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A pseudo-penis is any structure found on an animal that, while superficially appearing to be a penis, is derived from a different developmental path.

Mammals

In mammals, all intact developmentally typical males have a penis, but the clitoris in the females of the following species is sufficiently enlarged that it is usually termed a pseudo-penis: spotted hyena,^[1]^[2] juvenile fossa,^[3] binturong,^[4] lemur^[5] and spider monkey. The fossa's enlarged clitoris is supported by an os clitoridis, a bone similar to the male's os penis.^[3] The juvenile female fossa's os clitoridis and pseudo-penis shrink as she grows, unlike that of other pseudo-penis species.^[3]

The mammalian pseudo-penis appears to be simply for display, though the spotted hyena is an exception: the female spotted hyena urinates, copulates, and gives birth through her pseudo-penis. This prevents males from mating without the full co-operation of females, which means that mating preferences of the female are predominant.^[6]

Female spotted hyenas urinate through a pseudo-penis.^[7]

Spotted hyenas are a matriarchal society, where adult females dominate adult males^[8] and are also more aggressive than males. When a male hyena leaves his natal clan, he behaves submissively to all newly encountered hyenas; as a result, when he settles down with a new clan as a breeding male, he is submissive to all natal clan members.^[9] It was hypothesized that the male hyena erected his penis to show submissiveness.^[10] During greetings, hyenas would stand parallel to each other and sniff or lick the erect penis or anal scent gland.^[10]

The male spotted hyena inserts his penis into the female's pseudo-penis during copulation.^[7]

The spotted hyena's pseudo-penis has severe reproductive costs. Nearly all of her first-born cubs are stillborn, as the placenta is not long enough for the extended penile birth canal. In addition, the first birthing process is time-consuming, as it requires the meatus of the pseudo-penis to tear, allowing the fetus to pass through; as a result, the first-born often die of hypoxia.^[11]

The female spider monkey has a clitoris that is called a pseudo-penis because it is especially developed and has a shallow perineal groove that retains and distributes urine droplets as she moves around.^[12] The clitoris of female Geoffroy's spider monkeys is large and protrudes, looking like a penis.^[13] This organ, called a pendulous clitoris because of the way it dangles externally, is actually larger than the male flaccid penis.^[14] As a result, females are sometimes mistaken for males by human observers.^[13] The enlarged clitoris is believed to aid males in determining sexual receptiveness, allowing them to touch the clitoris and smell their fingers to pick up chemical or olfactory cues to the female's reproductive status.^[14]

Birds

Only 3% of avian species have a phallus. The most common genitalia among birds is the cloaca; a direct tract for elimination and reproduction in both of the sexes.^{[citation needed]} Certain bird species, such as the ratites, screamers, waterfowl, and cracids (a family of arboreal galliformes) exhibit a phallus in males. A notable example of a bird with a pseudo-penis is the red-billed buffalo weaver, which do not use their pseudo-penis for direct insertion during copulation; however it does play a part in successful mating and stimulation.^[15] Similarly to the red-billed buffalo weaver, the cassowary, a ratite, exhibits a pseudo-penis in both males and females. The male's pseudo-phallus is used to "invaginate", or to push the female's pseudo-phallus inside-out, and then ejaculates from the cloaca to ensure a successful mating.^{[failed verification]}

Insects

The only known example of a pseudo-penis to occur in the insects is found in the barklouse genus Neotrogla. In this genus, the pseudo-penis plays a part in their rarely-seen sexual reversal. In this case, the male has a vagina-like structure while the female has a pseudo-penis. Some may be confused, wondering why the individuals with a vagina-like structure are considered male while the ones with the pseudo-penis are considered female.^[16] This is because even though they have visible outer structures of the opposite sex, their inner structures are consistent to their sex. More important, the males produce sperm and the females produce eggs.

Females have a penis-like structure, called the gynosome that has a tube leading into their body to where their genitalia are located.^[17] Neotrogla males have a structure resembling that of a vagina. However, on the inside of their body, they have male genitalia. When the female inserts her organ into the male, the tip of the pseudo penis inflates. When this tip inflates, species specific ridges and spines flare up that match up with the walls of the male's genital chamber. This serves two functions, to stimulate the male's reproductive organs, and to keep the male and female locked together. After they have been locked together, the only way to get the two to part would be to rip off the abdomen of the male. During the lengthy, 40- to 70-hour copulation process, these male genitalia structures ejaculate inside of the male's body.^[18] The sperm is then deposited into the female's structure and then travels through a spermathecal duct to where it can fertilize the eggs.

Possible role of androstenedione

Androstenedione is a hormone that is converted to testosterone by enzymatic activity.^[19] It is theorized that the dominance and morphological phenotype of a pseudo-penis observed in female hyenas is due to the presence of prenatal and postnatal androstenedione levels. Prenatal androgen levels dictate genitalia differences between male and female. Higher levels of androgen are observed in the second half of gestation which is theorized to cause masculinization in terms of dominance and aggression in hyenas.^[20] Large amounts of androstenedione are produced in hyena ovarian tissues with little aromatase activity allowing the placenta to convert androstenedione to testosterone.^[21] High concentrations of androgens is theorized to virilize the female hyena genitalia and kill ovarian follicles.^[21] Postnatal androgen levels dictate growth in genitalia during puberty. Postnatal androgen levels are higher in females than males when they are younger; especially in infancy.^[22] These high levels of androstenedione contribute to aggression and dominance and the masculinization of genitalia during puberty.^[22]

