Penis

Penis
Penis
	Penis of anAsian elephant
Details
Precursor	Genital tubercle(amniotes)
System	Reproductive system,sometimes with thegenitourinary system
Identifiers
Latin	penis
	Anatomical terminology [edit on Wikidata]

Apenis(/ˈpiːnɪs/;pl.:penisesorpenes) is a malesex organthat is used toinseminatefemale orhermaphroditeanimals duringcopulation.^[1]^[2]Such organs occur in bothvertebratesandinvertebrates,but not in all male animals.

The termpenisapplies to manyintromittent organs,but not to all. As an example, theintromittent organof mostCephalopodais thehectocotylus,a specialized arm, and male spiders use theirpedipalps.Even within theVertebrata,there are morphological variants with specific terminology, such ashemipenes.

Etymology

The word "penis" is taken from theLatinword for "tail".Some derive that fromIndo-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". TheOxford English Dictionarycites an example of the wordyardused in this sense from 1379,^[3]and notes that in hisPhysical Dictionaryof 1684,Steven Blankaartdefined the wordpenisas "the Yard, made up of two nervous Bodies, the Channel, Nut, Skin, and Fore-skin, etc."^[4]According toWiktionary,this term meant (among other senses) "rod" or "bar".

As with nearly any aspect of the body involved in sexual orexcretoryfunctions, the penis is the subject of manyslangwords andeuphemismsfor it, a particularly common and enduring one being "cock". SeeWikiSaurus:penisfor a list of alternative words for penis.

The Latin word"phallus"(fromGreekφαλλος) is sometimes used to describe the penis, although "phallus" originally was used to describerepresentations,pictorial or carved, of the penis.^[5]

Evolution and function

Atiger's penisis aimed backward during urination. Tigersscent-marktheir territories withpheromonesin urine.^[6]

Dogs mark their territorieswith urine and secretions from thepreputial glandsin front of the penis.^[7]

The external genital organs appeared in theDevonian,about 410 million years ago, whentetrapodsbegan to abandon the aquatic environment.^[8]In fact, the necessity to overcome the absence of a liquid phase in which to release the gametes was achieved through the transition tointernal fertilization.

Among amniotes, the development of an erectile penis occurred independently formammals,squamates(lizardsandsnakes),testudines(turtles), andarchosaurs(crocodilesandbirds).

Over time, birds have lost this organ, with the exception ofPaleognathaeandAnseriformes.^[9]

The penis is anintromittent organused to transfersperminto the femalegenital tract(i.e.,vaginaorcloaca) for potentialfertilizationand, in the case ofplacentals,also for the excretion ofurine.^[10]^[11]The penises of different animal groups are nothomologouswith each other, but were created several times independently of each other in the course of evolution.

Anerectionis the stiffening and rising of the penis, which occurs duringsexual arousal,though it can also happen in non-sexual situations. Duringejaculation,a series of muscular contractions delivers semen, containing malegametesknown as sperm cells orspermatozoa,from the penis. Ejaculation is usually accompanied byorgasm.

The last common ancestor of all livingamniotes(mammals, birds and reptiles) likely possessed a penis.^[12]

Vertebrates

Birds

Maleduckshave a corkscrew-shaped penis while female ducks have corkscrew vaginas with many blind pockets evolved for difficult penetration and to prevent becoming pregnant.^[13]This reduced the likelihood of fertilization by unwanted aggressors in favor of fitter mates.

Most male birds (e.g.,roostersandturkeys) have acloaca(also present on the female), but not a penis. Among bird species with a penis arepaleognaths(tinamousandratites)^[14]andAnatidae(ducks, geese and swans).^[15]Themagpie goosein the familyAnseranatidaealso 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 bylymph,not blood.^[16]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 ahamadryas baboon

Penis of ahorse

Penis of acat

Penis of adog(Great Dane)

Penis of agiraffe

As with any other bodily attribute, the length and girth of the penis can be highly variable betweenmammalsof differentspecies.^[17]^[18]In many mammals, the penis is retracted into aprepucewhen not erect. Mammals have eithermusculocavernouspenises, which expand while erect, orfibroelasticpenises, which become erect by straightening without expanding.^[19]Preputial glandsare present in some prepuces. The penis bearsdistalpart of theurethrainplacental mammals.^[10]Theperineumoftesticondmammals (mammals without ascrotum) separates theanusand the penis.

A bone called thebaculumis present in most placentals but absent in humans, cattle and horses.

In mammals, the penis is divided into three parts:^[20]

Roots (crura): these begin at thecaudalborder of thepelvic ischial arch.
Body:the part of the penis extending from the roots.
Glans:the free end of the penis. The penile glans is absent inchimpanzeesandbonobos.^[21]

The internal structures of the penis consist mainly of cavernous,erectile tissue,which is a collection ofblood sinusoidsseparated by sheets ofconnective tissue(trabeculae).

