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Harem (zoology)

A harem is an animal group consisting of one or two males, a number of females, and their offspring. The dominant male drives off other males and maintains the unity of the group. If present, the second male is subservient to the dominant male. As juvenile males grow, they leave the group and roam as solitary individuals or join bachelor herds. Females in the group may be inter-related. The dominant male mates with the females as they become sexually active and drives off competitors, until he is displaced by another male. In some species, incoming males that achieve dominant status may commit infanticide.

Large male northern fur seal and harem of smaller females

For the male, the primary benefit of the harem system is obtaining exclusive access to a group of mature females. The females benefit from being in a stable social group and the associated benefits of grooming, predator avoidance and cooperative defense of territory. The disadvantages for the male are the energetic costs of gaining or defending a harem which may leave him with reduced reproductive success. The females are disadvantaged if their offspring are killed during dominance battles or by incoming males.

Overview

The term harem is used in zoology to distinguish social organization consisting of a group of females, their offspring, and one to two males.[1]

The single male, called the dominant male, may be accompanied by another young male, called a "follower" male. Females that closely associate with the dominant male are called "central females," while females who associate less frequently with the dominant male are called "peripheral females."[2] Juvenile male offspring leave the harem and live either solitarily, or, with other young males in groups known as bachelor herds.[3] Sexually mature female offspring may stay within their natal harem, or may join another harem.[4] The females in a harem may be, but are not exclusively, genetically related.[1][5][6] For instance, the females in hamadryas baboon harems are not usually genetically related because their harems are formed by "kidnapping" females from other harems and subsequent herding.[1] In contrast, gelada harems are based on kinship ties to genetically related females.[7] Multiple harems may assemble into larger groups known as "clans" or "teams".[8]

Harem cohesiveness is mediated by the dominant male who fights off invading males to keep claim over the harem.[9][10][11] In some harem-forming species, when a dominant male vacates his harem (due to death, defection to another harem, or usurpation) the incoming male sometimes commits infanticide of the offspring.[12] Because time and resources are no longer being devoted to the offspring, infanticide often stimulates the female to return to sexual receptivity and fertility sooner than if the offspring were to survive. Furthermore, while lactating, females do not ovulate and consequently are not fertile. Infanticide therefore has the potential to increase the incoming male's reproductive success.[12][13]

Benefits

Harems are a beneficial social structure for the dominant male, as it allows him access to several reproductively available females at a time.[10] Harems provide protection for the females within a particular harem, as dominant males will fiercely ward off potential invaders.[11] This level of protection may also, such in the case of the common pheasant, reduce the energy expended by females on remaining alert to, or fleeing from, invading males.[11] Harems allow bonding and socialization among the female members, which can result in greater control over access to females as determined by the females' preferences. Harems also facilitate socialized behavior such as grooming and cooperative defense of territory.[1][14]

Costs

Harems can prove energetically costly for both males and females. Males spend substantial amounts of energy engaging in battles to invade a harem, or to keep hold of a harem once dominance has been established.[9] Such energy expenditure can result in reduced reproductive success such as in the case of red deer.[9] This is especially true when there is high turnover rates of dominant males, as frequent intense fighting can result in great expenditure of energy.[9] High turnover rate of dominant males can also be energetically costly for the females as their offspring are frequently killed in harems where infanticide occurs. Harems can also negatively affect females if there is intra-harem competition among females for resources.[15]

A lower-cost alternative mating strategy, useful to bachelors without a harem, is kleptogyny or the "sneaky fucker strategy",[a][17] sneaking in to mate while the harem owner is distracted: in the case of red deer, when the harem stag is involved in a fight with another older stag.[17] The strategy is also recorded in the elephant seal.[18]

Examples

Animals that form harems include:

Mammals

Primates

Birds

Insects

Fish

Explanatory notes

  1. ^ The term was coined by the evolutionary biologist John Maynard Smith.[16]