References

^ Place, Ned J.; Glickman, Stephen E. (2004). "Masculinization of Female Mammals: Lessons from Nature". Hypospadias and Genital Development. Advances in Experimental Medicine and Biology. Vol. 545. pp. 243–253. CiteSeerX 10.1.1.501.721. doi:10.1007/978-1-4419-8995-6_15. ISBN 978-1-4613-4752-1. PMID 15086031.
^ ADW: Crocuta crocuta: Information
^ ^a ^b ^c Goodman S (2009). "Family Eupleridae (Madagascar Carnivores)". In Wilson D; Mittermeier R (eds.). Handbook of the Mammals of the World. Volume 1: Carnivores. Barcelona: Lynx Edicions. ISBN 978-84-96553-49-1. Archived from the original on 2011-07-25. Retrieved 2010-05-21.
^ [1] Archived May 16, 2006, at the Wayback Machine
^ Angier 1999, pp. 68–69; Hall 2005, p. 344; Goodman 2009.
^ Owen, James (15 August 2007). "Hyenas Encourage Sex With Strangers to Prevent Incest". National Geographic News. Archived from the original on 8 February 2008.
^ ^a ^b Szykman, M.; Van Horn, R. C.; Engh, A.L.; Boydston, E. E.; Holekamp, K. E. (2007). "Courtship and mating in free-living spotted hyenas" (PDF). Behaviour. 144 (7): 815–846. CiteSeerX 10.1.1.630.5755. doi:10.1163/156853907781476418. Archived from the original (PDF) on 30 November 2012. Retrieved 24 December 2011.
^ Frank, Laurence G. (October 1986). "Social organization of the spotted hyaena Crocuta crocuta. II. Dominance and reproduction". Animal Behaviour. 34 (5): 1510–1527. doi:10.1016/S0003-3472(86)80221-4. S2CID 53193864.
^ Holekamp, Kay E.; Dantzer, Ben; Stricker, Gregory; Shaw Yoshida, Kathryn C.; Benson-Amram, Sarah (May 2015). "Brains, brawn and sociality: a hyaena's tale". Animal Behaviour. 103: 237–248. doi:10.1016/j.anbehav.2015.01.023. PMC 4493912. PMID 26160980.
^ ^a ^b East, MarionL.; Hofer, Heribert; Wickler, Wolfgang (December 1993). "The erect 'penis' is a flag of submission in a female-dominated society: greetings in Serengeti spotted hyenas". Behavioral Ecology and Sociobiology. 33 (6): 355–370. doi:10.1007/BF00170251. JSTOR 4600895. S2CID 23727803.
^ Glickman, Stephen E.; Cunha, Gerald R.; Drea, Christine M.; Conley, Alan J.; Place, Ned J. (November 2006). "Mammalian sexual differentiation: lessons from the spotted hyena". Trends in Endocrinology & Metabolism. 17 (9): 349–356. doi:10.1016/j.tem.2006.09.005. PMID 17010637. S2CID 18227659.
^ Dixson, Alan F. (2012). Primate Sexuality: Comparative Studies of the Prosimians, Monkeys, Apes, and Humans. Oxford University Press. p. 364. ISBN 978-0-19-954464-6. Retrieved 2012-11-22.
^ ^a ^b Wainwright, M. (2002). The Natural History of Costa Rican Mammals. Zona Tropical. pp. 146–149. ISBN 0-9705678-1-2.
^ ^a ^b Campbell, C. & Gibson, K. (2008). "Spider monkey reproduction and sexual behavior". In Campbell, C. (ed.). Spider Monkeys. Cambridge University Press. pp. 266–283. ISBN 978-0-521-86750-4.
^ Winterbottom, M.; Burke, T.; Birkhead, T. R. (May 1999). "A stimulatory phalloid organ in a weaver bird". Nature. 399 (6731): 28. Bibcode:1999Natur.399...28W. doi:10.1038/19884. S2CID 4431318.
^ Ed Yong (April 17, 2014). "In This Insect, Females Have Penises And Males Have Vaginas". Not Exactly Rocket Science. National Geographic. Archived from the original on April 20, 2014.
^ Yoshizawa, Kazunori; Ferreira, Rodrigo L.; Kamimura, Yoshitaka; Lienhard, Charles (May 2014). "Female Penis, Male Vagina, and Their Correlated Evolution in a Cave Insect". Current Biology. 24 (9): 1006–1010. doi:10.1016/j.cub.2014.03.022. hdl:2115/56857. PMID 24746797.
^ Jason G. Goldman (April 17, 2014). "Scientists Discover the Gynosome, a New Kind of Sexual Organ". Gizmodo.
^ Skinner, J. D.; Chimimba, Christian T. (2005-11-15). The Mammals of the Southern African Sub-region. Cambridge University Press. ISBN 9781107394056.
^ French, Jeffrey A.; Mustoe, Aaryn C.; Cavanaugh, Jon; Birnie, Andrew K. (5 December 2013). "The influence of androgenic steroid hormones on female aggression in 'atypical' mammals". Philosophical Transactions of the Royal Society B: Biological Sciences. 368 (1631): 20130084. doi:10.1098/rstb.2013.0084. PMC 3826213. PMID 24167314.
^ ^a ^b Yalcinkaya, T.; Siiteri, P.; Vigne, J.; Licht, P; Pavgi, S; Frank, L.; Glickman, S. (25 June 1993). "A mechanism for virilization of female spotted hyenas in utero". Science. 260 (5116): 1929–1931. Bibcode:1993Sci...260.1929Y. doi:10.1126/science.8391165. PMID 8391165.
^ ^a ^b Glickman, S E; Frank, L G; Davidson, J M; Smith, E R; Siiteri, P K (May 1987). "Androstenedione may organize or activate sex-reversed traits in female spotted hyenas". Proceedings of the National Academy of Sciences of the United States of America. 84 (10): 3444–3447. Bibcode:1987PNAS...84.3444G. doi:10.1073/pnas.84.10.3444. PMC 304887. PMID 3472215.

[1] Place, Ned J.; Glickman, Stephen E. (2004). "Masculinization of Female Mammals: Lessons from Nature". Hypospadias and Genital Development. Advances in Experimental Medicine and Biology. Vol. 545. pp. 243–253. CiteSeerX 10.1.1.501.721. doi:10.1007/978-1-4419-8995-6_15. ISBN 978-1-4613-4752-1. PMID 15086031.