Canine peniseshave a structure at the base called thebulbus glandis.^[22]^[23]During copulation, thespotted hyenainserts his penis throughthe female's pseudo-penisinstead of directly through thevagina,which is blocked by the false scrotum. Thepseudo-penisand pseudo-scrotum, which are actually amasculinized vulva,closely resemble the male hyena's genitalia, but can be distinguished from the male by the female's greater thickness and more roundedglans.^[24]Domestic catshave barbed penises, with about 120–150 one millimetre longbackwards-pointing spines.^[25]

Marsupialsusually have bifurcated penises^[26]that are retracted into a preputial sheath in the male'surogenital sinuswhen not erect.^[27]Monotremesandmarsupial molesare the only mammals in which the penis is located inside the cloaca.^[28]^[29]

Reptiles

Maleturtlesandcrocodilianshave a penis, while male specimens of the reptile orderSquamata,which aresnakesandlizards,have two paired organs calledhemipenes.Tuatarasmust use their cloacae for reproduction.^[30]Due toevolutionary convergence,turtle and mammal penises have a similar structure.^[31]

Fish

In some fish, the gonopodium, andropodium, andclaspersare intromittent organs (to introduce sperm into the female) developed from modified fins.^[32]

Invertebrates

Harvestmenare the only malearachnidsthathave a penis.

In maleinsects,the structure analogous to a penis is known as anaedeagus.The male copulatory organ of various lower invertebrate animals is often called thecirrus.^[33]

In 2010, entomologist Charles Linehard described a new genus ofbarkfliescalledNeotrogla.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.^[34]A similar female structure has also been described in the closely relatedAfrotrogla.^[35]Scientists who study these insects have occasionally called the gynosome a "female penis"^[36]^[37]and insisted to drop the definition of penis as "the male copulatory organ".^[38]Motivations for using the term “female penis” include that such a term "is easier to understand and much more eye-catching"^[39]and that the gynosome have "analogous features" with male penises.^[38]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".^[40]

Heraldry

Pizzles are represented inheraldry,where the adjectivepizzled(orvilené^[41]) indicates that part of an animatecharge's anatomy, especially if coloured differently.

