1990:
1985:
1983:
1978:
1977:
1974:
1964:
1960:
1955:
1954:2027.42/46876
1950:
1946:
1942:
1938:
1934:
1931:(2): 99–109.
1930:
1926:
1919:
1917:
1908:
1900:
1896:
1891:
1890:2027.42/46874
1886:
1882:
1878:
1874:
1870:
1866:
1862:
1855:
1853:
1844:
1836:
1832:
1828:
1824:
1819:
1814:
1810:
1806:
1802:
1798:
1794:
1787:
1779:
1775:
1770:
1765:
1761:
1757:
1753:
1749:
1745:
1738:
1730:
1726:
1722:
1718:
1715:(3): 127–34.
1714:
1710:
1703:
1693:
1685:
1681:
1676:
1671:
1666:
1661:
1657:
1653:
1650:(6): e39866.
1649:
1645:
1641:
1639:
1630:
1616:on 2016-03-04
1615:
1611:
1607:
1603:
1599:
1595:
1591:
1587:
1583:
1579:
1572:
1564:
1560:
1556:
1552:
1548:
1544:
1540:
1536:
1532:
1528:
1524:
1520:
1513:
1505:
1501:
1496:
1491:
1487:
1483:
1479:
1472:
1470:
1461:
1457:
1453:
1449:
1445:
1441:
1437:
1433:
1426:
1419:
1417:
1408:
1404:
1400:
1396:
1392:
1388:
1381:
1379:
1370:
1366:
1361:
1356:
1352:
1348:
1344:
1340:
1336:
1329:
1321:
1317:
1313:
1309:
1305:
1301:
1294:
1292:
1283:
1279:
1275:
1271:
1267:
1263:
1259:
1252:
1244:
1240:
1236:
1232:
1228:
1224:
1220:
1213:
1205:
1201:
1196:
1191:
1186:
1181:
1177:
1173:
1169:
1165:
1161:
1157:
1150:
1148:
1146:
1137:
1133:
1128:
1123:
1119:
1115:
1111:
1107:
1103:
1099:
1095:
1088:
1086:
1084:
1075:
1071:
1067:
1063:
1059:
1055:
1051:
1044:
1042:
1040:
1031:
1027:
1023:
1019:
1015:
1011:
1007:
1003:
995:
984:
980:
976:
972:
968:
964:
960:
953:
946:
938:
934:
929:
924:
920:
916:
912:
908:
904:
900:
896:
894:
885:
883:
875:
869:
867:
859:
853:
851:
849:
847:
838:
834:
829:
824:
820:
816:
812:
810:
801:
793:
789:
785:
781:
777:
773:
769:
765:
761:
753:
751:
749:
740:
736:
732:
728:
724:
717:
715:
706:
702:
697:
692:
688:
684:
680:
676:
672:
665:
663:
653:
645:
641:
637:
633:
629:
625:
621:
617:
610:
601:
596:
592:
588:
584:
577:
568:
563:
559:
555:
551:
544:
542:
540:
538:
529:
525:
521:
517:
513:
509:
505:
501:
494:
486:
482:
478:
474:
467:
465:
457:
451:
447:
438:
435:
433:
430:
428:
425:
423:
420:
419:
413:
411:
407:
403:
399:
395:
385:
382:
381:green turtles
372:
370:
366:
355:
353:
349:
345:
341:
337:
333:
328:
326:
322:
316:
314:
313:
307:
305:
301:
297:
293:
289:
278:
276:
275:
274:Gerris buenoi
268:
265:
261:
260:
253:
250:
246:
245:
238:
236:
232:
228:
224:
220:
217:According to
215:
213:
212:
207:
203:
198:
193:
190:
183:
179:
175:
170:
166:
163:
162:sperm storage
159:
155:
150:
148:
147:
142:
138:
134:
133:Lepidopterans
130:
129:
117:
115:
105:
103:
99:
95:
91:
90:
85:
81:
80:
75:
71:
67:
66:
61:
57:
53:
52:mating system
49:
45:
38:
34:
33:
28:
22:
2762:olive baboon
2710:Hippopotamus
2698:domestic cat
2646:domestic dog
2574:golden eagle
2541:Crocodilians
2189:polygynandry
2178:
2110:cloacal kiss
1928:
1924:
1915:
1907:
1864:
1860:
1851:
1843:
1800:
1796:
1786:
1751:
1747:
1737:
1712:
1708:
1702:
1692:
1647:
1643:
1637:
1629:
1618:. Retrieved
1614:the original
1585:
1581:
1571:
1522:
1518:
1512:
1485:
1481:
1435:
1431:
1390:
1386:
1342:
1338:
1328:
1303:
1299:
1265:
1261:
1257:
1251:
1229:(1): 51–56.
