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Internal fertilization is the union of an egg and sperm cell during sexual reproduction inside the female body. Internal fertilization, unlike its counterpart, external fertilization, brings more control to the female with reproduction.^[1] For internal fertilization to happen there needs to be a method for the male to introduce the sperm into the female's reproductive tract.

Most taxa that reproduce by internal fertilization are gonochoric.^[2]^{: 124–125} Male mammals, reptiles, and certain other vertebrates transfer sperm into the female's vagina or cloaca through an intromittent organ during copulation.^[3]^[4]^[5] In most birds, the cloacal kiss is used, the two animals pressing their cloacas together while transferring sperm.^[6] Salamanders, spiders, some insects and some molluscs undertake internal fertilization by transferring a spermatophore, a bundle of sperm, from the male to the female. Following fertilization, the embryos are laid as eggs in oviparous organisms, or continue to develop inside the reproductive tract of the mother to be born later as live young in viviparous organisms.

Evolution of internal fertilization

Internal fertilization evolved many times in animals.^[2]^: 2 According to David B. Dusenbery all the features with internal fertilization were most likely a result from oogamy.^[7] It has been argued that internal fertilization evolve because of sexual selection through sperm competition.^[8]

In amphibians, internal fertilization evolved from external fertilization.^[9]

Methods of internal fertilization

Fertilization which takes place inside the female body is called internal fertilization in animals is done through the following different ways:^[10]^[11]^[12]

Copulation,^[13] which involves the insertion of the penis or other intromittent organ into the vagina (in most mammals)^[14] or to the cloaca in monotremes, most reptiles, some birds, the tailed frog, some fish, the disappeared dinosaurs, as well as in other non-vertebrate animals.^[5]^[15]
Cloacal kiss, which consists in that the two animals touch their cloacae together in order to transfer the sperm of the male to the female. It is used in most birds and in the tuatara, that do not have an intromittent organ.^[6]
Via spermatophore, a sperm-containing cap placed by the male in the female's cloaca. Usually, the sperm is stored in spermathecae on the roof of the cloaca until it is needed at the time of oviposition. It is used by some salamander and newt species, by the Arachnida, some insects and some mollusks.^[16]^[17]
In sponges, sperm cells are released into the water to fertilize ova that are retained by the female. Some species of sponge participate in external fertilization where the ova is released.^[18]

Expulsion

At some point, the growing egg or offspring must be expelled. There are several possible modes of reproduction. These are traditionally classified as follows:

Oviparity, as in most invertebrates and reptiles, monotremes, dinosaurs and all birds which lay eggs that continue to develop after being laid, and hatch later.^[19]
Viviparity, as in almost all mammals (such as whales, kangaroos and humans) which bear their young live. The developing young spend proportionately more time within the female's reproductive tract. The young are later released to survive on their own, with varying amounts of help from the parent(s) of the species.^[20]
Ovoviviparity, as in the garter snake, most vipers, and the Madagascar hissing cockroach, which have eggs (with shells) that hatch as they are laid, making it resemble live birth.^[21]

Advantages to internal fertilization

Internal fertilization allows for:

Female mate choice, which gives the female the ability to choose her partner before and after mating. The female cannot do this with external fertilization because she may have limited control of who is fertilizing her eggs, and when they are being fertilized.^[1]
Making a decision for the conditions of reproduction, like location and time.^[22] In external fertilization a female can only choose the time in which she releases her eggs, but not when they are fertilized. This is similar, in ways, to cryptic female choice.
Egg protection on dry land.^[23] While oviparous animals either have a jelly like ovum or a hard shell enclosing their egg, internally fertilizing animals grow their eggs and offspring inside themselves. This offers protection from predators and from dehydration on land.^[24] This allows for a higher chance of survival when there is a regulated temperature and protected area within the mother.