References

Citations

^Janet Leonard; Alex Cordoba-Aguilar R (18 June 2010).The Evolution of Primary Sexual Characters in Animals.Oxford University Press.ISBN 978-0-19-971703-3.Archivedfrom the original on 11 October 2013.Retrieved20 July2013.
^Schmitt, V.; Anthes, N.; Michiels, N. K. (2007)."Mating behaviour in the sea slug Elysia timida (Opisthobranchia, Sacoglossa): hypodermic injection, sperm transfer and balanced reciprocity".Frontiers in Zoology.4:17.doi:10.1186/1742-9994-4-17.ISSN 1742-9994.PMC1934903.PMID 17610714.
^Basu, S. C. (2011).Male Reproductive Dysfunction.JP Medical Ltd. p. 101.ISBN 9789350252208.
^Simpson, John,ed. (1989)."penis,n.".Oxford English Dictionary(second ed.).Oxford University Press.ISBN 978-0-19-861186-8.^{[dead link]}
^"Online Etymology Dictionary".Etymonline.Archivedfrom the original on 2011-06-06.Retrieved2011-05-28.
^Watson, Lyall (2000-04-17).Jacobson's Organ: And the Remarkable Nature of Smell.W. W. Norton & Company.ISBN 978-0-393-24493-9.
^Mech, L. David;Boitani, Luigi,eds. (2003).Wolves: Behaviour, Ecology and Conservation.University of Chicago Press.ISBN 0-226-51696-2.
^Dunlop, Jason A.; Penney, David (2012).Fossil Arachnids.Siri Scientific Press. p. 44.ISBN 978-0-95677-954-0.
^Cordoba-Aguilar, Alex; Leonard, Janet (2010).The Evolution of Primary Sexual Characters in Animals.Oxford University Press. pp. 216–221.ISBN 978-0-19971-703-3.
^^a ^bMarvalee H. Wake (15 September 1992).Hyman's Comparative Vertebrate Anatomy.University of Chicago Press. p. 583.ISBN 978-0-226-87013-7.Archivedfrom the original on 31 December 2013.Retrieved23 July2013.
^Feldhamer, George A. (2007-09-07).Mammalogy: Adaptation, Diversity, Ecology.JHU Press.ISBN 978-0-8018-8695-9.
^Sanger TJ, Gredler ML, Cohn MJ (October 2015)."Resurrecting embryos of the tuatara, Sphenodon punctatus, to resolve vertebrate phallus evolution".Biology Letters.11(10): 20150694.doi:10.1098/rsbl.2015.0694.PMC4650183.PMID 26510679.
^Brennan, Patricia L. R.; Clark, Christopher J.; Prum, Richard O. (2010-05-07)."Explosive eversion and functional morphology of the duck penis supports sexual conflict in waterfowl genitalia".Proceedings of the Royal Society B: Biological Sciences.277(1686): 1309–1314.doi:10.1098/rspb.2009.2139.ISSN 0962-8452.PMC2871948.PMID 20031991.
^Julian Lombardi (1998).Comparative Vertebrate Reproduction.Springer.ISBN 978-0-7923-8336-9.Archivedfrom the original on 26 March 2014.Retrieved5 December2012.
^MobileReference (15 December 2009).The Illustrated Encyclopedia of European Birds: An Essential Guide to Birds of Europe.MobileReference.ISBN 978-1-60501-557-6.Archived fromthe originalon 26 March 2014.Retrieved5 December2012.
^Frank B. Gill (6 October 2006).Ornithology.Macmillan. pp. 414–.ISBN 978-0-7167-4983-7.Archivedfrom the original on 7 January 2014.Retrieved5 December2012.
^Tim Birkhead (2000).Promiscuity: An Evolutionary History of Sperm Competition.Harvard University Press. p. 102.ISBN 978-0-674-00666-9.Archivedfrom the original on 26 March 2014.Retrieved23 July2013.
^Virginia Douglass Hayssen; Ari Van Tienhoven (1993).Asdell's Patterns of Mammalian Reproduction: A Compendium of Species-Specific Data.Cornell University Press.ISBN 978-0-8014-1753-5.Retrieved23 July2013.
^Chenoweth, Peter J.; Lorton, Steven P. (2022-02-03).Manual of Animal Andrology.CABI.ISBN 978-1-78924-350-5.
^William O. Reece (2009-03-04).Functional Anatomy and Physiology of Domestic Animals.John Wiley & Sons.ISBN 9780813814513.Archivedfrom the original on 2018-03-20.
^Dixson, Alan F. (2009).Sexual Selection and the Origins of Human Mating Systems.John OUP Oxford. p. 68.ISBN 978-0-19156-973-9.
^Susan Long (2006).Veterinary Genetics and Reproductive Physiology.Churchill Livingstone Elsevier.ISBN 978-0-7506-8877-2.Archivedfrom the original on 2014-03-26.Retrieved2013-11-08.
^R. F. Ewer (1998).The Carnivores.Cornell University Press. p. 116.ISBN 978-0-8014-8493-3.Archivedfrom the original on 26 March 2014.Retrieved23 July2013.
^Glickman, SE; Cunha, GR; Drea, CM; Conley, AJ; Place, NJ (2006)."Mammalian sexual differentiation: lessons from the spotted hyena"(PDF).Trends Endocrinol Metab.17(9): 349–356.doi:10.1016/j.tem.2006.09.005.PMID 17010637.S2CID 18227659.Archived(PDF)from the original on 2013-02-22.
^Aronson, L. R.; Cooper, M. L. (1967)."Penile spines of the domestic cat: their endocrine-behavior relations"(PDF).Anat. Rec.157(1): 71–8.doi:10.1002/ar.1091570111.PMID 6030760.S2CID 13070242.Archived fromthe original(PDF)on 2017-06-20.
^Tyndale-Biscoe, C. Hugh; Renfree, Marilyn (1987-01-30).Reproductive Physiology of Marsupials.Cambridge University Press.ISBN 978-0-521-33792-2.
^Armati, Patricia J.; Dickman, Chris R.; Hume, Ian D. (2006-08-17).Marsupials.Cambridge University Press.ISBN 978-1-139-45742-2.
^Gadow, H. On the systematic position of Notoryctes typhlops. Proc. Zool. Soc. London 1892, 361–370 (1892).
^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.
^Lutz, Dick (2005), Tuatara: A Living Fossil, Salem, Oregon: DIMI PRESS,ISBN 0-931625-43-2
^Kelly, D. A. (2004)."Turtle and mammal penis designs are anatomically convergent".Proceedings. Biological Sciences.271(Suppl 5): S293–S295.doi:10.1098/rsbl.2004.0161.PMC1810052.PMID 15503998.
^Ruschenberger, William Samuel Waithman (1846).Elements of Herpetology, and of Ichthyology: Prepared for the Use of Schools and Colleges.Grigg & Elliot. pp. 129–145.
^"Penis | Description, Anatomy, & Physiology | Britannica".Encyclopædia Britannica.January 2024.
^Lienhard, Charles; Oliveira do Carmo, Thais; Lopes Ferreira, Rodrigo (2010)."A new genus of Sensitibillini from Brazilian caves (Psocodea: 'Psocoptera': Prionoglarididae)".Revue suisse de Zoologie.117(4): 611–635.doi:10.5962/bhl.part.117600.ISSN 0035-418X.Archivedfrom the original on 2014-11-03.
^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 Letters14(11):doi.org/10.1098/rsbl.2018.0533
^Yoshizawa, Kazunori; Ferreira, Rodrigo L.; Kamimura, Yoshitaka; Lienhard, Charles (2014)."Female Penis, Male Vagina, and Their Correlated Evolution in a Cave Insect".Current Biology.24(9): 1006–10.doi:10.1016/j.cub.2014.03.022.hdl:2115/56857.ISSN 0960-9822.PMID 24746797.
^Yoshizawa K, Ferreira R.L., Lienhard C & Kamimura Y. (2019). "Why Did a Female Penis Evolve in a Small Group of Cave Insects?".BioEssays41(6):doi.org/10.1002/bies.201900005
^^a ^bYoshizawa K, Ferreira R.L., Kamimura Y & Lienhard C."A Transgender Brazilian Cave Insect".The Winnower3/9/2014
^Hollier J & Hollier A. (2020)."The retired taxonomist and the gynosome – the discovery of the female penis".Antenna44(3): p. 122-125
^Hayssen V. (2020). "Misconceptions about Conception and Other Fallacies: Historical Bias in Reproductive Biology".Integrative and Comparative Biology60(3): p. 683-791:doi.org/10.1093/icb/icaa035
^Rietstap, J. B. (1884)."Armorial général; précédé d'un Dictionnaire des termes du blason".G. B. van Goor zonen: XXXI.Vilené: se dit un animal qui a la marque du sexe d'un autre émail que le corps{{cite journal}}:Cite journal requires|journal=(help)