[2] ADW: Crocuta crocuta: Information

[HMW1-3] Goodman S (2009). "Family Eupleridae (Madagascar Carnivores)". In Wilson D; Mittermeier R (eds.). Handbook of the Mammals of the World. Volume 1: Carnivores. Barcelona: Lynx Edicions. ISBN 978-84-96553-49-1. Archived from the original on 2011-07-25. Retrieved 2010-05-21.

[4] [1] Archived May 16, 2006, at the Wayback Machine

[Angier_Hall_Goodman-5] Angier 1999, pp. 68–69; Hall 2005, p. 344; Goodman 2009.

[6] Owen, James (15 August 2007). "Hyenas Encourage Sex With Strangers to Prevent Incest". National Geographic News. Archived from the original on 8 February 2008.

[courtship-7] Szykman, M.; Van Horn, R. C.; Engh, A.L.; Boydston, E. E.; Holekamp, K. E. (2007). "Courtship and mating in free-living spotted hyenas" (PDF). Behaviour. 144 (7): 815–846. CiteSeerX 10.1.1.630.5755. doi:10.1163/156853907781476418. Archived from the original (PDF) on 30 November 2012. Retrieved 24 December 2011.

[8] Frank, Laurence G. (October 1986). "Social organization of the spotted hyaena Crocuta crocuta. II. Dominance and reproduction". Animal Behaviour. 34 (5): 1510–1527. doi:10.1016/S0003-3472(86)80221-4. S2CID 53193864.

[9] Holekamp, Kay E.; Dantzer, Ben; Stricker, Gregory; Shaw Yoshida, Kathryn C.; Benson-Amram, Sarah (May 2015). "Brains, brawn and sociality: a hyaena's tale". Animal Behaviour. 103: 237–248. doi:10.1016/j.anbehav.2015.01.023. PMC 4493912. PMID 26160980.

[jstor4600895-10] East, MarionL.; Hofer, Heribert; Wickler, Wolfgang (December 1993). "The erect 'penis' is a flag of submission in a female-dominated society: greetings in Serengeti spotted hyenas". Behavioral Ecology and Sociobiology. 33 (6): 355–370. doi:10.1007/BF00170251. JSTOR 4600895. S2CID 23727803.

[11] Glickman, Stephen E.; Cunha, Gerald R.; Drea, Christine M.; Conley, Alan J.; Place, Ned J. (November 2006). "Mammalian sexual differentiation: lessons from the spotted hyena". Trends in Endocrinology & Metabolism. 17 (9): 349–356. doi:10.1016/j.tem.2006.09.005. PMID 17010637. S2CID 18227659.

[Dixson-12] Dixson, Alan F. (2012). Primate Sexuality: Comparative Studies of the Prosimians, Monkeys, Apes, and Humans. Oxford University Press. p. 364. ISBN 978-0-19-954464-6. Retrieved 2012-11-22.

[costa-13] Wainwright, M. (2002). The Natural History of Costa Rican Mammals. Zona Tropical. pp. 146–149. ISBN 0-9705678-1-2.

[spidersex-14] Campbell, C. & Gibson, K. (2008). "Spider monkey reproduction and sexual behavior". In Campbell, C. (ed.). Spider Monkeys. Cambridge University Press. pp. 266–283. ISBN 978-0-521-86750-4.

[15] Winterbottom, M.; Burke, T.; Birkhead, T. R. (May 1999). "A stimulatory phalloid organ in a weaver bird". Nature. 399 (6731): 28. Bibcode:1999Natur.399...28W. doi:10.1038/19884. S2CID 4431318.

[16] Ed Yong (April 17, 2014). "In This Insect, Females Have Penises And Males Have Vaginas". Not Exactly Rocket Science. National Geographic. Archived from the original on April 20, 2014.

[17] Yoshizawa, Kazunori; Ferreira, Rodrigo L.; Kamimura, Yoshitaka; Lienhard, Charles (May 2014). "Female Penis, Male Vagina, and Their Correlated Evolution in a Cave Insect". Current Biology. 24 (9): 1006–1010. doi:10.1016/j.cub.2014.03.022. hdl:2115/56857. PMID 24746797.

[18] Jason G. Goldman (April 17, 2014). "Scientists Discover the Gynosome, a New Kind of Sexual Organ". Gizmodo.

[19] Skinner, J. D.; Chimimba, Christian T. (2005-11-15). The Mammals of the Southern African Sub-region. Cambridge University Press. ISBN 9781107394056.

[20] French, Jeffrey A.; Mustoe, Aaryn C.; Cavanaugh, Jon; Birnie, Andrew K. (5 December 2013). "The influence of androgenic steroid hormones on female aggression in 'atypical' mammals". Philosophical Transactions of the Royal Society B: Biological Sciences. 368 (1631): 20130084. doi:10.1098/rstb.2013.0084. PMC 3826213. PMID 24167314.

[pmid8391165-21] Yalcinkaya, T.; Siiteri, P.; Vigne, J.; Licht, P; Pavgi, S; Frank, L.; Glickman, S. (25 June 1993). "A mechanism for virilization of female spotted hyenas in utero". Science. 260 (5116): 1929–1931. Bibcode:1993Sci...260.1929Y. doi:10.1126/science.8391165. PMID 8391165.

[pmid3472215-22] Glickman, S E; Frank, L G; Davidson, J M; Smith, E R; Siiteri, P K (May 1987). "Androstenedione may organize or activate sex-reversed traits in female spotted hyenas". Proceedings of the National Academy of Sciences of the United States of America. 84 (10): 3444–3447. Bibcode:1987PNAS...84.3444G. doi:10.1073/pnas.84.10.3444. PMC 304887. PMID 3472215.