References

  1. ^ a b c d e f g h Maestripieri, Dario; Mayhew, Jessica; Carlson, Cindy L.; Hoffman, Christy L.; and Radtke, Jennifer M. "." Folia Primatologica 78.1 (2007): 56-68
  2. ^ Kummer, Hans. Social Organization of Hamadryas Baboons. A Field Study. Basel: Karger, (1968.) Print.
  3. ^ David, J. H. M. "The Behaviour of the Bontebok, Damaliscis Dorcas Dorcas, (Pallas 1766), with Special Reference to Territorial Behaviour." Zeitschrift für Tierpsychologie 33 (1973): 38-107
  4. ^ Qi, Xiao-Guang; Li, Bau-Guo; Garber, Paul A.; Ji, Weihong; and Wanatabe, Kunio. "Social Dynamics of the Golden Snub-Nosed Monkey (Rhinopithecus Roxellana): Female Transfer and One-Male Unit Succession." American Journal of Primatology 71 (2009): 670-79
  5. ^ a b Ortega, Jorge; Maldonado, Jesus E.; Wilkinson, Gerald S.; Arita, Hector T.; and Fleischer, Robert C. "Male Dominance, Paternity, and Relatedness in the Jamaican Fruit-eating Bat (Artibeus Jamaicensis) 2018-08-23 at the Wayback Machine." Molecular Ecology 12.9 (2003): 2409-415
  6. ^ Greenwood, Paul J. "Mating Systems, Philopatry and Dispersal in Birds and Mammals." Animal Behaviour 28.4 (1980): 1140-162
  7. ^ Mori, Akio; Iwamoto, Toshitaka; Mori, Umeyo; and Bekele, Afework. "Sociological and Demographic Characteristics of a Recently Found Arsi Gelada Population in Ethiopia." Primates 40.2 (1999): 365-81
  8. ^ Schreier, Amy L.; and Swedell, Larissa. "The Fourth Level of Social Structure in a Multi-level Society: Ecological and Social Functions of Clans in Hamadryas Baboons." American Journal of Primatology 71.11 (2009): 948-55
  9. ^ a b c d e Bonenfant, Christophe; Gaillard, Jean-Michel; Klein, François; and Maillard, Daniel. "Variation in Harem Size of Red Deer (Cervus Elaphus L.): The Effects of Adult Sex Ratio and Age-structure." Journal of Zoology 264.1 (2004): 77-85
  10. ^ a b c McCann, T. S. "Aggression and Sexual Activity of Male Southern Elephant Seals, Mirounga Leonina." Journal of Zoology 195 (1981): 295-310. Web.
  11. ^ a b c d Ridley, M. W.; and Hill, D. A. "Social Organization in the Pheasant (Phasianus Colchicus): Harem Formation, Mate Selection and the Role of Mate Guarding." Journal of Zoology 211 (1987): 619-30
  12. ^ a b Swedell, Larissa; and Tesfaye, Teklu. "Infant Mortality after Takeovers in Wild Ethiopian Hamadryas Baboons." American Journal of Primatology 60.3 (2003): 113-18
  13. ^ Horev, Aviad; Yosef, Reuven; Tryjanowski, Piotr; and Ovidia; Ofer. "Consequences of Variation in Male Harem Size to Population Persistence: Modeling Poaching and Extinction Risk of Bengal Tigers (Panthera Tigris)." Biological Conservation 147.1 (2012): 22-31
