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{{Short description|Primary sexual organ of male animals}}
PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!PENIS!!!!!!!
{{Merge from|Human penis|date=November 2024|discuss=Talk:Human penis#Penis}}
{{Other uses}}
{{About|penises in general|the human organ|Human penis}}
{{Redirect2|Penile|Penes|the community|Penile, Louisville|other uses|Penes (disambiguation)}}
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<!--To avoid edit wars, please see relevant discussions on this article's talk page before unilaterally changing the lead picture.-->
{{Infobox anatomy
| Name = Penis
| Latin = penis
| Greek =
| Image = Penis asiatischer Elefant.JPG
| Caption = Penis of an ]
| Precursor = ] (]s)
| System = ], sometimes with the ]
| Artery =
| Vein =
| Nerve =
| Lymph =
}}
A '''penis''' ({{IPAc-en|ˈ|p|iː|n|ɪ|s}}; {{plural form}}: '''penises''' or '''penes''') is a male ] that is used to ] female or ] animals during ].<ref name="SciencesMexico2010">{{cite book|author1=Janet Leonard|author2=Alex Cordoba-Aguilar R|title=The Evolution of Primary Sexual Characters in Animals|url=https://books.google.com/books?id=PgtXj5R6OfMC|access-date=20 July 2013|date=18 June 2010|publisher=Oxford University Press|isbn=978-0-19-971703-3|url-status=live|archive-url=https://web.archive.org/web/20131011093915/http://books.google.com/books?id=PgtXj5R6OfMC&printsec=frontcover#v=onepage&q=penis&f=false|archive-date=11 October 2013}}</ref><ref>{{cite journal|title=Mating behaviour in the sea slug Elysia timida (Opisthobranchia, Sacoglossa): hypodermic injection, sperm transfer and balanced reciprocity|year=2007|pmc=1934903|last1=Schmitt|first1=V.|last2=Anthes|first2=N.|last3=Michiels|first3=N. K.|journal=Frontiers in Zoology|volume=4|page=17|doi=10.1186/1742-9994-4-17|pmid=17610714 |doi-access=free |issn = 1742-9994 }}</ref> Such organs occur in both ]s and ]s, including humans, but not in all male animals.

The term ''penis'' applies to many ]s, but not to all. As an example, the ] of most ]a is the ], a specialized arm, and male spiders use their ]s. Even within the ], there are morphological variants with specific terminology, such as ].

{{TOC limit|5}}

==Etymology==
The word "penis" is taken from the ] word for "]". Some derive that from ] ''*pesnis'', and the Greek word πέος = "penis" from Indo-European ''*pesos''. Prior to the adoption of the Latin word in English, the penis was referred to as a "yard". The ] cites an example of the word ''yard'' used in this sense from 1379,<ref name="oed-yard">{{cite book |last1=Basu |first1=S. C. |title=Male Reproductive Dysfunction |date=2011 |publisher=JP Medical Ltd |isbn=9789350252208 |page=101 |url=https://books.google.com/books?id=BoL2I7yg67cC&q=Simpson,+John;+Weiner,+Edmund,+eds.+(1989).+%22yard,+n.2%22.+Oxford+English+Dictionary&pg=PA101 |language=en}}</ref> and notes that in his ''Physical Dictionary'' of 1684, ] defined the word ''penis'' as "the Yard, made up of two nervous Bodies, the Channel, Nut, Skin, and Fore-skin, etc."<ref name="oed-penis">{{Cite book|title=Oxford English Dictionary|contribution=penis, ''n''.|contribution-url=http://dictionary.oed.com/cgi/entry/50174565|publisher=]|editor-last=Simpson|editor-first=John|editor-link=John Simpson (lexicographer)|isbn=978-0-19-861186-8<!--|edition=Draft revision September 2005|year=2005-->|year=1989|edition=second|title-link=Oxford English Dictionary}}{{Dead link|date=June 2013}}</ref> According to , this term meant (among other senses) "rod" or "bar".

As with nearly any aspect of the body involved in sexual or ] functions, the penis is the subject of many ] words and ]s for it, a particularly common and enduring one being "cock". See ] for a list of alternative words for penis.

The Latin word ''"]"'' (from ] φαλλος) is sometimes used to describe the penis, although "phallus" originally was used to describe ''representations'', pictorial or carved, of the penis.<ref>{{cite web |url=http://www.etymonline.com/index.php?search=penis&searchmode=none |title=Online Etymology Dictionary |publisher=Etymonline.com |access-date=2011-05-28 |url-status=live |archive-url=https://web.archive.org/web/20110606085622/http://www.etymonline.com/index.php?search=penis&searchmode=none |archive-date=2011-06-06 }}</ref>

==Evolution and function==
{{CSS image crop|Image=Tiger_Tadoba_NP.jpg|bSize=880|cWidth=280|cHeight=110|oTop=230|oLeft=560|Description=A ] is aimed backward during urination. Tigers ] their territories with ]s in urine.<ref>{{Cite book |last=Watson |first=Lyall |url=https://books.google.com/books?id=e_KPAgAAQBAJ&pg=PT36 |title=Jacobson's Organ: And the Remarkable Nature of Smell |date=2000-04-17 |publisher=W. W. Norton & Company |isbn=978-0-393-24493-9 |language=en}}</ref>}}
The external genital organs appeared in the ], about 410 million years ago, when ]s began to abandon the aquatic environment.<ref>{{cite book|last1=Dunlop|first1=Jason A.|last2=Penney|first2=David|title=Fossil Arachnids|date=2012 |url=https://books.google.com/books?id=16Zed-dC1OYC&pg=PA44|page=44|publisher=Siri Scientific Press|isbn=978-0-95677-954-0}}</ref> In fact, the necessity to overcome the absence of a liquid phase in which to release the gametes was achieved through the transition to ].

Among amniotes, the development of an erectile penis occurred independently for ]s, ] (]s and ]), ] (turtles), and ]s (] and ]).