General and cited references

Horses

Walker, Donald F.; Vaughan, John T. (1 June 1980).Bovine and equine urogenital surgery.Lea & Febiger.ISBN 978-0-8121-0284-0.Retrieved23 July2013.
"The Stallion: Breeding Soundness Examination & Reproductive Anatomy".University of Wisconsin-Madison. Archived fromthe originalon 2007-07-16.Retrieved7 July2007.
Munroe, Graham; Weese, Scott (15 March 2011).Equine Clinical Medicine, Surgery and Reproduction.Manson Publishing.ISBN 978-1-84076-608-0.Retrieved18 February2014.
Budras, Klaus Dieter; Sack (1 March 2012).Anatomy of the Horse.Manson Publishing.ISBN 978-3-8426-8368-6.Retrieved1 July2013.
England, Gary (15 April 2008).Fertility and Obstetrics in the Horse.John Wiley & Sons.ISBN 978-0-470-75041-4.Retrieved18 February2014.
Equine Research (2004).Horse Conformation: Structure, Soundness, and Performance.Lyons Press.ISBN 978-1-59228-487-0.Archived fromthe originalon 7 July 2014.Retrieved23 July2013.
Evans, James Warren (15 February 1990).The Horse.W. H. Freeman.ISBN 978-0-7167-1811-6.Retrieved23 July2013.
Hayes, M. Horace; Rossdale, Peter D. (March 1988).Veterinary Notes for Horse Owners: An Illustrated Manual of Horse Medicine and Surgery.Simon & Schuster.ISBN 978-0-671-76561-3.Retrieved1 July2013.
McBane, Susan (2001).Modern Horse Breeding: A Guide for Owners.Globe Pequot Press.ISBN 978-1-58574-389-6.Archived fromthe originalon 7 July 2014.Retrieved18 February2014.

Marsupials

Parker, Rick (13 January 2012).Equine Science(4 ed.). Cengage Learning.ISBN 978-1-111-13877-6.Retrieved18 February2014.
Flannery, Tim (2008).Chasing Kangaroos: A Continent, a Scientist, and a Search for the World's Most Extraordinary Creature.Grove/Atlantic, Incorporated. pp. 60–.ISBN 9780802143716.Retrieved5 May2013.
Hunsaker, Don II (2 December 2012).The Biology of Marsupials.Elsevier Science.ISBN 978-0-323-14620-3.Retrieved18 February2014.
Jones, Menna E.; Dickman, Chris R.; Archer, Mike; Archer, Michael (2003).Predators With Pouches: The Biology of Carnivorous Marsupials.Csiro Publishing.ISBN 9780643066342.Retrieved5 May2013.
King, Anna (2001)."Discoveries about Marsupial Reproduction".Iowa State University Biology Dept. Archived fromthe originalon September 5, 2012.Retrieved2012-11-22.
Stonehouse, Bernard; Gilmore, Desmond (1977).The Biology of marsupials.University Park Press.ISBN 978-0-8391-0852-8.Retrieved25 July2013.
Tyndale-Biscoe, C. Hugh (2005).Life of Marsupials.Csiro Publishing.ISBN 978-0-643-06257-3.Retrieved18 February2014.