Disadvantages to internal fertilization

Gestation can and will add additional risks for the mother.^[25] The additional risks from gestation come from extra energy demands.
Along with internal fertilization comes sexual reproduction, in most cases. Sexual reproduction comes with some risks as well. The risks with sexual reproduction are with intercourse, it is infrequent and only works well during peak fertility. While animals which externally fertilize are able to release egg and sperm, usually into the water, not needing a specific partner to reproduce.^[25]
Fewer offspring are produced through internal fertilization in comparison to external fertilization. This is both because the mother cannot hold and grow as many offspring as eggs, and the mother cannot provide and obtain enough resources for a larger amount of offspring.^[26]

Fish

Some species of fish like guppies have the ability to internally fertilize, this process happens by the male inserting a tubular fin into the female's reproductive opening and then will deposit sperm into her reproductive tract. There are other species of fish that are mouthbrooders which means that one fish puts the eggs in its mouth for incubation. A certain type of fish that is a mouthbrooder is called cichlids and many of them are maternal mouthbrooders. The process for this is the female would lay the egg and pick it up in her mouth. Then the males will encourage the female to open her mouth so they can fertilize the eggs while it is in the female's mouth.^[27] Internal fertilization in cartilaginous fishes contains the same evolutionary origin as reptiles, birds, and mammals that internally fertilize. Also in these internally fertilizing fish while the sperm is transferred to the reproductive tract there is no noticeable change in tonality.^[28]

Amphibians

Most amphibians have external fertilization but there is an exception to some like salamanders which mostly have internal fertilization. Salamanders do not use intercourse for sexual reproduction due to their lack of external penis. Rather, the male salamander produces an encased capsule of sperm and nutrients called a spermatophore. The male deposits a spermatophore on the ground and the female will pick it up with her cloaca (a combined urinary and genital opening) and fertilize her eggs with it.^[27] Over time amphibians have been found evolving to increasing internal fertilization.^{[citation needed]} Within amphibians, it is common for high vertebrates to internally fertilize because of the transition from water to land during vertebrate evolution. There is an advantage for the amphibians who are internally fertilizing allowing for the selection of a time and place for reproduction.^[22]

Birds

Most birds do not have penises, but achieve internal fertilization via cloacal contact (or "cloaca kiss"). In these birds, males and females contact their cloacas together, typically briefly, and transfer sperm to the female. However, water fowls such as ducks and geese have penises and are able to use them for internal fertilization. ^[27] While birds have internal fertilization, most species no longer have phallus structures. This makes them the only vertebrate taxon to fall into both categories of lacking the phallus but participating in internal fertilization.^[29]