Over time, birds have lost this organ, with the exception of ] and ].<ref>{{cite book|last1=Cordoba-Aguilar|first1=Alex|last2=Leonard|first2=Janet|title=The Evolution of Primary Sexual Characters in Animals|year=2010|pages=216–221|publisher=Oxford University Press|url=https://books.google.com/books?id=PgtXj5R6OfMC|isbn=978-0-19971-703-3}}</ref>

The penis is an ] used to transfer ] into the female ] (i.e., ] or ]) for potential ]. The penises of different animal groups are not ] with each other, but were created several times independently of each other in the course of evolution.

An ] is the stiffening and rising of the penis, which occurs during ], though it can also happen in non-sexual situations.
During ], a series of muscular contractions delivers ], containing male ] known as sperm cells or ], from the penis. Ejaculation is usually accompanied by ].

The last common ancestor of all living ]s (mammals, birds and reptiles) likely possessed a penis.<ref>{{cite journal |vauthors=Sanger TJ, Gredler ML, Cohn MJ |date=October 2015 |title=Resurrecting embryos of the tuatara, Sphenodon punctatus, to resolve vertebrate phallus evolution |journal=] |volume=11 |issue=10 |pages=20150694 |doi=10.1098/rsbl.2015.0694 |pmc=4650183 |pmid=26510679}}</ref>

==Vertebrates==
===Birds===
{{anchor|Birds}}
{{multiple image
| total_width = 300px
| image1 = Bird sex - Two copulating ducks showing a corkscrew penis inserted in a cloaca in Don Det Laos.jpg|thumb|Copulating ducks showing a corkscrew ] inserted in a ].
| image2 = Duck Reproduction.svg|thumb
| footer = Male ]s have a corkscrew-shaped penis to match the females' corkscrew vaginas. This favors fertilization by fitter mates over unwanted aggressors.<ref>{{Cite journal |last1=Brennan |first1=Patricia L. R. |last2=Clark |first2=Christopher J. |last3=Prum |first3=Richard O. |date=2010-05-07 |title=Explosive eversion and functional morphology of the duck penis supports sexual conflict in waterfowl genitalia |journal=Proceedings of the Royal Society B: Biological Sciences |volume=277 |issue=1686 |pages=1309–1314 |doi=10.1098/rspb.2009.2139 |issn=0962-8452 |pmc=2871948 |pmid=20031991}}</ref>}}
Most male birds (e.g., ]s and ]s) have a ] (also present on the female), but not a penis. Among bird species with a penis are ] (]s and ])<ref name="Lombardi1998">{{cite book|author=Julian Lombardi|title=Comparative Vertebrate Reproduction|url=https://books.google.com/books?id=cqQX9RMPAegC&q=penis|access-date=5 December 2012|year=1998|publisher=Springer|isbn=978-0-7923-8336-9|url-status=live|archive-url=https://web.archive.org/web/20140326171659/http://books.google.com/books?id=cqQX9RMPAegC&q=penis&f=false|archive-date=26 March 2014}}</ref> and ] (ducks, geese and swans).<ref name="MobileReference2009">{{cite book|author=MobileReference|title=The Illustrated Encyclopedia of European Birds: An Essential Guide to Birds of Europe|url=https://books.google.com/books?id=RgPTUkhiSmkC&q=penis|archive-url=https://web.archive.org/web/20140326174050/http://books.google.com/books?id=RgPTUkhiSmkC&q=penis&f=false|url-status=dead|archive-date=26 March 2014|access-date=5 December 2012|date=15 December 2009|publisher=MobileReference|isbn=978-1-60501-557-6}}</ref> The ] in the family ] also has a penis. A bird penis is different in structure from mammal penises, being an erectile expansion of the cloacal wall (in ducks) and being erected by ], not blood.<ref name="Gill2006">{{cite book|author=Frank B. Gill|title=Ornithology|url=https://books.google.com/books?id=zM0tG5ApO0UC&q=penis&pg=PA414|access-date=5 December 2012|date=6 October 2006|publisher=Macmillan|isbn=978-0-7167-4983-7|pages=414–|url-status=live|archive-url=https://web.archive.org/web/20140107223055/http://books.google.com/books?id=zM0tG5ApO0UC&pg=PA414&q=penis&f=false|archive-date=7 January 2014}}</ref> It is usually partially feathered and in some species features spines and brush-like filaments, and in a flaccid state, curls up inside the cloaca.

=== Mammals ===
{{anchor|Mammals}}

{{multiple image
| perrow = 2/1/2
| total_width = 300px
| image1 = Papio_hamadryas-adult_penis-Lisbon_07.JPG
| caption1 = Penis of a ]
| image2 = Caloocan Annual Tilbury Race 45.jpg
| caption2 = Penis of a ]
| image3 = Penis-cat.jpg
| caption3 = Penis of a ]
| image4 = Great Dane penis.jpg
| caption4 = Penis of a ] (])
| image5 = Retreat (5543779030).jpg
| caption5 = Penis of a ]
}}