Other animals

Austin, Colin Russell; Short, Roger Valentine (21 March 1985).Reproduction in Mammals: Volume 4, Reproductive Fitness.Cambridge University Press.ISBN 978-0-521-31984-3.Retrieved22 July2013.
Bassert, Joanna M.; McCurnin, Dennis M. (1 April 2013).McCurnin's Clinical Textbook for Veterinary Technicians.Elsevier Health Sciences.ISBN 978-1-4557-2884-8.Retrieved18 February2014.
Bates, Paul J. J.; Ratrimomanarivo, Fanja H.; Harrison, David L.; Goodman, Steven M. (December 2006). "A description of a new species of Pipistrellus (Chiroptera: Vespertilionidae) from Madagascar with a review of related Vespertilioninae from the island".Acta Chiropterologica.8(2): 299–324.doi:10.3161/1733-5329(2006)8[299:ADOANS]2.0.CO;2.S2CID 85825521.
Beck, Benjamin B.; Wemmer, Christen M. (1983).The Biology and management of an extinct species: Père David's deer.Noyes Publications.ISBN 978-0-8155-0938-7.Retrieved5 July2013.
Burns, Eugene (1953).The sex life of wild animals: a North American study.Rinehart.Retrieved23 July2013.
Carnaby, Trevor (22 January 2007).Beat About the Bush: Mammals.Jacana Media.ISBN 978-1-77009-240-2.Retrieved19 July2013.
Brehm, Alfred Edmund (1895)."Brehm's Life of Animals".Chicago: A. N. Marquis & Company.Retrieved2013-11-08.{{cite journal}}:Cite journal requires|journal=(help)
Elbroch, Lawrence Mark; Kresky, Michael Raymond; Evans, Jonah Wy (7 April 2012).Field Guide to Animal Tracks and Scat of California.University of California Press.ISBN 978-0-520-95164-8.Retrieved5 July2013.
Eltringham, Stewart Keith (1979).The ecology and conservation of large African mammals.Macmillan.ISBN 978-0-333-23580-5.Retrieved20 July2013.
Estes, Richard (1998).The safari companion: a guide to watching African mammals, including hoofed mammals, carnivores, and primates.Chelsea Green Publishing.ISBN 978-1-890132-44-6.
Frandson, Rowen D.; Wilke, W. Lee; Fails, Anna Dee (30 June 2009).Anatomy and Physiology of Farm Animals.John Wiley & Sons.ISBN 978-0-8138-1394-3.Retrieved1 July2013.
Geist, Valerius (1993).Elk Country.T&N Children's Publishing.ISBN 978-1-55971-208-8.Retrieved5 July2013.
Hayssen, Virginia Douglass; Tienhoven, Ari Van (1993).Asdell's Patterns of Mammalian Reproduction: A Compendium of Species-Specific Data.Cornell University Press.ISBN 978-0-8014-1753-5.Retrieved23 July2013.
Heptner, V. G.; Sludskii, A. A. (2002).Mammals of the Soviet Union. Vol. II, part 1b, Carnivores (Mustelidae and Procyonidae).Washington, D.C.: Smithsonian Institution Libraries and National Science Foundation.ISBN 978-90-04-08876-4.Retrieved2013-11-08.
Hoffmeister, Donald F. (2002).Mammals of Illinois.University of Illinois Press.ISBN 978-0-252-07083-9.Retrieved22 July2013.
Hooper, E.T.; Musser, G.G. (1964)."The glans penis in Neotropical cricetines (Family Muridae) with comments on classification of muroid rodents".Miscellaneous Publications of the University of Michigan Museum of Zoology.123:1–57.hdl:2027.42/56367.
Horowitz, Barbara N.; Bowers, Kathryn (12 June 2012).Zoobiquity: What Animals Can Teach Us About Being Human.Doubleday Canada.ISBN 978-0-385-67061-6.Retrieved25 July2013.
Horwich, Robert H. (June 1972).The ontogeny of social behavior in the gray squirrel (Sciurus carolinensis).P. Parey.ISBN 978-3-489-68036-9.Retrieved23 July2013.
Jackson, Hartley H. (January 1961).Mammals of Wisconsin.Univ of Wisconsin Press.ISBN 978-0-299-02150-4.Retrieved22 July2013.
Journal of the Mammalogical Society of Japan.The Society. 1986.Retrieved5 July2013.
Khanna, Dev Raj; Yadav, P. R. (1 January 2005).Biology Of Mammals.Discovery Publishing House.ISBN 978-81-7141-934-0.Retrieved20 July2013.
Kingdon, Jonathan (January 1984).East African Mammals: An Atlas of Evolution in Africa. Vol. I.University of Chicago Press.ISBN 978-0-226-43718-7.Retrieved22 July2013.
Kingdon, Jonathan (1984).East African Mammals: An Atlas of Evolution in Africa.University of Chicago Press.ISBN 9780226437187.