References

^ ^a ^b Alonzo SH, Stiver KA, Marsh-Rollo SE (August 2016). "Ovarian fluid allows directional cryptic female choice despite external fertilization". Nature Communications. 7 (1): 12452. Bibcode:2016NatCo...712452A. doi:10.1038/ncomms12452. PMC 4990696. PMID 27529581.
^ ^a ^b Leonard, Janet; Cordoba-Aguilar, Alex (2010-07-16). The Evolution of Primary Sexual Characters in Animals. Oxford University Press. ISBN 978-0-19-971703-3.
^ Naguib, Marc (2020-04-19). Advances in the Study of Behavior. Academic Press. ISBN 978-0-12-820726-0.
^ Hyman LH (15 September 1992). Hyman's Comparative Vertebrate Anatomy. University of Chicago Press. ISBN 978-0-226-87013-7.
^ ^a ^b Austin CR (1984). "Evolution of the copulatory apparatus". Bolletino di Zoologia. 51 (1–2): 249–269. doi:10.1080/11250008409439463.
^ ^a ^b Romer AS, Parsons TS (1977). The Vertebrate Body. Philadelphia, PA: Holt-Saunders International. pp. 396–399. ISBN 978-0-03-910284-5.
^ Dusenbery, David B. (2009). Living at Micro Scale: The Unexpected Physics of Being Small. Harvard University Press. p. 326. ISBN 978-0-674-06021-0.
^ Smith, Robert L. (2012-12-02). Sperm Competition and the Evolution of Animal Mating systems. Elsevier. pp. 7–8. ISBN 978-0-323-14313-4.
^ Sawada, Hitoshi; Inoue, Naokazu; Iwano, Megumi (2014-02-07). Sexual Reproduction in Animals and Plants. Springer. pp. 97–98. ISBN 978-4-431-54589-7.
^ Reichard UH (2002). "Monogamy—A variable relationship" (PDF). Max Planck Research. 3: 62–7. Archived from the original (PDF) on 14 May 2011. Retrieved 24 April 2013.
^ Lipton JE, Barash DP (2001). The Myth of Monogamy: Fidelity and Infidelity in Animals and People. San Francisco: W.H. Freeman and Company. ISBN 978-0-7167-4004-9.
^ Morell V (September 1998). "A new look at monogamy". Science. 281 (5385): 1982–3. doi:10.1126/science.281.5385.1982. PMID 9767050. S2CID 31391458.
^ Lombardi J (6 December 2012). Comparative Vertebrate Reproduction. Springer Science & Business Media. ISBN 978-1-4615-4937-6.
^ Naguib, Marc (2020-04-19). Advances in the Study of Behavior. Academic Press. ISBN 978-0-12-820726-0.
^ Diamond J (1991). The Rise and Fall of the Third Chimpanzee. Radius. pp. 360 pages. ISBN 978-0091742683.
^ Wedell N, Tregenza T, Simmons LW (July 2008). "Nuptial gifts fail to resolve a sexual conflict in an insect". BMC Evolutionary Biology. 8 (1): 204. Bibcode:2008BMCEE...8..204W. doi:10.1186/1471-2148-8-204. PMC 2491630. PMID 18627603.
^ Sozou PD, Seymour RM (September 2005). "Costly but worthless gifts facilitate courtship". Proceedings. Biological Sciences. 272 (1575): 1877–84. doi:10.1098/rspb.2005.3152. PMC 1559891. PMID 16191592.
^ Bergquist PR (1978). Sponges. London: Hutchinson. ISBN 9780520036581.
^ Lodé T (2001). Les stratégies de reproduction des animaux [Reproduction Strategies in the Animal Kingdom] (in French). Paris: Dunod Sciences.
^ Blackburn DG (January 2000). "Classification of the reproductive patterns of amniotes". Herpetological Monographs. 14: 371–7. doi:10.2307/1467051. JSTOR 1467051.
^ Carrier JC, Musick JA, Heithaus MR, eds. (2012). Biology of Sharks and Their Relatives. CRC Press. pp. 296–301. ISBN 978-1439839249.
^ ^a ^b Yokoe M, Takayama-Watanabe E, Saito Y, Kutsuzawa M, Fujita K, Ochi H, et al. (2016-08-31). Klymkowsky M (ed.). "A Novel Cysteine Knot Protein for Enhancing Sperm Motility That Might Facilitate the Evolution of Internal Fertilization in Amphibians". PLOS ONE. 11 (8): e0160445. Bibcode:2016PLoSO..1160445Y. doi:10.1371/journal.pone.0160445. PMC 5007030. PMID 27579691.
^ Altig R, McDiarmid RW (December 2007). "Morphological diversity and evolution of egg and clutch structure in amphibians". Herpetological Monographs. 21 (1): 1–32. doi:10.1655/06-005.1. S2CID 55728625.
^ "43.2A: External and Internal Fertilization". Biology LibreTexts. 2018-07-17. Retrieved 2020-11-09.
^ ^a ^b Wallen K, Zehr JL (February 2004). "Hormones and history: the evolution and development of primate female sexuality". Journal of Sex Research. 41 (1): 101–12. doi:10.1080/00224490409552218. PMC 1255935. PMID 15216429.
^ Parker, G. A. (1970). "Sperm Competition and Its Evolutionary Consequences in the Insects". Biological Reviews. 45 (4): 525–567. doi:10.1111/j.1469-185X.1970.tb01176.x. ISSN 1469-185X. S2CID 85156929.
^ ^a ^b ^c Cotner, Sehoya; Wassenberg, Deena (2020). The Evolution and Biology of Sex.
^ Engel, Kathrin M; Dzyuba, Viktoriya; Ninhaus-Silveira, Alexandre; Veríssimo-Silveira, Rosicleire; Dannenberger, Dirk; Schiller, Jürgen; Steinbach, Christoph; Dzyuba, Borys (February 2020). "Sperm Lipid Composition in Early Diverged Fish Species: Internal vs. External Mode of Fertilization". Biomolecules. 10 (2): 172. doi:10.3390/biom10020172. PMC 7072473. PMID 31979037.
^ Brennan, Patricia L. R.; Birkhead, Tim R.; Zyskowski, Kristof; Van Der Waag, Jessica; Prum, Richard O. (10 September 2008). "Independent evolutionary reductions of the phallus in basal birds". Journal of Avian Biology. 39 (5): 487–492. doi:10.1111/j.0908-8857.2008.04610.x.