As with any other bodily attribute, the length and girth of the penis can be highly variable between ]s of different ].<ref>{{cite book|author=Tim Birkhead|title=Promiscuity: An Evolutionary History of Sperm Competition|url=https://books.google.com/books?id=3i4Q8SvohfEC&pg=PA102|access-date=23 July 2013|year=2000|publisher=Harvard University Press|isbn=978-0-674-00666-9|page=102|url-status=live|archive-url=https://web.archive.org/web/20140326175533/http://books.google.com/books?id=3i4Q8SvohfEC&pg=PA102#v=onepage&q=penis&f=false|archive-date=26 March 2014}}</ref><ref name="HayssenTienhoven1993">{{cite book|author1=Virginia Douglass Hayssen|author2=Ari Van Tienhoven|title=Asdell's Patterns of Mammalian Reproduction: A Compendium of Species-Specific Data|url=https://archive.org/details/asdellspatternso00hays|url-access=registration|access-date=23 July 2013|year=1993|publisher=Cornell University Press|isbn=978-0-8014-1753-5}}</ref> In many mammals, the penis is retracted into a ] when not erect. Mammals have either ''musculocavernous'' penises, which expand while erect, or ''fibroelastic'' penises, which become erect by straightening without expanding.<ref>{{Cite book |last1=Chenoweth |first1=Peter J. |url=https://books.google.com/books?id=3JtcEAAAQBAJ&dq=musculocavernous+fibroelastic+penis&pg=PA16 |title=Manual of Animal Andrology |last2=Lorton |first2=Steven P. |date=2022-02-03 |publisher=CABI |isbn=978-1-78924-350-5 |language=en}}</ref> ]s are present in some prepuces. In ]s, the ], which is connected to the ], travels through and exits the penis, thus both ] and semen are expelled from this organ.<ref name="Wake1992">{{cite book|author=Marvalee H. Wake|title=Hyman's Comparative Vertebrate Anatomy|url=https://books.google.com/books?id=VKlWjdOkiMwC&pg=PA583|access-date=23 July 2013|date=15 September 1992|publisher=University of Chicago Press|isbn=978-0-226-87013-7|page=583|url-status=live|archive-url=https://web.archive.org/web/20131231143932/http://books.google.com/books?id=VKlWjdOkiMwC&pg=PA583#v=onepage&q=penis&f=false|archive-date=31 December 2013}}</ref><ref>{{Cite book |last=Feldhamer |first=George A. |url=https://books.google.com/books?id=udCnKce9hfoC&pg=PA194 |title=Mammalogy: Adaptation, Diversity, Ecology |date=2007-09-07 |publisher=JHU Press |isbn=978-0-8018-8695-9 |language=en}}</ref> The ] of ] mammals (mammals without a ]) separates the ] and the penis.

A bone called the ] is present in most placentals but absent in humans, cattle and horses.

In mammals, the penis is divided into three parts:<ref name="Reece">{{cite book|title=Functional Anatomy and Physiology of Domestic Animals|url=https://books.google.com/books?id=naSWWxJLcd0C&q=penis%20OR%20glans|date=2009-03-04|publisher=John Wiley & Sons|isbn=9780813814513|author=William O. Reece|url-status=live|archive-url=https://web.archive.org/web/20180320145847/https://books.google.com/books?id=naSWWxJLcd0C&q=penis%20OR%20glans&f=false|archive-date=2018-03-20}}</ref>

* Roots (]): these begin at the ] border of the ].
* ]: the part of the penis extending from the roots.
* ]: the free end of the penis and where the urethra opens into in placentals. The penile glans is absent in ]s and ]s.<ref>{{cite book|last=Dixson|first=Alan F.|title= Sexual Selection and the Origins of Human Mating Systems|page=68|url=https://books.google.com/books?id=VRTniKE2liYC&pg=PA68|year=2009|publisher=John OUP Oxford|isbn=978-0-19156-973-9}}</ref>

The internal structures of the penis consist mainly of cavernous, ], which is a collection of ] separated by sheets of ] (trabeculae).

]es have a structure at the base called the ].<ref name="Long2006">{{cite book|author=Susan Long|title=Veterinary Genetics and Reproductive Physiology|url=https://books.google.com/books?id=Gre0VUZuhGYC|year=2006|publisher=Churchill Livingstone Elsevier|isbn=978-0-7506-8877-2|access-date=2013-11-08|url-status=live|archive-url=https://web.archive.org/web/20140326180538/http://books.google.com/books?id=Gre0VUZuhGYC&printsec=frontcover#v=onepage&q=bulbus%20glandis&f=false|archive-date=2014-03-26}}</ref><ref name=autogenerated3>{{cite book|author=R. F. Ewer|title=The Carnivores|url=https://books.google.com/books?id=IETMd3-lSlkC&pg=PA116|access-date=23 July 2013|year=1998|publisher=Cornell University Press|isbn=978-0-8014-8493-3|page=116|url-status=live|archive-url=https://web.archive.org/web/20140326181627/http://books.google.com/books?id=IETMd3-lSlkC&pg=PA116#v=onepage&q=penis&f=false|archive-date=26 March 2014}}</ref> During copulation, the ] inserts his penis through ] instead of directly through the ], which is blocked by the false scrotum. The ] and pseudo-scrotum, which are actually a ] ], closely resemble the male hyena's genitalia, but can be distinguished from the male by the female's greater thickness and more rounded ].<ref name="differentiation">{{cite journal | last1 = Glickman | first1 = SE | last2 = Cunha | first2 = GR | last3 = Drea | first3 = CM | last4 = Conley | first4 = AJ | last5 = Place | first5 = NJ | year = 2006 | title = Mammalian sexual differentiation: lessons from the spotted hyena | url = http://courses.washington.edu/pbio509/Glickman_etal.pdf | journal = Trends Endocrinol Metab | volume = 17 | issue = 9| pages = 349–356 | doi = 10.1016/j.tem.2006.09.005 | pmid = 17010637 | s2cid = 18227659 | url-status = live | archive-url = https://web.archive.org/web/20130222022824/http://courses.washington.edu/pbio509/Glickman_etal.pdf | archive-date = 2013-02-22 }}</ref> {{anchor|Felidae}}] have barbed penises, with about 120–150 one millimetre long ].<ref>{{Cite journal |last1=Aronson |first1=L. R. |last2=Cooper |first2=M. L. |title=Penile spines of the domestic cat: their endocrine-behavior relations |journal=Anat. Rec. |volume=157 |issue=1 |pages=71–8 |year=1967 |pmid=6030760 |doi=10.1002/ar.1091570111 |s2cid=13070242 |url=http://www.catcollection.org/files/PenileSpines.pdf |url-status=dead |archive-url=https://web.archive.org/web/20170620212227/http://www.catcollection.org/files/PenileSpines.pdf |archive-date=2017-06-20 }}</ref>