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+{{Short description|Anatomical structure resembling a penis}}
 A '''pseudo-penis''' is any structure found on an animal that, while superficially appearing to be a [[penis]], is derived from a different developmental path.
-==Mammals==
+== Mammals ==
-{{further|Fossa (animal)#External genitalia|Clitoris#Spotted hyena|Spider monkey#Pseudo-penis}}
+{{further|Fossa (animal)#External genitalia|Clitoris#Spotted hyena|Spider monkey#Pseudo-penis|Pseudo-penises of spotted hyenas}}
+[[File:Anatomischer_Anzeiger_(1922)_(18006271698).jpg|thumb|Male and female reproductive systems of the spotted hyena]]
-In mammals, all intact developmentally typical males have a penis, but the [[clitoris]] in the females of the following species is sufficiently enlarged that it is usually termed a pseudo-penis: [[spotted hyena]],<ref>[http://animaldiversity.ummz.umich.edu/site/accounts/information/Crocuta_crocuta.html ADW: Crocuta crocuta: Information<!-- Bot generated title -->]</ref> [[squirrel monkey]],{{citation needed|date=April 2013}} [[lemur]],<ref name="Angier_Hall_Goodman">
+In mammals, all intact developmentally typical males have a penis, but the [[Animal clitoris|clitoris]] in the females of the following species is sufficiently enlarged that it is usually termed a pseudo-penis: [[spotted hyena]],<ref>{{cite book |last1=Place |first1=Ned J. |last2=Glickman |first2=Stephen E. |chapter=Masculinization of Female Mammals: Lessons from Nature |title=Hypospadias and Genital Development |series=Advances in Experimental Medicine and Biology |date=2004 |volume=545 |pages=243–253 |doi=10.1007/978-1-4419-8995-6_15 |pmid=15086031 |isbn=978-1-4613-4752-1 |citeseerx=10.1.1.501.721 }}</ref><ref>[http://animaldiversity.ummz.umich.edu/site/accounts/information/Crocuta_crocuta.html ADW: Crocuta crocuta: Information<!-- Bot generated title -->]</ref>  juvenile [[fossa (animal)|fossa]],<ref name = HMW1>{{cite book | author = Goodman S | editor1 = Wilson D | editor2 = Mittermeier R | chapter = Family Eupleridae (Madagascar Carnivores) | title = Handbook of the Mammals of the World. Volume 1: Carnivores | year = 2009 | place = Barcelona | publisher = Lynx Edicions | isbn = 978-84-96553-49-1 | chapter-url = http://www.lynxeds.com/hmw/species-accounts/hmw-1-species-accounts-red-panda-ailurus-fulgens | title-link = Handbook of the Mammals of the World | access-date = 2010-05-21 | archive-date = 2011-07-25 | archive-url = https://web.archive.org/web/20110725081653/http://www.lynxeds.com/hmw/species-accounts/hmw-1-species-accounts-red-panda-ailurus-fulgens | url-status = dead }}</ref> [[binturong]],<ref>[http://www.conservatorscenter.org/speciesinfo/bint.pdf]  {{webarchive|url=https://web.archive.org/web/20060516213601/http://www.conservatorscenter.org/speciesinfo/bint.pdf|date=May 16, 2006}}</ref> [[lemur]]<ref name="Angier_Hall_Goodman">
 {{harvnb|Angier|1999|pp=[https://books.google.com/books?id=NIonAf-m9OwC&pg=PT95 68–69]}};
 {{harvnb|Hall|2005|p=[https://books.google.com/books?id=y-RWPGDONlIC&pg=PA344 344]}};
+{{harvnb|Goodman|2009}}.</ref> and [[spider monkey]]. The fossa's enlarged clitoris is supported by an [[os clitoridis]], a bone similar to the male's [[os penis]].<ref name=HMW1/> The juvenile female fossa's os clitoridis and pseudo-penis shrink as she grows, unlike that of other pseudo-penis species.<ref name=HMW1/>
-{{harvnb|Goodman|2009}}.
-</ref> juvenile [[fossa (animal)|fossa]]<ref name = HMW1>{{cite book | author = Goodman S | editors = Wilson D & Mittermeier R | chapter= Family Eupleridae (Madagascar Carnivores) | title = [[Handbook of the Mammals of the World]]. Volume 1: Carnivores | year = 2009 | pages =  | place = Barcelona | publisher = Lynx Edicions | isbn = 978-84-96553-49-1 | chapter-url = http://www.lynxeds.com/hmw/species-accounts/hmw-1-species-accounts-red-panda-ailurus-fulgens}}</ref> and [[binturong]].<ref>[http://www.conservatorscenter.org/speciesinfo/bint.pdf ]  {{webarchive |url=https://web.archive.org/web/20060516213601/http://www.conservatorscenter.org/speciesinfo/bint.pdf |date=May 16, 2006 }}</ref> The enlarged clitoris in the fossa is supported by an [[Baculum|os clitoridis]], a bone similar to the [[os penis]] found in most mammals.<ref name=HMW1/> However, the fossa's os clitoridis and pseudo-penis shrink as the juvenile female grows, unlike that of other pseudo-penis species.<ref name=HMW1/> The [[labia]] of the [[spider monkey]] are elongated and may be similarly confused during display. [[Elongated labia]] are also observed in humans.
+The mammalian pseudo-penis appears to be simply for display, though the spotted hyena is an exception: the female spotted hyena urinates, copulates, and gives birth through her pseudo-penis. This prevents males from mating without the full co-operation of females, which means that mating preferences of the female are predominant.<ref>{{cite news |first1=James |last1=Owen |date=15 August 2007 |title=Hyenas Encourage Sex With Strangers to Prevent Incest |work=National Geographic News |url=http://news.nationalgeographic.com/news/2007/08/070815-hyenas-sex.html |archive-url=https://web.archive.org/web/20080208115722/http://news.nationalgeographic.com/news/2007/08/070815-hyenas-sex.html |archive-date=8 February 2008 }}</ref>{{CSS image crop|Image=Spotted_hyena_-_Addo_Elephant_National_Park_(33948058355).jpg|bSize=660|cWidth=220|cHeight=120|oTop=250|oLeft=120|Description=Female spotted hyenas [[urination|urinate]] through a pseudo-penis.