[Alonzo_2016-1] Alonzo SH, Stiver KA, Marsh-Rollo SE (August 2016). "Ovarian fluid allows directional cryptic female choice despite external fertilization". Nature Communications. 7 (1): 12452. Bibcode:2016NatCo...712452A. doi:10.1038/ncomms12452. PMC 4990696. PMID 27529581.

[:2-2] Leonard, Janet; Cordoba-Aguilar, Alex (2010-07-16). The Evolution of Primary Sexual Characters in Animals. Oxford University Press. ISBN 978-0-19-971703-3.

[3] Naguib, Marc (2020-04-19). Advances in the Study of Behavior. Academic Press. ISBN 978-0-12-820726-0.

[Hyman1992-4] Hyman LH (15 September 1992). Hyman's Comparative Vertebrate Anatomy. University of Chicago Press. ISBN 978-0-226-87013-7.

[Austin-5] Austin CR (1984). "Evolution of the copulatory apparatus". Bolletino di Zoologia. 51 (1–2): 249–269. doi:10.1080/11250008409439463.

[VB-6] Romer AS, Parsons TS (1977). The Vertebrate Body. Philadelphia, PA: Holt-Saunders International. pp. 396–399. ISBN 978-0-03-910284-5.

[7] Dusenbery, David B. (2009). Living at Micro Scale: The Unexpected Physics of Being Small. Harvard University Press. p. 326. ISBN 978-0-674-06021-0.

[8] Smith, Robert L. (2012-12-02). Sperm Competition and the Evolution of Animal Mating systems. Elsevier. pp. 7–8. ISBN 978-0-323-14313-4.

[9] Sawada, Hitoshi; Inoue, Naokazu; Iwano, Megumi (2014-02-07). Sexual Reproduction in Animals and Plants. Springer. pp. 97–98. ISBN 978-4-431-54589-7.

[10] Reichard UH (2002). "Monogamy—A variable relationship" (PDF). Max Planck Research. 3: 62–7. Archived from the original (PDF) on 14 May 2011. Retrieved 24 April 2013.

[Barash,Lipton,2001-11] Lipton JE, Barash DP (2001). The Myth of Monogamy: Fidelity and Infidelity in Animals and People. San Francisco: W.H. Freeman and Company. ISBN 978-0-7167-4004-9.

[Gowaty,Morell,1998-12] Morell V (September 1998). "A new look at monogamy". Science. 281 (5385): 1982–3. doi:10.1126/science.281.5385.1982. PMID 9767050. S2CID 31391458.

[Lombardi2012-13] Lombardi J (6 December 2012). Comparative Vertebrate Reproduction. Springer Science & Business Media. ISBN 978-1-4615-4937-6.

[14] Naguib, Marc (2020-04-19). Advances in the Study of Behavior. Academic Press. ISBN 978-0-12-820726-0.

[Diamond-15] Diamond J (1991). The Rise and Fall of the Third Chimpanzee. Radius. pp. 360 pages. ISBN 978-0091742683.

[wedell-16] Wedell N, Tregenza T, Simmons LW (July 2008). "Nuptial gifts fail to resolve a sexual conflict in an insect". BMC Evolutionary Biology. 8 (1): 204. Bibcode:2008BMCEE...8..204W. doi:10.1186/1471-2148-8-204. PMC 2491630. PMID 18627603.

[17] Sozou PD, Seymour RM (September 2005). "Costly but worthless gifts facilitate courtship". Proceedings. Biological Sciences. 272 (1575): 1877–84. doi:10.1098/rspb.2005.3152. PMC 1559891. PMID 16191592.

[18] Bergquist PR (1978). Sponges. London: Hutchinson. ISBN 9780520036581.

[19] Lodé T (2001). Les stratégies de reproduction des animaux [Reproduction Strategies in the Animal Kingdom] (in French). Paris: Dunod Sciences.

[20] Blackburn DG (January 2000). "Classification of the reproductive patterns of amniotes". Herpetological Monographs. 14: 371–7. doi:10.2307/1467051. JSTOR 1467051.