]s usually have bifurcated penises<ref>{{Cite book |last1=Tyndale-Biscoe |first1=C. Hugh |url=https://books.google.com/books?id=HpjovN0vXW4C |title=Reproductive Physiology of Marsupials |last2=Renfree |first2=Marilyn |date=1987-01-30 |publisher=Cambridge University Press |isbn=978-0-521-33792-2 |language=en}}</ref> that are retracted into a preputial sheath in the male's ] when not erect.<ref>{{Cite book |last1=Armati |first1=Patricia J. |url=https://books.google.com/books?id=x3S5v971Nk0C |title=Marsupials |last2=Dickman |first2=Chris R. |last3=Hume |first3=Ian D. |date=2006-08-17 |publisher=Cambridge University Press |isbn=978-1-139-45742-2 |language=en}}</ref> ]s and ]s are the only mammals in which the penis is located inside the cloaca.<ref>Gadow, H. On the systematic position of Notoryctes typhlops. Proc. Zool. Soc. London 1892, 361–370 (1892).</ref><ref>Riedelsheimer, B., Unterberger, P., Künzle, H. and U. Welsch. 2007. Histological study of the cloacal region and associated structures in the hedgehog tenrec Echinops telfairi. Mammalian Biology 72(6): 330-341.</ref>

===Reptiles===
]]]
Male ]s and ]ns have a penis, while male specimens of the reptile order ], which are ]s and ]s, have two paired organs called ]. ]s must use their cloacae for reproduction.<ref>Lutz, Dick (2005), Tuatara: A Living Fossil, Salem, Oregon: DIMI PRESS, {{ISBN|0-931625-43-2}}</ref> Due to ], turtle and mammal penises have a similar structure.<ref>{{cite journal | pmc=1810052 | year=2004 | last1=Kelly | first1=D. A. | title=Turtle and mammal penis designs are anatomically convergent | journal=Proceedings. Biological Sciences | volume=271 | issue=Suppl 5 | pages=S293–S295 | doi=10.1098/rsbl.2004.0161 | pmid=15503998 }}</ref>

===Fish===
In some fish, the gonopodium, andropodium, and ]s are intromittent organs (to introduce sperm into the female) developed from modified fins.<ref>{{Cite book |last=Ruschenberger |first=William Samuel Waithman |url=https://books.google.com/books?id=700AAAAAYAAJ |title=Elements of Herpetology, and of Ichthyology: Prepared for the Use of Schools and Colleges |date=1846 |publisher=Grigg & Elliot |pages=129–145 |language=en}}</ref>

==Invertebrates==
{{Redirect|Female penis|the homologue to the penis in female amniotes|Clitoris}}
]]]

] are the only male ]s that ].

In male ]s, the structure analogous to a penis is known as an ]. The male copulatory organ of various lower invertebrate animals is often called the ''cirrus''.<ref name="Britannica">{{Cite encyclopedia |title=Penis {{!}} Description, Anatomy, & Physiology {{!}} Britannica |encyclopedia=Encyclopædia Britannica |date=January 2024 |url=https://www.britannica.com/science/penis |language=en}}</ref>

In 2010, entomologist Charles Linehard described a new genus of ] called '']''. Species of this genus have sex-reversed genitalia: females have penis-like organs called gynosomes that are inserted into vagina-like openings of males during mating.<ref name="Lienhard2010">{{cite journal|first1=Charles|last1=Lienhard|first2=Thais|last2=Oliveira do Carmo|first3=Rodrigo|last3=Lopes Ferreira|year=2010|url=http://biostor.org/reference/134593|title=A new genus of Sensitibillini from Brazilian caves (Psocodea: 'Psocoptera': Prionoglarididae)|journal=]|volume=117|issue=4|pages=611–635|issn=0035-418X|url-status=live|archive-url=https://web.archive.org/web/20141103042552/http://biostor.org/reference/134593|archive-date=2014-11-03|doi=10.5962/bhl.part.117600|doi-access=free}}</ref> A similar female structure has also been described in the closely related '']''.<ref name="Yoshizawa_et.al.2018">Yoshizawa K, Ferreira R.L., Yao I, Lienhard C & Kamimura Y. "Independent origins of female penis and its coevolution with male vagina in cave insects (Psocodea: Prionoglarididae)". ''Biology Letters'' '''14'''(11): </ref> Scientists who study these insects have occasionally called the gynosome a "female penis"<ref name="YoshizawaFerreira2014">{{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=]|volume=24|issue=9|pages=1006–10|year=2014|issn=0960-9822|doi=10.1016/j.cub.2014.03.022|pmid=24746797|doi-access=free|bibcode=2014CBio...24.1006Y |hdl=2115/56857|hdl-access=free}}</ref><ref name="Yoshizawa _et.al.2019">Yoshizawa K, Ferreira R.L., Lienhard C & Kamimura Y. (2019). "Why Did a Female Penis Evolve in a Small Group of Cave Insects?". ''BioEssays'' '''41'''(6): </ref> and insisted to drop the definition of penis as "the male copulatory organ".<ref name="Yoshizawa_et.al._2014">Yoshizawa K, Ferreira R.L., Kamimura Y & Lienhard C. . ''The Winnower'' 3/9/2014</ref> Motivations for using the term "female penis" include that such a term "is easier to understand and much more eye-catching"<ref name="Hollier&Hollier_2020">Hollier J & Hollier A. (2020). . ''Antenna'' '''44'''(3): p. 122-125</ref> and that the gynosome have "analogous features" with male penises.<ref name="Yoshizawa_et.al._2014"></ref> Meanwhile, critics have argued that it does not fit the intromittent organ definition of "a structure that enters the female genital tract and deposits sperm".<ref name="Hayssen2020">Hayssen V. (2020). "Misconceptions about Conception and Other Fallacies: Historical Bias in Reproductive Biology". ''Integrative and Comparative Biology'' '''60'''(3): p. 683-791: </ref>

==Heraldry==
{{Main|Pizzle}}
Pizzles are represented in ], where the adjective ''pizzled'' (or ''vilené''<ref name="rietstap-1884">{{Cite journal |last=Rietstap |first=J. B. |title=Armorial général; précédé d'un Dictionnaire des termes du blason |year=1884 |page=XXXI |publisher=G. B. van Goor zonen |quote=Vilené: se dit un animal qui a la marque du sexe d'un autre émail que le corps |url= https://archive.org/details/armorialgnralpr00rollgoog }}</ref>) indicates that part of an animate ]'s anatomy, especially if coloured differently.