König, Horst Erich; Liebich, Hans-Georg (2007).Veterinary Anatomy of Domestic Mammals: Textbook and Atlas.Schattauer Verlag.ISBN 978-3-7945-2485-3.Retrieved23 July2013.
Kotpal, R. L. (2010).Modern Text Book Of Zoology Vertebrates.Rastogi Publications.ISBN 978-81-7133-891-7.Retrieved23 July2013.
Krause, William J. (1 March 2008).An Atlas of Opossum Organogenesis.Universal-Publishers.ISBN 978-1-58112-969-4.Retrieved20 July2013.
Linzey, Donald W. (28 December 2011).Vertebrate Biology.JHU Press.ISBN 978-1-4214-0040-2.Retrieved20 July2013.
Lukefahr, Steven D.; Cheeke, Peter R.; Patton, Nephi M. (2013).Rabbit Production.CABI.ISBN 978-1-78064-012-9.Retrieved20 July2013.
Natural History Bulletin of the Siam Society.1975.Retrieved5 July2013.
Percequillo, A.R.; Weksler, M.; Costa, L.P. (2011)."A new genus and species of rodent from the Brazilian Atlantic Forest (Rodentia: Cricetidae: Sigmodontinae: Oryzomyini), with comments on oryzomyine biogeography".Zoological Journal of the Linnean Society.161(2): 357–390.doi:10.1111/j.1096-3642.2010.00643.x.
Rose, Kenneth D.; Archibald, J. David (22 February 2005).The Rise of Placental Mammals: Origins and Relationships of the Major Extant Clades.JHU Press.ISBN 978-0-8018-8022-3.Retrieved22 July2013.
Roze, Uldis (2009).The North American Porcupine.Cornell University Press.ISBN 978-0-8014-4646-7.Retrieved25 July2013.
Sarkar, Amita (1 January 2003).Sexual Behaviour In Animals.Discovery Publishing House.ISBN 978-81-7141-746-9.Retrieved20 July2013.
Schatten, Heide; Constantinescu, Gheorghe M. (21 March 2008).Comparative Reproductive Biology.John Wiley & Sons.ISBN 978-0-470-39025-2.Retrieved23 July2013.
Small, Meredith F. (1993).Female Choices: Sexual Behavior of Female Primates.Cornell University Press.ISBN 978-0-8014-8305-9.Retrieved23 July2013.
Skinner, J. D.; Chimimba, Christian T. (15 November 2005).The Mammals of the Southern African Sub-region.Cambridge University Press.ISBN 978-0-521-84418-5.Retrieved19 July2013.
Staker, Lynda (2006).The Complete Guide to the Care of Macropods: A Comprehensive Guide to the Handrearing, Rehabilitation and Captive Management of Kangaroo Species.Lynda Staker.ISBN 978-0-9775751-0-7.Retrieved19 July2013.
Strum, Shirley C.; Fedigan, Linda Marie (15 August 2000).Primate Encounters: Models of Science, Gender, and Society.University of Chicago Press.ISBN 978-0-226-77754-2.Retrieved22 July2013.
Sturtz, Robin; Asprea, Lori (30 July 2012).Anatomy and Physiology for Veterinary Technicians and Nurses: A Clinical Approach.John Wiley & Sons.ISBN 978-1-118-40585-7.Retrieved22 July2013.
Peter J Chenoweth; Steven Lorton (30 April 2014).Animal Andrology: Theories and Applications.CABI.ISBN 978-1-78064-316-8.
Verts, B. J.; Carraway, Leslie N. (1998).Land Mammals of Oregon.University of California Press. p. 41.ISBN 978-0-520-21199-5.Retrieved20 July2013.
Voss, R.S.; Linzey, A.V. (1981)."Comparative gross morphology of male accessory glands among Neotropical Muridae (Mammalia: Rodentia) with comments on systematic implications".Miscellaneous Publications of the University of Michigan Museum of Zoology.159:1–41.hdl:2027.42/56403.
Voss, R.S (2003)."A new species ofThomasomys(Rodentia: Muridae) from eastern Ecuador, with remarks on mammalian diversity and biogeography in the Cordillera Oriental ".American Museum Novitates(3421): 1–47.doi:10.1206/0003-0082(2003)421<0001:ansotr>2.0.co;2.hdl:2246/2850.S2CID 62795333.
Weksler, M. (2006)."Phylogenetic relationships of oryzomyine rodents (Muroidea: Sigmodontinae): separate and combined analyses of morphological and molecular data".Bulletin of the American Museum of Natural History.296:1–149.doi:10.1206/0003-0090(2006)296[0001:PROORM]2.0.CO;2.hdl:2246/5777.S2CID 86057173.
Wilson, Don E.; Reeder, DeeAnn M. (16 November 2005).Mammal Species of the World: A Taxonomic and Geographic Reference.JHU Press.ISBN 978-0801882210.Retrieved20 July2013.