[21] Carrier JC, Musick JA, Heithaus MR, eds. (2012). Biology of Sharks and Their Relatives. CRC Press. pp. 296–301. ISBN 978-1439839249.

[cysteine_knot-22] Yokoe M, Takayama-Watanabe E, Saito Y, Kutsuzawa M, Fujita K, Ochi H, et al. (2016-08-31). Klymkowsky M (ed.). "A Novel Cysteine Knot Protein for Enhancing Sperm Motility That Might Facilitate the Evolution of Internal Fertilization in Amphibians". PLOS ONE. 11 (8): e0160445. Bibcode:2016PLoSO..1160445Y. doi:10.1371/journal.pone.0160445. PMC 5007030. PMID 27579691.

[23] Altig R, McDiarmid RW (December 2007). "Morphological diversity and evolution of egg and clutch structure in amphibians". Herpetological Monographs. 21 (1): 1–32. doi:10.1655/06-005.1. S2CID 55728625.

[24] "43.2A: External and Internal Fertilization". Biology LibreTexts. 2018-07-17. Retrieved 2020-11-09.

[:1-25] Wallen K, Zehr JL (February 2004). "Hormones and history: the evolution and development of primate female sexuality". Journal of Sex Research. 41 (1): 101–12. doi:10.1080/00224490409552218. PMC 1255935. PMID 15216429.

[26] Parker, G. A. (1970). "Sperm Competition and Its Evolutionary Consequences in the Insects". Biological Reviews. 45 (4): 525–567. doi:10.1111/j.1469-185X.1970.tb01176.x. ISSN 1469-185X. S2CID 85156929.

[:0-27] Cotner, Sehoya; Wassenberg, Deena (2020). The Evolution and Biology of Sex.

[28] Engel, Kathrin M; Dzyuba, Viktoriya; Ninhaus-Silveira, Alexandre; Veríssimo-Silveira, Rosicleire; Dannenberger, Dirk; Schiller, Jürgen; Steinbach, Christoph; Dzyuba, Borys (February 2020). "Sperm Lipid Composition in Early Diverged Fish Species: Internal vs. External Mode of Fertilization". Biomolecules. 10 (2): 172. doi:10.3390/biom10020172. PMC 7072473. PMID 31979037.

[29] Brennan, Patricia L. R.; Birkhead, Tim R.; Zyskowski, Kristof; Van Der Waag, Jessica; Prum, Richard O. (10 September 2008). "Independent evolutionary reductions of the phallus in basal birds". Journal of Avian Biology. 39 (5): 487–492. doi:10.1111/j.0908-8857.2008.04610.x.