==See also==
{{Div col|colwidth=15em}}
* ]
* ]
* ]
* ]
* ]
* ]
{{Div col end}}

==References==
=== Citations ===
{{reflist|colwidth=30em}}

=== General and cited references ===
====Horses====
{{refbegin|30em}}
*{{cite book|first1=Donald F.|last1=Walker|first2=John T.|last2=Vaughan|title=Bovine and equine urogenital surgery|url=https://books.google.com/books?id=dpo1VZBIORUC|access-date=23 July 2013|date=1 June 1980|publisher=]|isbn=978-0-8121-0284-0}}
*{{cite web|title=The Stallion: Breeding Soundness Examination & Reproductive Anatomy|publisher=University of Wisconsin-Madison|url=http://www.wisc.edu/ansci_repro/101equinelab/reproduction/stallion_exam.html|access-date=7 July 2007|archive-url=https://web.archive.org/web/20070716140038/http://www.wisc.edu/ansci_repro/101equinelab/reproduction/stallion_exam.html|archive-date=2007-07-16}}
*{{cite book|last1=Munroe|first1=Graham|last2=Weese|first2=Scott|title=Equine Clinical Medicine, Surgery and Reproduction|url=https://books.google.com/books?id=Cu-XUjVS85QC|access-date=18 February 2014|date=15 March 2011|publisher=Manson Publishing|isbn=978-1-84076-608-0}}
*{{cite book|first1=Klaus Dieter|last1=Budras|first12=W. O.|last2=Sack|title=Anatomy of the Horse|url=https://books.google.com/books?id=2FjNASBMUqgC|access-date=1 July 2013|date=1 March 2012|publisher=Manson Publishing|isbn=978-3-8426-8368-6}}
*{{cite book|last=England|first=Gary|title=Fertility and Obstetrics in the Horse|url=https://books.google.com/books?id=A-R2ZhT_oHQC|access-date=18 February 2014|date=15 April 2008|publisher=John Wiley & Sons|isbn=978-0-470-75041-4}}
*{{cite book|author=Equine Research|title=Horse Conformation: Structure, Soundness, and Performance|url=https://books.google.com/books?id=krD1bAEqypcC|archive-url=https://web.archive.org/web/20140707070110/http://books.google.com/books?id=krD1bAEqypcC|url-status=dead|archive-date=7 July 2014|access-date=23 July 2013|year=2004|publisher=Lyons Press|isbn=978-1-59228-487-0}}
*{{cite book|first=James Warren|last=Evans|title=The Horse|url=https://archive.org/details/isbn_9780716718116|url-access=registration|access-date=23 July 2013|date=15 February 1990|publisher=W. H. Freeman|isbn=978-0-7167-1811-6}}
*{{cite book|first1=M. Horace|last1=Hayes|first2=Peter D.|last2=Rossdale|title=Veterinary Notes for Horse Owners: An Illustrated Manual of Horse Medicine and Surgery|url=https://books.google.com/books?id=LHlwaJnOVCEC&q=penis|access-date=1 July 2013|date=March 1988|publisher=Simon & Schuster|isbn=978-0-671-76561-3}}
*{{cite book|last=McBane|first=Susan|title=Modern Horse Breeding: A Guide for Owners|url=https://books.google.com/books?id=BBoMV-CIHpYC|archive-url=https://web.archive.org/web/20140707073433/http://books.google.com/books?id=BBoMV-CIHpYC|url-status=dead|archive-date=7 July 2014|access-date=18 February 2014|year=2001|publisher=Globe Pequot Press|isbn=978-1-58574-389-6}}
{{refend}}

====Marsupials====
{{refbegin|30em}}
*{{cite book|last=Parker|first=Rick|title=Equine Science|url=https://books.google.com/books?id=cCt9EKwu9r0C|access-date=18 February 2014|edition=4|date=13 January 2012|publisher=Cengage Learning|isbn=978-1-111-13877-6}}
*{{cite book|last=Flannery|first=Tim|title=Chasing Kangaroos: A Continent, a Scientist, and a Search for the World's Most Extraordinary Creature|url=https://books.google.com/books?id=ep-PUlwyxDMC|access-date=5 May 2013|year=2008|publisher=Grove/Atlantic, Incorporated|isbn=9780802143716|pages=60–}}
*{{cite book|last=Hunsaker|first=Don II|title=The Biology of Marsupials|url=https://books.google.com/books?id=cESCLrRJGm0C|access-date=18 February 2014|date=2 December 2012|publisher=Elsevier Science|isbn=978-0-323-14620-3}}
*{{cite book|last1=Jones|first1=Menna E.|last2=Dickman|first2=Chris R.|last3=Archer|first3=Mike|first4=Michael|last4=Archer|title=Predators With Pouches: The Biology of Carnivorous Marsupials|url=https://books.google.com/books?id=3YQSDiWHfD0C|access-date=5 May 2013|year=2003|publisher=Csiro Publishing|isbn=9780643066342}}
*{{cite web|url=http://www.biology.iastate.edu/InternationalTrips/1Australia/Australia%20papers/Discoveries%20about%20Marsupial%20Rep |publisher=Iowa State University Biology Dept. |title=Discoveries about Marsupial Reproduction |first=Anna |last=King |year=2001 |access-date=2012-11-22 |url-status=dead |archive-url=https://web.archive.org/web/20120905123347/http://www.biology.iastate.edu/InternationalTrips/1Australia/Australia%20papers/Discoveries%20about%20Marsupial%20Rep |archive-date=September 5, 2012 }}
*{{cite book|first1=Bernard|last1=Stonehouse|first2=Desmond|last2=Gilmore|title=The Biology of marsupials|url=https://books.google.com/books?id=l63wAAAAMAAJ&q=penis|access-date=25 July 2013|year=1977|publisher=University Park Press|isbn=978-0-8391-0852-8}}
*{{cite book|last=Tyndale-Biscoe|first=C. Hugh|title=Life of Marsupials|url=https://books.google.com/books?id=KqtlPZJ9y8EC|access-date=18 February 2014|year=2005|publisher=Csiro Publishing|isbn=978-0-643-06257-3}}
{{refend}}