External links

Kinsey Institute on the penis

[SciencesMexico2010-1] Janet Leonard; Alex Cordoba-Aguilar R (18 June 2010).The Evolution of Primary Sexual Characters in Animals.Oxford University Press.ISBN 978-0-19-971703-3.Archivedfrom the original on 11 October 2013.Retrieved20 July2013.

[2] Schmitt, V.; Anthes, N.; Michiels, N. K. (2007)."Mating behaviour in the sea slug Elysia timida (Opisthobranchia, Sacoglossa): hypodermic injection, sperm transfer and balanced reciprocity".Frontiers in Zoology.4:17.doi:10.1186/1742-9994-4-17.ISSN 1742-9994.PMC1934903.PMID 17610714.

[oed-yard-3] Basu, S. C. (2011).Male Reproductive Dysfunction.JP Medical Ltd. p. 101.ISBN 9789350252208.

[oed-penis-4] Simpson, John,ed. (1989)."penis,n.".Oxford English Dictionary(second ed.).Oxford University Press.ISBN 978-0-19-861186-8.^{[dead link]}

[5] "Online Etymology Dictionary".Etymonline.Archivedfrom the original on 2011-06-06.Retrieved2011-05-28.

[6] Watson, Lyall (2000-04-17).Jacobson's Organ: And the Remarkable Nature of Smell.W. W. Norton & Company.ISBN 978-0-393-24493-9.

[Mech-7] Mech, L. David;Boitani, Luigi,eds. (2003).Wolves: Behaviour, Ecology and Conservation.University of Chicago Press.ISBN 0-226-51696-2.

[8] Dunlop, Jason A.; Penney, David (2012).Fossil Arachnids.Siri Scientific Press. p. 44.ISBN 978-0-95677-954-0.

[9] Cordoba-Aguilar, Alex; Leonard, Janet (2010).The Evolution of Primary Sexual Characters in Animals.Oxford University Press. pp. 216–221.ISBN 978-0-19971-703-3.

[Wake1992-10] Marvalee H. Wake (15 September 1992).Hyman's Comparative Vertebrate Anatomy.University of Chicago Press. p. 583.ISBN 978-0-226-87013-7.Archivedfrom the original on 31 December 2013.Retrieved23 July2013.

[11] Feldhamer, George A. (2007-09-07).Mammalogy: Adaptation, Diversity, Ecology.JHU Press.ISBN 978-0-8018-8695-9.

[12] Sanger TJ, Gredler ML, Cohn MJ (October 2015)."Resurrecting embryos of the tuatara, Sphenodon punctatus, to resolve vertebrate phallus evolution".Biology Letters.11(10): 20150694.doi:10.1098/rsbl.2015.0694.PMC4650183.PMID 26510679.

[13] Brennan, Patricia L. R.; Clark, Christopher J.; Prum, Richard O. (2010-05-07)."Explosive eversion and functional morphology of the duck penis supports sexual conflict in waterfowl genitalia".Proceedings of the Royal Society B: Biological Sciences.277(1686): 1309–1314.doi:10.1098/rspb.2009.2139.ISSN 0962-8452.PMC2871948.PMID 20031991.

[Lombardi1998-14] Julian Lombardi (1998).Comparative Vertebrate Reproduction.Springer.ISBN 978-0-7923-8336-9.Archivedfrom the original on 26 March 2014.Retrieved5 December2012.

[MobileReference2009-15] MobileReference (15 December 2009).The Illustrated Encyclopedia of European Birds: An Essential Guide to Birds of Europe.MobileReference.ISBN 978-1-60501-557-6.Archived fromthe originalon 26 March 2014.Retrieved5 December2012.

[Gill2006-16] Frank B. Gill (6 October 2006).Ornithology.Macmillan. pp. 414–.ISBN 978-0-7167-4983-7.Archivedfrom the original on 7 January 2014.Retrieved5 December2012.

[17] Tim Birkhead (2000).Promiscuity: An Evolutionary History of Sperm Competition.Harvard University Press. p. 102.ISBN 978-0-674-00666-9.Archivedfrom the original on 26 March 2014.Retrieved23 July2013.

[HayssenTienhoven1993-18] Virginia Douglass Hayssen; Ari Van Tienhoven (1993).Asdell's Patterns of Mammalian Reproduction: A Compendium of Species-Specific Data.Cornell University Press.ISBN 978-0-8014-1753-5.Retrieved23 July2013.

[19] Chenoweth, Peter J.; Lorton, Steven P. (2022-02-03).Manual of Animal Andrology.CABI.ISBN 978-1-78924-350-5.

[Reece-20] William O. Reece (2009-03-04).Functional Anatomy and Physiology of Domestic Animals.John Wiley & Sons.ISBN 9780813814513.Archivedfrom the original on 2018-03-20.

[21] Dixson, Alan F. (2009).Sexual Selection and the Origins of Human Mating Systems.John OUP Oxford. p. 68.ISBN 978-0-19156-973-9.

[Long2006-22] Susan Long (2006).Veterinary Genetics and Reproductive Physiology.Churchill Livingstone Elsevier.ISBN 978-0-7506-8877-2.Archivedfrom the original on 2014-03-26.Retrieved2013-11-08.