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@@ Line 1: / Line 1: @@
 {{short description|Union of an egg and sperm to form a zygote within the female body}}
 {{Sex (biology) sidebar}}
-'''Internal fertilization''' is the union of an [[egg cell]] with a [[sperm]] during [[sexual reproduction]] inside the female body. Internal fertilization, unlike its counterpart, [[external fertilization]], brings more control to the female with reproduction.<ref name="Alonzo_2016">{{cite journal | vauthors = Alonzo SH, Stiver KA, Marsh-Rollo SE | title = Ovarian fluid allows directional cryptic female choice despite external fertilization | journal = Nature Communications | volume = 7 | issue = 1 | pages = 12452 | date = August 2016 | pmid = 27529581 | pmc = 4990696 | doi = 10.1038/ncomms12452 | bibcode = 2016NatCo...712452A }}</ref> For internal fertilization to happen there needs to be a method for the male to introduce the sperm into the female's reproductive tract. In [[mammal]]s, [[reptile]]s, and certain other groups of animals, this is done by [[copulation]], an [[intromittent organ]] being introduced into the [[vagina]] or [[cloaca]].<ref name="Hyman1992">{{cite book| vauthors = Hyman LH |title=Hyman's Comparative Vertebrate Anatomy|url=https://books.google.com/books?id=VKlWjdOkiMwC&q=fertilization|date=15 September 1992|publisher=University of Chicago Press|isbn=978-0-226-87013-7}}</ref><ref name="Austin" /> In most birds, the [[cloacal kiss]] is used, the two animals pressing their cloacas together while transferring sperm.<ref name="VB" /> [[Salamander]]s, [[spider]]s, some [[insect]]s and some [[mollusc]]s undertake internal fertilization by transferring a [[spermatophore]], a bundle of sperm, from the male to the female. Following fertilization, the embryos are laid as eggs in [[Oviparity|oviparous]] organisms, or continue to develop inside the reproductive tract of the mother to be born later as live young in [[Viviparity|viviparous]] organisms.
+'''Internal fertilization''' is the union of an [[egg cell|egg]] and [[sperm|sperm cell]] during [[sexual reproduction]] inside the female body. Internal fertilization, unlike its counterpart, [[external fertilization]], brings more control to the female with reproduction.<ref name="Alonzo_2016">{{cite journal | vauthors = Alonzo SH, Stiver KA, Marsh-Rollo SE | title = Ovarian fluid allows directional cryptic female choice despite external fertilization | journal = Nature Communications | volume = 7 | issue = 1 | pages = 12452 | date = August 2016 | pmid = 27529581 | pmc = 4990696 | doi = 10.1038/ncomms12452 | bibcode = 2016NatCo...712452A }}</ref> For internal fertilization to happen there needs to be a method for the male to introduce the sperm into the female's reproductive tract.
+Most [[Taxon|taxa]] that reproduce by internal fertilization are [[Gonochorism|gonochoric]].<ref name=":2" />{{Rp|124-125}} Male [[mammal]]s, [[reptile]]s, and certain other vertebrates transfer sperm into the female's [[vagina]] or [[cloaca]] through an [[intromittent organ]] during [[copulation (zoology)|copulation]].<ref>{{Cite book |last=Naguib |first=Marc |url=https://books.google.com/books?id=KgTeDwAAQBAJ&pg=PA65 |title=Advances in the Study of Behavior |date=2020-04-19 |publisher=Academic Press |isbn=978-0-12-820726-0 |language=en}}</ref><ref name="Hyman1992">{{cite book| vauthors = Hyman LH |title=Hyman's Comparative Vertebrate Anatomy|url=https://books.google.com/books?id=VKlWjdOkiMwC&q=fertilization|date=15 September 1992|publisher=University of Chicago Press|isbn=978-0-226-87013-7}}</ref><ref name="Austin" /> In most birds, the [[cloacal kiss]] is used, the two animals pressing their cloacas together while transferring sperm.<ref name="VB" /> [[Salamander]]s, [[spider]]s, some [[insect]]s and some [[mollusc]]s undertake internal fertilization by transferring a [[spermatophore]], a bundle of sperm, from the male to the female. Following fertilization, the embryos are laid as eggs in [[Oviparity|oviparous]] organisms, or continue to develop inside the reproductive tract of the mother to be born later as live young in [[Viviparity|viviparous]] organisms.
+== Evolution of internal fertilization ==
+{{Main|Evolution of sexual reproduction}}