====Other animals====
{{refbegin|30em}}
*{{cite book|first1=Colin Russell|last1=Austin|first2=Roger Valentine|last2=Short|title=Reproduction in Mammals: Volume 4, Reproductive Fitness|url=https://books.google.com/books?id=P5Al0X60UAIC|access-date=22 July 2013|date=21 March 1985|publisher=Cambridge University Press|isbn=978-0-521-31984-3}}
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* {{cite journal |last1=Bates |first1=Paul J. J. |last2=Ratrimomanarivo |first2=Fanja H. |last3=Harrison |first3=David L. |last4=Goodman |first4=Steven M. |title=A description of a new species of Pipistrellus (Chiroptera: Vespertilionidae) from Madagascar with a review of related Vespertilioninae from the island |journal=Acta Chiropterologica |date=December 2006 |volume=8 |issue=2 |pages=299–324 |doi=10.3161/1733-5329(2006)82.0.CO;2|s2cid=85825521 }}
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*{{cite book|first=Eugene|last=Burns|title=The sex life of wild animals: a North American study|url=https://books.google.com/books?id=2mFBAAAAYAAJ|access-date=23 July 2013|year=1953|publisher=Rinehart}}
*{{cite book|first=Trevor|last=Carnaby|title=Beat About the Bush: Mammals|url=https://books.google.com/books?id=4u-VroUwC6QC|access-date=19 July 2013|date=22 January 2007|publisher=Jacana Media|isbn=978-1-77009-240-2}}
*{{Cite journal|last=Brehm|first=Alfred Edmund|url=https://archive.org/details/brehmslifeofanim00breh|title=Brehm's Life of Animals|publisher=Chicago: A. N. Marquis & Company|year=1895|access-date=2013-11-08}}
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*{{cite book|first=Stewart Keith|last=Eltringham|title=The ecology and conservation of large African mammals|url=https://books.google.com/books?ei=yAHrUe-DNtOkyAGQkIGQDw|access-date=20 July 2013|year=1979|publisher=Macmillan|isbn=978-0-333-23580-5}}
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*{{cite book|first1=Rowen D.|last1=Frandson|first2=W. Lee|last2=Wilke|first3=Anna Dee|last3=Fails|title=Anatomy and Physiology of Farm Animals|url=https://books.google.com/books?id=I9ZZkwnFLN0C|access-date=1 July 2013|date=30 June 2009|publisher=John Wiley & Sons|isbn=978-0-8138-1394-3}}
*{{cite book|first=Valerius|last=Geist|title=Elk Country|url=https://books.google.com/books?id=b0K9OG6znDgC|access-date=5 July 2013|year=1993|publisher=T&N Children's Publishing|isbn=978-1-55971-208-8}}
*{{cite book|first1=Virginia Douglass|last1=Hayssen|first2=Ari Van|last2=Tienhoven|title=Asdell's Patterns of Mammalian Reproduction: A Compendium of Species-Specific Data|url=https://archive.org/details/asdellspatternso00hays|url-access=registration|access-date=23 July 2013|year=1993|publisher=Cornell University Press|isbn=978-0-8014-1753-5}}
*{{Cite book|last1=Heptner|first1=V. G.|last2=Sludskii|first2=A. A.|url=https://archive.org/details/mammalsofsov212001gept|title=Mammals of the Soviet Union. Vol. II, part 1b, Carnivores (Mustelidae and Procyonidae)|publisher=Washington, D.C. : Smithsonian Institution Libraries and National Science Foundation|year=2002|isbn=978-90-04-08876-4|access-date=2013-11-08}}
*{{cite book|first=Donald F.|last=Hoffmeister|title=Mammals of Illinois|url=https://books.google.com/books?id=IH4iv6MrrW4C|access-date=22 July 2013|year=2002|publisher=University of Illinois Press|isbn=978-0-252-07083-9}}
* {{cite journal |last1=Hooper |first1=E.T. |last2=Musser |first2=G.G. |year=1964 |url=http://hdl.handle.net/2027.42/56367 |title=The glans penis in Neotropical cricetines (Family Muridae) with comments on classification of muroid rodents |journal=Miscellaneous Publications of the University of Michigan Museum of Zoology |volume=123 |pages=1–57|hdl=2027.42/56367 }}
*{{cite book|first1=Barbara N.|last1=Horowitz|first2=Kathryn|last2=Bowers|title=Zoobiquity: What Animals Can Teach Us About Being Human|url=https://books.google.com/books?id=sQraNhrbX2IC|access-date=25 July 2013|date=12 June 2012|publisher=Doubleday Canada|isbn=978-0-385-67061-6}}
*{{cite book|first=Robert H.|last=Horwich|title=The ontogeny of social behavior in the gray squirrel (Sciurus carolinensis)|url=https://books.google.com/books?id=-nMeAQAAMAAJ|access-date=23 July 2013|date=June 1972|publisher=P. Parey|isbn=978-3-489-68036-9}}
*{{cite book|first1=Hartley H.|last1=Jackson|title=Mammals of Wisconsin|url=https://books.google.com/books?id=qQfigtpJ11gC|access-date=22 July 2013|date=January 1961|publisher=Univ of Wisconsin Press|isbn=978-0-299-02150-4}}