  14. ^ Searcy, William A.; and Yasukawa, Ken. "Alternative Models of Territorial Polygyny in Birds." The American Naturalist 134.3 (1989): 323-43
  15. ^ a b Latty, Tanya M.; Magrath, Michael J. L.; and Symonds, Matthew R. E. "Harem Size and Oviposition Behaviour in a Polygynous Bark Beetle". Ecological Entomology 34.5 (2009): 562-68
  16. ^ Pallen, Mark (2011). The Rough Guide to Evolution. Rough Guides. pp. 182–. ISBN 978-1-4093-5855-8.
  17. ^ a b Cherfas, Jeremy (15 September 1977). "The games animals play". New Scientist. pp. 672–673.
  18. ^ Frankenhuis, Willem E.; Fraley, R. Chris (2017). "What Do Evolutionary Models Teach Us About Sensitive Periods in Psychological Development?". European Psychologist. 22 (3): 141–150. doi:10.1027/1016-9040/a000265. S2CID 96439286.
  19. ^ Storz, Jay F.; Bhat, Hari R.; and Kunz, Thomas H. "Social Structure of a Polygynous Tent-making Bat, Cynopterus Sphinx (Megachiroptera)." Journal of Zoology 251.2 (2000): 151-65
  20. ^ Agoramoorthy, Govindasamy. "Adult Male Replacement and Social Change in Two Troops of Hanuman Langurs (Presbytis entellus) at Jodhpur, India 2017-12-10 at the Wayback Machine." International Journal of Primatology 15.2 (1994): 225-38
  21. ^ Watts, D. P. (1996). "Comparative socio-ecology of gorillas". In McGrew, W. C.; Marchant, L. F.; Nishida, T. (eds.). Great ape societies. Cambridge, UK: Cambridge University Press. pp. 16–28.
  22. ^ Codenotti, Thaïs L.; and Alvarez, Fernando. "Mating Behavior Of The Male Greater Rhea." The Wilson Bulletin 113.1 (2001): 85-89
  23. ^ Kelly, Clint D.; and Jennions, Michael D. "Sexually Dimorphic Immune Response in the Harem Polygynous Wellington Tree Weta Hemideina Crassidens". Physiological Entomology 34.2 (2009): 174-79
  24. ^ Baker, Richard H.; Ashwell, Robert I. S.; Richards, Thomas A.; Fowler, Kevin; Chapman, Tracey; Pomiankowski, Andrew (2001-11-01). "Effects of multiple mating and male eye span on female reproductive output in the stalk-eyed fly, Cyrtodiopsis dalmanni". Behavioral Ecology. 12 (6): 732–739. doi:10.1093/beheco/12.6.732. ISSN 1045-2249.
  25. ^ Colin, P. L. "Spawning and larval development of the hogfish, Lachnolaimus maximus (Pisces: Labridae)". Fish. Bull. 80 (1982): 853–862.
  26. ^ Coleman, Ron. "Something Old Doing Something New". Cichlid News Magazine (1998): 30-31
  27. ^ Froeschke, John (2006). "The Fish Assemblages Inside and Outside of a Temperate Marine Reserve in Southern California". Bulletin, Southern California Academy of Sciences. 10 (3): 128–142. doi:10.3160/0038-3872(2006)105[128:tfaiao]2.0.co;2.