<ref name="courtship"/>}} Spotted hyenas are a matriarchal society, where adult females dominate adult males<ref>{{cite journal |last1=Frank |first1=Laurence G. |title=Social organization of the spotted hyaena Crocuta crocuta. II. Dominance and reproduction |journal=Animal Behaviour |date=October 1986 |volume=34 |issue=5 |pages=1510–1527 |doi=10.1016/S0003-3472(86)80221-4 |s2cid=53193864 }}</ref> and are also more aggressive than males. When a male hyena leaves his natal clan, he behaves submissively to all newly encountered hyenas; as a result, when he settles down with a new clan as a breeding male, he is submissive to all natal clan members.<ref>{{cite journal |last1=Holekamp |first1=Kay E. |last2=Dantzer |first2=Ben |last3=Stricker |first3=Gregory |last4=Shaw Yoshida |first4=Kathryn C. |last5=Benson-Amram |first5=Sarah |title=Brains, brawn and sociality: a hyaena's tale |journal=Animal Behaviour |date=May 2015 |volume=103 |pages=237–248 |doi=10.1016/j.anbehav.2015.01.023 |pmid=26160980 |pmc=4493912 }}</ref> It was hypothesized that the male hyena erected his penis to show submissiveness.<ref name="jstor4600895">{{cite journal |last1=East |first1=MarionL. |last2=Hofer |first2=Heribert |last3=Wickler |first3=Wolfgang |title=The erect 'penis' is a flag of submission in a female-dominated society: greetings in Serengeti spotted hyenas |journal=Behavioral Ecology and Sociobiology |date=December 1993 |volume=33 |issue=6 |pages=355–370 |doi=10.1007/BF00170251 |jstor=4600895 |s2cid=23727803 }}</ref> During greetings, hyenas would stand parallel to each other and sniff or lick the erect penis or anal scent gland.<ref name="jstor4600895" />{{CSS image crop|Image=Mating hyenas.jpg|bSize=300|cWidth=220|cHeight=160|oTop=60|oLeft=60|Location=right|Description=The male spotted hyena inserts his penis into the female's pseudo-penis during [[copulation (zoology)|copulation]].<ref name="courtship">{{cite journal | last1 = Szykman | first1 = M. | last2 = Van Horn | first2 = R. C. | last3 = Engh | first3 = A.L. | last4 = Boydston | first4 = E. E. | last5 = Holekamp |first5=K. E. | year = 2007 | title = Courtship and mating in free-living spotted hyenas | url = http://tuvalu.santafe.edu/~bowles/Dominance/Papers/SzykmanetalHyenaMatingBehaviour2007.pdf | journal = Behaviour | volume = 144 | issue = 7 | pages = 815–846 | doi = 10.1163/156853907781476418 | citeseerx = 10.1.1.630.5755 | access-date = 24 December 2011 | archive-date = 30 November 2012 | archive-url = https://web.archive.org/web/20121130193631/http://tuvalu.santafe.edu/~bowles/Dominance/Papers/SzykmanetalHyenaMatingBehaviour2007.pdf | url-status = dead }}</ref>}} The spotted hyena's pseudo-penis has severe reproductive costs. Nearly all of her first-born cubs are stillborn, as the placenta is not long enough for the extended penile birth canal. In addition, the first birthing process is time-consuming, as it requires the [[Urinary meatus|meatus]] of the pseudo-penis to tear, allowing the fetus to pass through; as a result, the first-born often die of [[Hypoxia (medical)|hypoxia]].<ref>{{cite journal |last1=Glickman |first1=Stephen E. |last2=Cunha |first2=Gerald R. |last3=Drea |first3=Christine M. |last4=Conley |first4=Alan J. |last5=Place |first5=Ned J. |title=Mammalian sexual differentiation: lessons from the spotted hyena |journal=Trends in Endocrinology & Metabolism |date=November 2006 |volume=17 |issue=9 |pages=349–356 |doi=10.1016/j.tem.2006.09.005 |pmid=17010637 |s2cid=18227659 }}</ref>
-The mammalian pseudo-penis appears to be simply for display, though the spotted hyena is an exception: the female spotted hyena additionally uses her pseudo-penis for urination, copulation, and childbirth. In addition, this makes it difficult for males to mate without the full cooperation of females, which means that mating preferences of the female are predominant.<ref>[http://news.nationalgeographic.com/news/2007/08/070815-hyenas-sex_2.html Hyenas Encourage Sex With Strangers to Prevent Incest], ''[[National Geographic (magazine)|National Geographic]].''</ref>
+The female [[spider monkey]] has a clitoris that is called a pseudo-penis because it is especially developed and has a shallow perineal groove that retains and distributes urine droplets as she moves around.<ref name="Dixson">{{cite book |last=Dixson |first=Alan F. |title=Primate Sexuality: Comparative Studies of the Prosimians, Monkeys, Apes, and Humans |isbn=978-0-19-954464-6 |publisher=[[Oxford University Press]] |year=2012 |access-date=2012-11-22|page=364|url=https://books.google.com/books?id=SY-PyKNQglIC&pg=PA364}}</ref> The clitoris of female [[Geoffroy's spider monkey]]s is large and protrudes, looking like a penis.<ref name=costa>{{cite book|title=The Natural History of Costa Rican Mammals|author=Wainwright, M.|year=2002|publisher=Zona Tropical|isbn=0-9705678-1-2|pages=146–149}}</ref> This organ, called a pendulous clitoris because of the way it dangles externally, is actually larger than the male flaccid penis.<ref name=spidersex>{{cite book|title=Spider Monkeys|editor=Campbell, C.|chapter=Spider monkey reproduction and sexual behavior|author1=Campbell, C.  |author2=Gibson, K. |name-list-style=amp |pages=266–283|year=2008|publisher=Cambridge University Press|isbn=978-0-521-86750-4}}</ref> As a result, females are sometimes mistaken for males by human observers.<ref name=costa/> The enlarged clitoris is believed to aid males in determining sexual receptiveness, allowing them to touch the clitoris and smell their fingers to pick up [[olfactic communication|chemical or olfactory cues]] to the female's reproductive status.<ref name=spidersex/>