*{{cite book|title=Journal of the Mammalogical Society of Japan|url=https://books.google.com/books?id=w91KAAAAYAAJ|access-date=5 July 2013|year=1986|publisher=The Society}}
*{{cite book|first1=Dev Raj|last1=Khanna|first2=P. R.|last2=Yadav|title=Biology Of Mammals|url=https://books.google.com/books?id=4snvlKZpPecC|access-date=20 July 2013|date=1 January 2005|publisher=Discovery Publishing House|isbn=978-81-7141-934-0}}
*{{cite book|first=Jonathan|last=Kingdon|title=East African Mammals: An Atlas of Evolution in Africa. Vol. I|url=https://books.google.com/books?id=x75kYjINSqUC|access-date=22 July 2013|date=January 1984|publisher=University of Chicago Press|isbn=978-0-226-43718-7}}
*{{cite book |last1=Kingdon |first1=Jonathan |title=East African Mammals: An Atlas of Evolution in Africa |date=1984 |publisher=University of Chicago Press |isbn=9780226437187 |url=https://books.google.com/books?id=x75kYjINSqUC |language=en}}
*{{cite book|first1=Horst Erich|last1=König|first2=Hans-Georg|last2=Liebich|title=Veterinary Anatomy of Domestic Mammals: Textbook and Atlas|url=https://books.google.com/books?id=QoXiBjSp368C|access-date=23 July 2013|year=2007|publisher=Schattauer Verlag|isbn=978-3-7945-2485-3}}
*{{cite book|first=R. L.|last=Kotpal|title=Modern Text Book Of Zoology Vertebrates|url=https://books.google.com/books?id=U7N1j-8LMsEC|access-date=23 July 2013|year=2010|publisher=Rastogi Publications|isbn=978-81-7133-891-7}}
*{{cite book|first=William J.|last=Krause|title=An Atlas of Opossum Organogenesis|url=https://books.google.com/books?id=fMpj9XNCss8C|access-date=20 July 2013|date=1 March 2008|publisher=Universal-Publishers|isbn=978-1-58112-969-4}}
*{{cite book|first=Donald W.|last=Linzey|title=Vertebrate Biology|url=https://books.google.com/books?id=qpQ9y-vXovoC|access-date=20 July 2013|date=28 December 2011|publisher=JHU Press|isbn=978-1-4214-0040-2}}
*{{cite book|first1=Steven D.|last1=Lukefahr|first2=Peter R.|last2=Cheeke|first3=Nephi M.|last3=Patton|title=Rabbit Production|url=https://books.google.com/books?id=ZY-8F9MwBWsC|access-date=20 July 2013|year=2013|publisher=CABI|isbn=978-1-78064-012-9}}
*{{cite book|title=Natural History Bulletin of the Siam Society|url=https://books.google.com/books?id=iWRMAQAAIAAJ|access-date=5 July 2013|year=1975}}
* {{Cite journal | doi = 10.1111/j.1096-3642.2010.00643.x | last1 = Percequillo | first1 = A.R. | last2 = Weksler | first2 = M. | last3 = Costa | first3 = L.P. | title = A new genus and species of rodent from the Brazilian Atlantic Forest (Rodentia: Cricetidae: Sigmodontinae: Oryzomyini), with comments on oryzomyine biogeography | journal = Zoological Journal of the Linnean Society | volume = 161 | issue = 2 | pages = 357–390 | year = 2011| doi-access = free }}
*{{cite book|first1=Kenneth D.|last1=Rose|first2=J. David|last2=Archibald|title=The Rise of Placental Mammals: Origins and Relationships of the Major Extant Clades|url=https://books.google.com/books?id=DhchVG_rbQ8C|access-date=22 July 2013|date=22 February 2005|publisher=JHU Press|isbn=978-0-8018-8022-3}}
*{{cite book|first=Uldis|last=Roze|title=The North American Porcupine|url=https://books.google.com/books?id=A3HuW_DMglQC|access-date=25 July 2013|year=2009|publisher=Cornell University Press|isbn=978-0-8014-4646-7}}
*{{cite book|first=Amita|last=Sarkar|title=Sexual Behaviour In Animals|url=https://books.google.com/books?id=bsCiWUiPY5UC|access-date=20 July 2013|date=1 January 2003|publisher=Discovery Publishing House|isbn=978-81-7141-746-9}}
*{{cite book|first1=Heide|last1=Schatten|first2=Gheorghe M.|last2=Constantinescu|title=Comparative Reproductive Biology|url=https://books.google.com/books?id=6iNdSk7gPf4C|access-date=23 July 2013|date=21 March 2008|publisher=John Wiley & Sons|isbn=978-0-470-39025-2}}
*{{cite book|first=Meredith F.|last=Small|title=Female Choices: Sexual Behavior of Female Primates|url=https://archive.org/details/femalechoicessex0000smal|url-access=registration|access-date=23 July 2013|year=1993|publisher=Cornell University Press|isbn=978-0-8014-8305-9}}
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*{{cite journal |last=Weksler |first=M. |year=2006 |url=http://hdl.handle.net/2246/5777 |title=Phylogenetic relationships of oryzomyine rodents (Muroidea: Sigmodontinae): separate and combined analyses of morphological and molecular data |journal=Bulletin of the American Museum of Natural History |volume=296 |pages=1–149|doi=10.1206/0003-0090(2006)2962.0.CO;2 |hdl=2246/5777 |s2cid=86057173 }}
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{{refend}}