[autogenerated3-23] R. F. Ewer (1998).The Carnivores.Cornell University Press. p. 116.ISBN 978-0-8014-8493-3.Archivedfrom the original on 26 March 2014.Retrieved23 July2013.

[differentiation-24] Glickman, SE; Cunha, GR; Drea, CM; Conley, AJ; Place, NJ (2006)."Mammalian sexual differentiation: lessons from the spotted hyena"(PDF).Trends Endocrinol Metab.17(9): 349–356.doi:10.1016/j.tem.2006.09.005.PMID 17010637.S2CID 18227659.Archived(PDF)from the original on 2013-02-22.

[25] Aronson, L. R.; Cooper, M. L. (1967)."Penile spines of the domestic cat: their endocrine-behavior relations"(PDF).Anat. Rec.157(1): 71–8.doi:10.1002/ar.1091570111.PMID 6030760.S2CID 13070242.Archived fromthe original(PDF)on 2017-06-20.

[26] Tyndale-Biscoe, C. Hugh; Renfree, Marilyn (1987-01-30).Reproductive Physiology of Marsupials.Cambridge University Press.ISBN 978-0-521-33792-2.

[27] Armati, Patricia J.; Dickman, Chris R.; Hume, Ian D. (2006-08-17).Marsupials.Cambridge University Press.ISBN 978-1-139-45742-2.

[28] Gadow, H. On the systematic position of Notoryctes typhlops. Proc. Zool. Soc. London 1892, 361–370 (1892).

[29] 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.

[30] Lutz, Dick (2005), Tuatara: A Living Fossil, Salem, Oregon: DIMI PRESS,ISBN 0-931625-43-2

[31] Kelly, D. A. (2004)."Turtle and mammal penis designs are anatomically convergent".Proceedings. Biological Sciences.271(Suppl 5): S293–S295.doi:10.1098/rsbl.2004.0161.PMC1810052.PMID 15503998.

[32] Ruschenberger, William Samuel Waithman (1846).Elements of Herpetology, and of Ichthyology: Prepared for the Use of Schools and Colleges.Grigg & Elliot. pp. 129–145.

[Britannica-33] "Penis | Description, Anatomy, & Physiology | Britannica".Encyclopædia Britannica.January 2024.

[Lienhard2010-34] Lienhard, Charles; Oliveira do Carmo, Thais; Lopes Ferreira, Rodrigo (2010)."A new genus of Sensitibillini from Brazilian caves (Psocodea: 'Psocoptera': Prionoglarididae)".Revue suisse de Zoologie.117(4): 611–635.doi:10.5962/bhl.part.117600.ISSN 0035-418X.Archivedfrom the original on 2014-11-03.

[Yoshizawa_et.al.2018-35] 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 Letters14(11):doi.org/10.1098/rsbl.2018.0533

[YoshizawaFerreira2014-36] Yoshizawa, Kazunori; Ferreira, Rodrigo L.; Kamimura, Yoshitaka; Lienhard, Charles (2014)."Female Penis, Male Vagina, and Their Correlated Evolution in a Cave Insect".Current Biology.24(9): 1006–10.doi:10.1016/j.cub.2014.03.022.hdl:2115/56857.ISSN 0960-9822.PMID 24746797.

[Yoshizawa_et.al.2019-37] Yoshizawa K, Ferreira R.L., Lienhard C & Kamimura Y. (2019). "Why Did a Female Penis Evolve in a Small Group of Cave Insects?".BioEssays41(6):doi.org/10.1002/bies.201900005

[Yoshizawa_et.al._2014-38] Yoshizawa K, Ferreira R.L., Kamimura Y & Lienhard C."A Transgender Brazilian Cave Insect".The Winnower3/9/2014

[Hollier&Hollier_2020-39] Hollier J & Hollier A. (2020)."The retired taxonomist and the gynosome – the discovery of the female penis".Antenna44(3): p. 122-125

[Hayssen2020-40] Hayssen V. (2020). "Misconceptions about Conception and Other Fallacies: Historical Bias in Reproductive Biology".Integrative and Comparative Biology60(3): p. 683-791:doi.org/10.1093/icb/icaa035

[rietstap-1884-41] Rietstap, J. B. (1884)."Armorial général; précédé d'un Dictionnaire des termes du blason".G. B. van Goor zonen: XXXI.Vilené: se dit un animal qui a la marque du sexe d'un autre émail que le corps{{cite journal}}:Cite journal requires|journal=(help)

[1]

[2]

[3]

[4]

[5]

[6]

[7]

[8]

[9]

[10]

[11]

[12]

[13]

[14]

[15]

[16]

[17]

[18]

[19]

[20]

[21]

[22]

[23]

[24]

[25]

[26]

[27]

[28]

[29]

[30]

[31]

[32]

[33]

[34]

[35]

[36]

[37]

[38]

[39]

[40]

[41]