-[[Spotted hyenas]] are a matriarchal society, where adult female hyenas dominate the adult male hyenas.<ref>{{cite journal | url = http://www.sciencedirect.com/science/article/pii/S0003347286802214 | doi=10.1016/S0003-3472(86)80221-4 | volume=34 | title=Social organization of the spotted hyaena Crocuta crocuta. II. Dominance and reproduction | journal=Animal Behaviour | pages=1510–1527}}</ref>  Female [[spotted hyena]] are also more aggressive than the male [[spotted hyena]]. When a male hyena leaves its natal clan, it behaves submissively to all newly encountered hyenas; as a result, when a male hyena settles down with a new clan as a breeding male, it is submissive to all natal clan members.<ref>{{cite journal | url = http://www.sciencedirect.com/science/article/pii/S0003347215000330 | doi=10.1016/j.anbehav.2015.01.023 | volume=103 | title=Brains, brawn and sociality: a hyaena's tale | journal=Animal Behaviour | pages=237–248}}</ref> As a result of the submissive behavior in males, it was hypothesized that the male hyena erected its penis as a show of submissiveness.<ref name="jstor.org">http://www.jstor.org/stable/4600895?seq=1#page_scan_tab_contents</ref> During greetings, hyenas would stand parallel to each other and sniff or lick the erect penis or anal scent gland.<ref name="jstor.org"/>
-There are severe reproductive costs to this androgen-fueled hyper masculinization of female spotted hyena. Nearly all female spotted hyena’s first-born cubs are stillborn, as the placenta is not long enough for the extended penile birth canal. In addition, the first birthing process is time consuming, as it requires the meatus of the pseudo-penis to tear, allowing the fetus to pass through; as a result, the first-born often die of anoxia.<ref>{{cite journal | url = http://www.sciencedirect.com/science/article/pii/S1043276006001767 | doi=10.1016/j.tem.2006.09.005 | volume=17 | title=Mammalian sexual differentiation: lessons from the spotted hyena | journal=Trends in Endocrinology | pages=349–356 | pmid=17010637 | date=November 2006 | last1 = Glickman | first1 = SE | last2 = Cunha | first2 = GR | last3 = Drea | first3 = CM | last4 = Conley | first4 = AJ | last5 = Place | first5 = NJ}}</ref>
+== Birds ==
-Female spider monkeys have a clitoris that is referred to as a pseudo-penis because it is especially developed and has a shallow perineal groove that retains and distributes urine droplets as the female moves around.<ref name="Dixson">{{cite book |last=Dixson |first=Alan F. |title=Primate Sexuality: Comparative Studies of the Prosimians, Monkeys, Apes, and Humans |isbn=978-0-19-954464-6 |publisher=[[Oxford University Press]] |year=2012 |accessdate=2012-11-22|page=364|url=https://books.google.com/books?id=SY-PyKNQglIC&pg=PA364}}</ref> The clitoris of female [[Geoffroy's spider monkey]]s is large and protrudes, looking like a penis.<ref name=costa>{{cite book|title=The Natural History of Costa Rican Mammals|author=Wainwright, M.|year=2002|publisher=Zona Tropical|isbn=0-9705678-1-2|pages=146–149}}</ref> This organ, called a pendulous clitoris because of the way it dangles externally, is actually larger than the male flaccid penis.<ref name=spidersex>{{cite book|title=Spider Monkeys|editor=Campbell, C.|chapter=Spider monkey reproduction and sexual behavior|author1=Campbell, C.  |author2=Gibson, K. |lastauthoramp=yes |pages=266–283|year=2008|publisher=Cambridge University Press|isbn=978-0-521-86750-4}}</ref> As a result, females are sometimes mistaken for males by human observers.<ref name=costa/> The enlarged clitoris is believed to aid males in determining sexual receptiveness, allowing them to touch the clitoris and smell their fingers to pick up chemical or olfactory cues to the female's reproductive status.<ref name=spidersex/>
+{{disputed section|date=December 2019}}
-==Birds==
 {{see also|Bird penis|Avian reproductive system}}
+Only 3% of avian species have a phallus. The most common genitalia among birds is the [[cloaca]]; a direct tract for elimination and reproduction in both of the sexes.{{citation needed|date=May 2019}} Certain bird species, such as the ratites, screamers, waterfowl, and cracids (a family of arboreal galliformes) exhibit a phallus in males. A notable example of a bird with a pseudo-penis is the [[red-billed buffalo weaver]], which do not use their pseudo-penis for direct insertion during copulation; however it does play a part in successful mating and stimulation.<ref>{{cite journal |last1=Winterbottom |first1=M. |last2=Burke |first2=T. |last3=Birkhead |first3=T. R. |title=A stimulatory phalloid organ in a weaver bird |journal=Nature |date=May 1999 |volume=399 |issue=6731 |pages=28 |doi=10.1038/19884 |bibcode=1999Natur.399...28W |s2cid=4431318 |doi-access=free }}</ref> Similarly to the red-billed buffalo weaver, the cassowary, a ratite, exhibits a pseudo-penis in both males and females. The male's pseudo-phallus is used to "invaginate", or to push the female's pseudo-phallus inside-out, and then ejaculates from the cloaca to ensure a successful mating.{{failed verification|date=May 2017}}
-[[File:Mallard with visible pseudo-penis (croped).jpg|thumb|right|Pseudo-penis of a mallard{{dubious|date=November 2015}}]]