1226:
1222:
1212:
1167:
1163:
1101:
1097:
1057:
1053:
1005:
1001:
994:
983:the original
962:
958:
945:
902:
898:
892:
873:
857:
818:
814:
808:
800:
767:
763:
759:
730:
726:
722:
678:
674:
652:
619:
615:
609:
590:
586:
576:
557:
553:
503:
499:
493:
476:
472:
455:
450:
405:
391:
378:
368:
361:
329:
324:
317:
310:
308:
303:
295:
287:
284:
272:
269:
263:
257:
254:
242:
239:
233:and thus to
216:
209:
205:
194:
187:
154:Testudinidae
151:
144:
137:Orthopterans
127:
123:
111:
87:
77:
63:
47:
41:
30:
2501:Salamanders
2381:Echinoderms
2364:sea anemone
2347:Cephalopods
2251:cannibalism
2174:promiscuity
2093:mating plug
2083:mating call
2078:mate choice
1754:(1): 24–6.
344:worker bees
2809:Categories
2476:Amphibians
2456:Salmon run
2386:Gastropods
2308:Arthropods
2273:intralocus
2268:interlocus
2246:bimaturism
2105:Copulation
2098:lek mating
1620:2014-11-21
443:References
365:Bank voles
114:viviparous
89:Antechinus
60:alpha male
37:Costa Rica
2835:Polyandry
2743:Primates
2732:Pinnipeds
2720:Marsupial
2651:gray wolf
2411:earthworm
2396:love dart
2369:jellyfish
2338:butterfly
2231:anisogamy
2219:synchrony
2209:cuckoldry
2179:polyandry
2029:evolution
1963:206782867
1282:0005-7959
1262:Behaviour
1243:1045-2249
1154:Lee, PL;
520:0003-3472
485:1522-0613
398:marmosets
340:queen bee
264:D. anilis
174:queen bee
84:marmosets
48:polyandry
21:Polyandry
2820:Ethology
2767:mandrill
2725:kangaroo
2668:Elephant
2579:seabirds
2511:Reptiles
2461:Seahorse
2359:Cnidaria
2323:scorpion
2256:coercion
2214:seasonal
2184:polygyny
2169:monogamy
2142:external
2137:internal
1899:32094448
1835:33428552
1827:18452517
1778:19675002
1729:16701243
1684:22768149
1644:PLOS ONE
1610:35627708
1504:12920101
1407:53188748
1369:19726479
1320:53185546
1204:15096623
1156:Hays, GC
1136:24273373
1074:53171933
1030:35179878
937:25478149
837:16489146
792:33260761
705:24266042
644:20395774
636:10973716
528:10617092
416:See also
402:tamarins
336:eusocial
102:pipefish
98:dunnocks
56:Polygyny
2784:Raccoon
2757:gorilla
2693:cheetah
2678:Felidae
2663:Dolphin
2656:red fox
2596:Mammals
2546:Tuatara
2524:lizards
2416:epitoky
2352:octopus
2011:General
1933:Bibcode
1869:Bibcode
1805:Bibcode
1769:2817239
1697:1371–9.
1675:3387204
1652:Bibcode
1590:Bibcode
1555:7209555
1547:1686202
1527:Bibcode
1519:Science
1460:2081814
1440:Bibcode
1360:2821345
1172:Bibcode
1127:3827896
1106:Bibcode
1010:Bibcode
967:Bibcode
928:4224532
907:Bibcode
772:Bibcode
696:3813325
292:cichlid
94:jacanas
2789:Rodent
2752:bonobo
2737:walrus
2636:coyote
2529:snakes
2466:Sharks
2407:Worms
2403:Sponge
2333:insect
2328:beetle
2318:spider
2236:oogamy
1961:
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294:fish (
231:cervix
182:drones
72:, the
2747:human
2705:Fossa
2688:tiger
2641:dingo
2626:Canid
2556:Birds
2489:frogs
2374:coral
2164:Modes
1959:S2CID
1921:(PDF)
1895:S2CID
1857:(PDF)
1831:S2CID
1606:S2CID
1559:S2CID
1543:JSTOR
1456:S2CID
1428:(PDF)
1403:S2CID
1316:S2CID
1070:S2CID
1026:S2CID
986:(PDF)
955:(PDF)
788:S2CID
640:S2CID
524:S2CID
330:Some
223:glans
2683:lion
2451:Eels
2433:Fish
2313:crab
1823:PMID
1774:PMID
1725:PMID
1680:PMID
1551:PMID
1500:PMID
1365:PMID
1278:ISSN
1239:ISSN
1200:PMID
1132:PMID
933:PMID
833:PMID
701:PMID
632:PMID
516:ISSN
481:ISSN
400:and
350:and
332:taxa
323:'s (
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135:and
96:and
2609:rut
1949:hdl
1941:doi
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