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A harem is an animal group consisting of one or two males a number of females and their offspring The dominant male drives off other males and maintains the unity of the group If present the second male is subservient to the dominant male As juvenile males grow they leave the group and roam as solitary individuals or join bachelor herds Females in the group may be inter related The dominant male mates with the females as they become sexually active and drives off competitors until he is displaced by another male In some species incoming males that achieve dominant status may commit infanticide Large male northern fur seal and harem of smaller females For the male the primary benefit of the harem system is obtaining exclusive access to a group of mature females The females benefit from being in a stable social group and the associated benefits of grooming predator avoidance and cooperative defense of territory The disadvantages for the male are the energetic costs of gaining or defending a harem which may leave him with reduced reproductive success The females are disadvantaged if their offspring are killed during dominance battles or by incoming males Contents 1 Overview 2 Benefits 3 Costs 4 Examples 4 1 Mammals 4 1 1 Primates 4 2 Birds 4 3 Insects 4 4 Fish 5 Explanatory notes 6 ReferencesOverview EditThe term harem is used in zoology to distinguish social organization consisting of a group of females their offspring and one to two males 1 The single male called the dominant male may be accompanied by another young male called a follower male Females that closely associate with the dominant male are called central females while females who associate less frequently with the dominant male are called peripheral females 2 Juvenile male offspring leave the harem and live either solitarily or with other young males in groups known as bachelor herds 3 Sexually mature female offspring may stay within their natal harem or may join another harem 4 The females in a harem may be but are not exclusively genetically related 1 5 6 For instance the females in hamadryas baboon harems are not usually genetically related because their harems are formed by kidnapping females from other harems and subsequent herding 1 In contrast gelada harems are based on kinship ties to genetically related females 7 Multiple harems may assemble into larger groups known as clans or teams 8 Harem cohesiveness is mediated by the dominant male who fights off invading males to keep claim over the harem 9 10 11 In some harem forming species when a dominant male vacates his harem due to death defection to another harem or usurpation the incoming male sometimes commits infanticide of the offspring 12 Because time and resources are no longer being devoted to the offspring infanticide often stimulates the female to return to sexual receptivity and fertility sooner than if the offspring were to survive Furthermore while lactating females do not ovulate and consequently are not fertile Infanticide therefore has the potential to increase the incoming male s reproductive success 12 13 Benefits EditHarems are a beneficial social structure for the dominant male as it allows him access to several reproductively available females at a time 10 Harems provide protection for the females within a particular harem as dominant males will fiercely ward off potential invaders 11 This level of protection may also such in the case of the common pheasant reduce the energy expended by females on remaining alert to or fleeing from invading males 11 Harems allow bonding and socialization among the female members which can result in greater control over access to females as determined by the females preferences Harems also facilitate socialized behavior such as grooming and cooperative defense of territory 1 14 Costs EditFurther information Alternative mating strategy Harems can prove energetically costly for both males and females Males spend substantial amounts of energy engaging in battles to invade a harem or to keep hold of a harem once dominance has been established 9 Such energy expenditure can result in reduced reproductive success such as in the case of red deer 9 This is especially true when there is high turnover rates of dominant males as frequent intense fighting can result in great expenditure of energy 9 High turnover rate of dominant males can also be energetically costly for the females as their offspring are frequently killed in harems where infanticide occurs Harems can also negatively affect females if there is intra harem competition among females for resources 15 A lower cost alternative mating strategy useful to bachelors without a harem is kleptogyny or the sneaky fucker strategy a 17 sneaking in to mate while the harem owner is distracted in the case of red deer when the harem stag is involved in a fight with another older stag 17 The strategy is also recorded in the elephant seal 18 Examples EditAnimals that form harems include Mammals Edit Red deer 9 Sika deer Elk Fallow deer Collared Peccary Fur seal Elephant seal 10 Greater short nosed fruit bat 19 Jamaican fruit bat 5 African lion harem referred to as a pride with the infrequent strategy of two coequal males often brothers who enter the territory of an existing harem and share the dominance which makes it easier to depose the existing alpha male Primates Edit