-Only 3% of avian species have a phallus. The most common genital among birds is the cloaca; a direct tract for elimination and reproduction in both of the sexes [10]. Certain bird species, such as the ratites, screamers, waterfowl, and cracids (a family of arboreal galliformes) exhibit a phallus in males. A notable example of a bird with a pseudo-penis is the [[red-billed buffalo weaver]], which do not use their pseudo-penis for direct insertion during copulation; however it does play a part in successful mating and stimulation.<ref>http://www.nature.com/news/2001/010716/full/news010719-4.html</ref> Similarly to the red-billed buffalo weaver, the cassowary, a ratite, exhibits a pseudo-penis in both males and females.{{not in citation given|date=May 2017}} The male’s pseudo-phallus is used to “invaginate”, or to push the female’s pseudo-phallus inside-out, and then ejaculates from the cloaca to ensure a successful mating.<ref>http://www.wholeearth.com/issue/2100/article/143/left-handed.bears.and.androgynous.cassowaries</ref>
-==Insects==
+== Insects ==
-One of the more known examples of a pseudo penis to occur in the insects is found in the species [[Neotrogla]]. In this species, the pseudo-penis plays a part in their rarely-seen sexual reversal. In this case, the male has a vagina-like structure while the female has a pseudo-penis. Some may be confused, wondering why the individuals with a vagina-like structure are considered male while the ones with the pseudo-penis are considered female.<ref>http://phenomena.nationalgeographic.com/2014/04/17/in-this-insect-females-have-penises-and-males-have-vaginas/</ref> This is because even though they have visible outer structures of the opposite gender, their inner structures are consistent to their genders.
+The only known example of a pseudo-penis to occur in the insects is found in the [[barklouse]] genus ''[[Neotrogla]]''. In this genus, the pseudo-penis plays a part in their rarely-seen sexual reversal. In this case, the male has a vagina-like structure while the female has a pseudo-penis. Some may be confused, wondering why the individuals with a vagina-like structure are considered male while the ones with the pseudo-penis are considered female.<ref>{{cite web |work=Not Exactly Rocket Science	|author=Ed Yong |title=In This Insect, Females Have Penises And Males Have Vaginas |date=April 17, 2014 |publisher=National Geographic |url=http://phenomena.nationalgeographic.com/2014/04/17/in-this-insect-females-have-penises-and-males-have-vaginas/|archive-url=https://web.archive.org/web/20140420012318/http://phenomena.nationalgeographic.com/2014/04/17/in-this-insect-females-have-penises-and-males-have-vaginas/|url-status=dead|archive-date=April 20, 2014}}</ref> This is because even though they have visible outer structures of the opposite sex, their inner structures are consistent to their sex. More important, the males produce sperm and the females produce eggs.
-Females have a penis-like structure, called the [[gynosome]] that has a tube leading into their body to where their genitalia are located.<ref>{{cite journal | url = http://www.sciencedirect.com/science/article/pii/S0960982214003145 | doi=10.1016/j.cub.2014.03.022 | volume=24 | title=Female Penis, Male Vagina, and Their Correlated Evolution in a Cave Insect | journal=Current Biology | pages=1006–1010 | pmid=24746797 | last1 = Yoshizawa | first1 = K | last2 = Ferreira | first2 = RL | last3 = Kamimura | first3 = Y | last4 = Lienhard | first4 = C}}</ref> [[Neotrogla]] males have a structure resembling that of a vagina. However, on the inside of their body, they have male genitalia. When the female inserts her organ into the male, the tip of the pseudo penis inflates. When this tip inflates, species specific ridges and spines flare up that match up with the walls of the male’s seminal duct. This serves two functions, to stimulate the male’s reproductive organs, and to keep the male and female locked together. After they have been locked together, the only way to get the two to part would be to rip off the abdomen of the male. During the lengthy, 40- to 70-hour copulation process, these male genitalia structures ejaculate inside of the male’s body.<ref>http://animals.io9.com/scientists-discover-the-gynosome-a-new-kind-of-sexual-1564382279</ref> The sperm is then deposited into the female’s structure and then travels through a spermathecal duct to where it can fertilize the eggs.
+Females have a penis-like structure, called the [[gynosome]] that has a tube leading into their body to where their genitalia are located.<ref>{{cite journal |last1=Yoshizawa |first1=Kazunori |last2=Ferreira |first2=Rodrigo L. |last3=Kamimura |first3=Yoshitaka |last4=Lienhard |first4=Charles |title=Female Penis, Male Vagina, and Their Correlated Evolution in a Cave Insect |journal=Current Biology |date=May 2014 |volume=24 |issue=9 |pages=1006–1010 |doi=10.1016/j.cub.2014.03.022 |pmid=24746797 |doi-access=free |hdl=2115/56857 |hdl-access=free }}</ref> ''Neotrogla'' males have a structure resembling that of a vagina. However, on the inside of their body, they have male genitalia. When the female inserts her organ into the male, the tip of the pseudo penis inflates. When this tip inflates, species specific ridges and spines flare up that match up with the walls of the male's genital chamber. This serves two functions, to stimulate the male's reproductive organs, and to keep the male and female locked together. After they have been locked together, the only way to get the two to part would be to rip off the abdomen of the male. During the lengthy, 40- to 70-hour copulation process, these male genitalia structures ejaculate inside of the male's body.<ref>{{cite web |title=Scientists Discover the Gynosome, a New Kind of Sexual Organ |author=Jason G. Goldman |date=April 17, 2014 |publisher=Gizmodo |url=https://gizmodo.com/scientists-discover-the-gynosome-a-new-kind-of-sexual-1564382279}}</ref> The sperm is then deposited into the female's structure and then travels through a spermathecal duct to where it can fertilize the eggs.