==External links==
{{Wiktionary|penis}}
{{Commons|Penis}}
{{Commons category|Male genitalia in heraldry}}
*

{{Authority control}}

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Latest revision as of 22:19, 30 December 2024

Primary sexual organ of male animals
It has been suggested that Human penis be merged into this article. (Discuss) Proposed since November 2024.
For other uses, see Penis (disambiguation). This article is about penises in general. For the human organ, see Human penis. "Penile" and "Penes" redirect here. For the community, see Penile, Louisville. For other uses, see Penes (disambiguation).

Penis
Penis of an Asian elephant
Details
PrecursorGenital tubercle (amniotes)
SystemReproductive system, sometimes with the genitourinary system
Identifiers
Latinpenis
Anatomical terminology[edit on Wikidata]

A penis (/ˈpiːnɪs/; pl.: penises or penes) is a male sex organ that is used to inseminate female or hermaphrodite animals during copulation. Such organs occur in both vertebrates and invertebrates, including humans, but not in all male animals.

The term penis applies to many intromittent organs, but not to all. As an example, the intromittent organ of most Cephalopoda is the hectocotylus, a specialized arm, and male spiders use their pedipalps. Even within the Vertebrata, there are morphological variants with specific terminology, such as hemipenes.

Etymology

The word "penis" is taken from the Latin word for "tail". Some derive that from Indo-European *pesnis, and the Greek word πέος = "penis" from Indo-European *pesos. Prior to the adoption of the Latin word in English, the penis was referred to as a "yard". The Oxford English Dictionary cites an example of the word yard used in this sense from 1379, and notes that in his Physical Dictionary of 1684, Steven Blankaart defined the word penis as "the Yard, made up of two nervous Bodies, the Channel, Nut, Skin, and Fore-skin, etc." According to Wiktionary, this term meant (among other senses) "rod" or "bar".

As with nearly any aspect of the body involved in sexual or excretory functions, the penis is the subject of many slang words and euphemisms for it, a particularly common and enduring one being "cock". See WikiSaurus:penis for a list of alternative words for penis.

The Latin word "phallus" (from Greek φαλλος) is sometimes used to describe the penis, although "phallus" originally was used to describe representations, pictorial or carved, of the penis.

Evolution and function

A tiger's penis is aimed backward during urination. Tigers scent-mark their territories with pheromones in urine. A tiger's penis is aimed backward during urination. Tigers scent-mark their territories with pheromones in urine.

The external genital organs appeared in the Devonian, about 410 million years ago, when tetrapods began to abandon the aquatic environment. In fact, the necessity to overcome the absence of a liquid phase in which to release the gametes was achieved through the transition to internal fertilization.

Among amniotes, the development of an erectile penis occurred independently for mammals, squamates (lizards and snakes), testudines (turtles), and archosaurs (crocodiles and birds).

Over time, birds have lost this organ, with the exception of Paleognathae and Anseriformes.

The penis is an intromittent organ used to transfer sperm into the female genital tract (i.e., vagina or cloaca) for potential fertilization. The penises of different animal groups are not homologous with each other, but were created several times independently of each other in the course of evolution.

An erection is the stiffening and rising of the penis, which occurs during sexual arousal, though it can also happen in non-sexual situations. During ejaculation, a series of muscular contractions delivers semen, containing male gametes known as sperm cells or spermatozoa, from the penis. Ejaculation is usually accompanied by orgasm.

The last common ancestor of all living amniotes (mammals, birds and reptiles) likely possessed a penis.

Vertebrates

Birds

Male ducks have a corkscrew-shaped penis to match the females' corkscrew vaginas. This favors fertilization by fitter mates over unwanted aggressors.

Most male birds (e.g., roosters and turkeys) have a cloaca (also present on the female), but not a penis. Among bird species with a penis are paleognaths (tinamous and ratites) and Anatidae (ducks, geese and swans). The magpie goose in the family Anseranatidae also has a penis. A bird penis is different in structure from mammal penises, being an erectile expansion of the cloacal wall (in ducks) and being erected by lymph, not blood. It is usually partially feathered and in some species features spines and brush-like filaments, and in a flaccid state, curls up inside the cloaca.

Mammals

Penis of a hamadryas baboonPenis of a horsePenis of a catPenis of a dog (Great Dane)Penis of a giraffe

As with any other bodily attribute, the length and girth of the penis can be highly variable between mammals of different species. In many mammals, the penis is retracted into a prepuce when not erect. Mammals have either musculocavernous penises, which expand while erect, or fibroelastic penises, which become erect by straightening without expanding. Preputial glands are present in some prepuces. In placentals, the urethra, which is connected to the vasa deferentia, travels through and exits the penis, thus both urine and semen are expelled from this organ. The perineum of testicond mammals (mammals without a scrotum) separates the anus and the penis.

A bone called the baculum is present in most placentals but absent in humans, cattle and horses.

In mammals, the penis is divided into three parts:

The internal structures of the penis consist mainly of cavernous, erectile tissue, which is a collection of blood sinusoids separated by sheets of connective tissue (trabeculae).

Canine penises have a structure at the base called the bulbus glandis. During copulation, the spotted hyena inserts his penis through the female's pseudo-penis instead of directly through the vagina, which is blocked by the false scrotum. The pseudo-penis and pseudo-scrotum, which are actually a masculinized vulva, closely resemble the male hyena's genitalia, but can be distinguished from the male by the female's greater thickness and more rounded glans. Domestic cats have barbed penises, with about 120–150 one millimetre long backwards-pointing spines.

Marsupials usually have bifurcated penises that are retracted into a preputial sheath in the male's urogenital sinus when not erect. Monotremes and marsupial moles are the only mammals in which the penis is located inside the cloaca.

Reptiles

Hemipenes of a gold tegu

Male turtles and crocodilians have a penis, while male specimens of the reptile order Squamata, which are snakes and lizards, have two paired organs called hemipenes. Tuataras must use their cloacae for reproduction. Due to evolutionary convergence, turtle and mammal penises have a similar structure.

Fish

In some fish, the gonopodium, andropodium, and claspers are intromittent organs (to introduce sperm into the female) developed from modified fins.

Invertebrates

"Female penis" redirects here. For the homologue to the penis in female amniotes, see Clitoris.
The spine-covered penis of Callosobruchus analis, a bean weevil

Harvestmen are the only male arachnids that have a penis.

In male insects, the structure analogous to a penis is known as an aedeagus. The male copulatory organ of various lower invertebrate animals is often called the cirrus.

In 2010, entomologist Charles Linehard described a new genus of barkflies called Neotrogla. Species of this genus have sex-reversed genitalia: females have penis-like organs called gynosomes that are inserted into vagina-like openings of males during mating. A similar female structure has also been described in the closely related Afrotrogla. Scientists who study these insects have occasionally called the gynosome a "female penis" and insisted to drop the definition of penis as "the male copulatory organ". Motivations for using the term "female penis" include that such a term "is easier to understand and much more eye-catching" and that the gynosome have "analogous features" with male penises. Meanwhile, critics have argued that it does not fit the intromittent organ definition of "a structure that enters the female genital tract and deposits sperm".

Heraldry

Main article: Pizzle

Pizzles are represented in heraldry, where the adjective pizzled (or vilené) indicates that part of an animate charge's anatomy, especially if coloured differently.

See also

References

Citations

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General and cited references

Horses

Marsupials

Other animals

External links

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