Hamadryas baboon 1 Gelada baboon 1 Golden snub nosed monkey 1 Guinea baboon 1 Gray langurs 20 Chimpanzee Gorilla 21 Birds Edit Common pheasant 11 Greater rhea 22 Northern harrier red winged blackbirdInsects Edit Bark beetle 15 Tree weta 23 Malaysian stalk eyed fly forms temporary harems even though mating frequency is high and the behavior of multiple mating partners is common 24 Fish Edit Hogfish 25 Several species of cichlid 26 California sheephead 27 Explanatory notes Edit The term was coined by the evolutionary biologist John Maynard Smith 16 References Edit a b c d e f g h Maestripieri Dario Mayhew Jessica Carlson Cindy L Hoffman Christy L and Radtke Jennifer M One Male Harems and Female Social Dynamics in Guinea Baboons Folia Primatologica 78 1 2007 56 68 Kummer Hans Social Organization of Hamadryas Baboons A Field Study Basel Karger 1968 Print David J H M The Behaviour of the Bontebok Damaliscis Dorcas Dorcas Pallas 1766 with Special Reference to Territorial Behaviour Zeitschrift fur Tierpsychologie 33 1973 38 107 Qi Xiao Guang Li Bau Guo Garber Paul A Ji Weihong and Wanatabe Kunio Social Dynamics of the Golden Snub Nosed Monkey Rhinopithecus Roxellana Female Transfer and One Male Unit Succession American Journal of Primatology 71 2009 670 79 a b Ortega Jorge Maldonado Jesus E Wilkinson Gerald S Arita Hector T and Fleischer Robert C Male Dominance Paternity and Relatedness in the Jamaican Fruit eating Bat Artibeus Jamaicensis Archived 2018 08 23 at the Wayback Machine Molecular Ecology 12 9 2003 2409 415 Greenwood Paul J Mating Systems Philopatry and Dispersal in Birds and Mammals Animal Behaviour 28 4 1980 1140 162 Mori Akio Iwamoto Toshitaka Mori Umeyo and Bekele Afework Sociological and Demographic Characteristics of a Recently Found Arsi Gelada Population in Ethiopia Primates 40 2 1999 365 81 Schreier Amy L and Swedell Larissa The Fourth Level of Social Structure in a Multi level Society Ecological and Social Functions of Clans in Hamadryas Baboons American Journal of Primatology 71 11 2009 948 55 a b c d e Bonenfant Christophe Gaillard Jean Michel Klein Francois and Maillard Daniel Variation in Harem Size of Red Deer Cervus Elaphus L The Effects of Adult Sex Ratio and Age structure Journal of Zoology 264 1 2004 77 85 a b c McCann T S Aggression and Sexual Activity of Male Southern Elephant Seals Mirounga Leonina Journal of Zoology 195 1981 295 310 Web a b c d Ridley M W and Hill D A Social Organization in the Pheasant Phasianus Colchicus Harem Formation Mate Selection and the Role of Mate Guarding Journal of Zoology 211 1987 619 30 a b Swedell Larissa and Tesfaye Teklu Infant Mortality after Takeovers in Wild Ethiopian Hamadryas Baboons American Journal of Primatology 60 3 2003 113 18 Horev Aviad Yosef Reuven Tryjanowski Piotr and Ovidia Ofer Consequences of Variation in Male Harem Size to Population Persistence Modeling Poaching and Extinction Risk of Bengal Tigers Panthera Tigris Biological Conservation 147 1 2012 22 31 Searcy William A and Yasukawa Ken Alternative Models of Territorial Polygyny in Birds The American Naturalist 134 3 1989 323 43 a b Latty Tanya M Magrath Michael J L and Symonds Matthew R E Harem Size and Oviposition Behaviour in a Polygynous Bark Beetle Ecological Entomology 34 5 2009 562 68 Pallen Mark 2011 The Rough Guide to Evolution Rough Guides pp 182 ISBN 978 1 4093 5855 8 a b Cherfas Jeremy 15 September 1977 The games animals play New Scientist pp 672 673 Frankenhuis Willem E Fraley R Chris 2017 What Do Evolutionary Models Teach Us About Sensitive Periods in Psychological Development European Psychologist 22 3 141 150 doi 10 1027 1016 9040 a000265 S2CID 96439286 Storz Jay F Bhat Hari R and Kunz Thomas H Social Structure of a Polygynous Tent making Bat Cynopterus Sphinx Megachiroptera Journal of Zoology 251 2 2000 151 65 Agoramoorthy Govindasamy Adult Male Replacement and Social Change in Two Troops of Hanuman Langurs Presbytis entellus at Jodhpur India Archived 2017 12 10 at the Wayback Machine International Journal of Primatology 15 2 1994 225 38 Watts D P 1996 Comparative socio ecology of gorillas In McGrew W C Marchant L F Nishida T eds Great ape societies Cambridge UK Cambridge University Press pp 16 28 Codenotti Thais L and Alvarez Fernando Mating Behavior Of The Male Greater Rhea The Wilson Bulletin 113 1 2001 85 89 Kelly Clint D and Jennions Michael D Sexually Dimorphic Immune Response in the Harem Polygynous Wellington Tree Weta Hemideina Crassidens Physiological Entomology 34 2 2009 174 79 Baker Richard H Ashwell Robert I S Richards Thomas A Fowler Kevin Chapman Tracey Pomiankowski Andrew 2001 11 01 Effects of multiple mating and male eye span on female reproductive output in the stalk eyed fly Cyrtodiopsis dalmanni Behavioral Ecology 12 6 732 739 doi 10 1093 beheco 12 6 732 ISSN 1045 2249 Colin P L Spawning and larval development of the hogfish Lachnolaimus maximus Pisces Labridae Fish Bull 80 1982 853 862 Coleman Ron Something Old Doing Something New Cichlid News Magazine 1998 30 31 Froeschke John 2006 The Fish Assemblages Inside and Outside of a Temperate Marine Reserve in Southern California Bulletin Southern California Academy of Sciences 10 3 128 142 doi 10 3160 0038 3872 2006 105 128 tfaiao 2 0 co 2 Retrieved from https en wikipedia org w index php title Harem zoology amp oldid 1088264239, wikipedia, wiki, book, books, library,

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