-==Possible Role of Androstenedione==
+== Possible role of androstenedione ==
-[[Androstenedione]] is a hormone that is converted to [[testosterone]] by enzymatic activity.<ref>https://books.google.com/books?id=F23lAgAAQBAJ&pg=PA376&lpg=PA376&dq=androstenedione+pseudopenis&source=bl&ots=ukSinY7jir&sig=6BfDMONSqoB6p24_KVrNQ67-rVI&hl=en&sa=X&ved=0ahUKEwiutai2l8XJAhUGwiYKHV10BWsQ6AEIUTAI#v=onepage&q=androstenedione%20pseudopenis&f=false)</ref> It is theorized that the dominance and morphological phenotype of a pseudo-penis observed in female hyenas is due to the presence of prenatal and postnatal androstenedione levels. Prenatal [[androgen]] levels dictate genitalia differences between male and female. Higher levels of [[androgen]] are observed in the second half of gestation which is theorized to cause masculinization in terms of dominance and aggression in hyenas.<ref>{{cite journal |pmid=24167314 | doi=10.1098/rstb.2013.0084 | pmc=3826213 | volume=368 | title=The influence of androgenic steroid hormones on female aggression in 'atypical' mammals | year=2013 | journal=Philos. Trans. R. Soc. Lond., B, Biol. Sci. | pages=20130084 | last1 = French | first1 = JA | last2 = Mustoe | first2 = AC | last3 = Cavanaugh | first3 = J | last4 = Birnie | first4 = AK}}</ref> Large amounts of androstenedione are produced in hyena ovarian tissues with little aromatase activity allowing the placenta to convert androstenedione to [[testosterone]].<ref name="sciencemag.org">http://www.sciencemag.org/content/260/5116/1929.abstract</ref> High concentrations of androgens is theorized to virilize the female hyena genitalia and kill ovarian follicles.<ref name="sciencemag.org"/>  Postnatal androgen levels dictate growth in genitalia during puberty. Postnatal androgen levels are higher in females than males when they are younger; especially in infancy.<ref name="pnas.org">http://www.pnas.org/content/84/10/3444.long</ref>  These high levels of androstenedione contribute to aggression and dominance and the masculinization of genitalia during puberty.<ref name="pnas.org"/>
+[[Androstenedione]] is a hormone that is converted to [[testosterone]] by enzymatic activity.<ref>{{Cite book | url=https://books.google.com/books?id=F23lAgAAQBAJ&q=androstenedione+pseudopenis%29&pg=PA376 | title=The Mammals of the Southern African Sub-region| isbn=9781107394056| last1=Skinner| first1=J. D.| last2=Chimimba| first2=Christian T.| date=2005-11-15| publisher=Cambridge University Press}}</ref> It is theorized that the dominance and morphological phenotype of a pseudo-penis observed in female hyenas is due to the presence of prenatal and postnatal androstenedione levels. Prenatal [[androgen]] levels dictate genitalia differences between male and female. Higher levels of [[androgen]] are observed in the second half of gestation which is theorized to cause masculinization in terms of dominance and aggression in hyenas.<ref>{{cite journal |last1=French |first1=Jeffrey A. |last2=Mustoe |first2=Aaryn C. |last3=Cavanaugh |first3=Jon |last4=Birnie |first4=Andrew K. |title=The influence of androgenic steroid hormones on female aggression in 'atypical' mammals |journal=Philosophical Transactions of the Royal Society B: Biological Sciences |date=5 December 2013 |volume=368 |issue=1631 |pages=20130084 |doi=10.1098/rstb.2013.0084 |pmid=24167314 |pmc=3826213 }}</ref> Large amounts of androstenedione are produced in hyena ovarian tissues with little aromatase activity allowing the placenta to convert androstenedione to [[testosterone]].<ref name=pmid8391165>{{cite journal |last1=Yalcinkaya |first1=T. |last2=Siiteri |first2=P. |last3=Vigne |first3=J. |last4=Licht |first4=P |last5=Pavgi |first5=S |last6=Frank |first6=L. |last7=Glickman |first7=S. |title=A mechanism for virilization of female spotted hyenas in utero |journal=Science |date=25 June 1993 |volume=260 |issue=5116 |pages=1929–1931 |doi=10.1126/science.8391165 |pmid=8391165 |bibcode = 1993Sci...260.1929Y }}</ref> High concentrations of androgens is theorized to virilize the female hyena genitalia and kill ovarian follicles.<ref name=pmid8391165/>  Postnatal androgen levels dictate growth in genitalia during puberty. Postnatal androgen levels are higher in females than males when they are younger; especially in infancy.<ref name=pmid3472215>{{cite journal |last1=Glickman |first1=S E |last2=Frank |first2=L G |last3=Davidson |first3=J M |last4=Smith |first4=E R |last5=Siiteri |first5=P K |title=Androstenedione may organize or activate sex-reversed traits in female spotted hyenas. |journal=Proceedings of the National Academy of Sciences of the United States of America |date=May 1987 |volume=84 |issue=10 |pages=3444–3447 |doi=10.1073/pnas.84.10.3444 |pmid=3472215 |pmc=304887 |bibcode=1987PNAS...84.3444G |doi-access=free }}</ref> These high levels of androstenedione contribute to aggression and dominance and the masculinization of genitalia during puberty.<ref name=pmid3472215/>
-==See also==
+== See also ==
+*[[Clitoromegaly]]
 *[[Pseudohermaphroditism]]
 *[[Sexual mimicry]]
-*[[Clitoromegaly]]
-*[[Spider monkey#Anatomy and physiology]]
-==References==
+== References ==
-{{reflist|30em}}
+{{reflist}}
 [[Category:Penis]]