+Internal fertilization evolved many times in [[animal]]s.<ref name=":2">{{Cite book|last1=Leonard|first1=Janet|url=https://books.google.com/books?id=PgtXj5R6OfMC&q=internal+fertilization+evolved|title=The Evolution of Primary Sexual Characters in Animals|last2=Cordoba-Aguilar|first2=Alex|date=2010-07-16|publisher=Oxford University Press|isbn=978-0-19-971703-3|language=en}}</ref>{{Rp|2}} According to [[David B. Dusenbery]] all the features with internal fertilization were most likely a result from [[oogamy]].<ref>{{Cite book|last=Dusenbery|first=David B.|url=https://books.google.com/books?id=9hgtEAAAQBAJ&q=Oogamy+internal+fertilization&pg=PA326|title=Living at Micro Scale: The Unexpected Physics of Being Small|date=2009|publisher=Harvard University Press|isbn=978-0-674-06021-0|pages=326|language=en}}</ref> It has been argued that internal fertilization evolve because of [[sexual selection]] through [[sperm competition]].<ref>{{Cite book|last=Smith|first=Robert L.|url=https://books.google.com/books?id=OIh4DX7tk_0C&q=evolution+of+gametes|title=Sperm Competition and the Evolution of Animal Mating systems|date=2012-12-02|publisher=Elsevier|isbn=978-0-323-14313-4|pages=7–8|language=en}}</ref>
+In [[amphibian]]s, internal fertilization evolved from [[external fertilization]].<ref>{{Cite book|last1=Sawada|first1=Hitoshi|url=https://books.google.com/books?id=Adm6BQAAQBAJ&q=internal+fertilization+evolved|title=Sexual Reproduction in Animals and Plants|last2=Inoue|first2=Naokazu|last3=Iwano|first3=Megumi|date=2014-02-07|publisher=Springer|isbn=978-4-431-54589-7|pages=97–98|language=en}}</ref>
 ==Methods of internal fertilization==
 [[Fertilization]] which takes place inside the female body is called internal fertilization in [[animals]] is done through the following different ways:<ref>{{cite journal| vauthors = Reichard UH |title=Monogamy—A variable relationship |journal=Max Planck Research |volume=3 |pages=62–7 |year=2002 |url=http://www.mpg.de/1028786/W001_Biology-Medicine_062_067.pdf |access-date=24 April 2013 |url-status=dead |archive-url=https://web.archive.org/web/20110514183645/http://www.mpg.de/1028786/W001_Biology-Medicine_062_067.pdf |archive-date=14 May 2011 }}</ref><ref name="Barash,Lipton,2001">{{cite book | vauthors = Lipton JE, Barash DP |title=The Myth of Monogamy: Fidelity and Infidelity in Animals and People |publisher=W.H. Freeman and Company |location=San Francisco |year=2001 |isbn=978-0-7167-4004-9 |url-access=registration |url=https://archive.org/details/isbn_9780716740049 }}</ref><ref name="Gowaty,Morell,1998">{{cite journal | vauthors = Morell V | title = A new look at monogamy | journal = Science | volume = 281 | issue = 5385 | pages = 1982–3 | date = September 1998 | pmid = 9767050 | doi = 10.1126/science.281.5385.1982 | s2cid = 31391458 }}</ref>
-* [[Copulation (zoology)|Copulation]],<ref name="Lombardi2012">{{cite book | vauthors = Lombardi J |title=Comparative Vertebrate Reproduction|url=https://books.google.com/books?id=tXvjBwAAQBAJ&q=%22internal+fertilization%22|date=6 December 2012|publisher=Springer Science & Business Media|isbn=978-1-4615-4937-6}}</ref> which involves the insertion of the [[penis]] or other [[intromittent organ]] into the [[vagina]] (in most [[mammal]]s) or to the [[cloaca]] in [[monotreme]]s, most [[reptile]]s, some [[bird]]s, the [[amphibian]] [[tailed frog]] and some [[fish]], the disappeared [[dinosaur]]s, as well as in other [[Invertebrate|non-vertebrate animals]].<ref name="Austin">{{Cite journal | doi=10.1080/11250008409439463|title = Evolution of the copulatory apparatus| journal=Bolletino di Zoologia| volume=51| issue=1–2| pages=249–269|year = 1984| vauthors = Austin CR | doi-access=free}}</ref><ref name="Diamond">{{Cite book| vauthors = Diamond J |author-link = Jared Diamond| title = The Rise and Fall of the Third Chimpanzee| year = 1991 | publisher =  Radius | pages = 360 pages<!-- Various pages in the book discuss all of these matters.-->|isbn = 978-0091742683|url=https://books.google.com/books?id=PQVFAQAAIAAJ}}</ref>
+* [[Copulation (zoology)|Copulation]],<ref name="Lombardi2012">{{cite book | vauthors = Lombardi J |title=Comparative Vertebrate Reproduction|url=https://books.google.com/books?id=tXvjBwAAQBAJ&q=%22internal+fertilization%22|date=6 December 2012|publisher=Springer Science & Business Media|isbn=978-1-4615-4937-6}}</ref> which involves the insertion of the [[penis]] or other [[intromittent organ]] into the [[vagina]] (in most [[mammal]]s)<ref>{{Cite book |last=Naguib |first=Marc |url=https://books.google.com/books?id=KgTeDwAAQBAJ&pg=PA65 |title=Advances in the Study of Behavior |date=2020-04-19 |publisher=Academic Press |isbn=978-0-12-820726-0 |language=en}}</ref> or to the [[cloaca]] in [[monotreme]]s, most [[reptile]]s, some [[bird]]s, the [[tailed frog]], some [[fish]], the disappeared [[dinosaur]]s, as well as in other [[Invertebrate|non-vertebrate animals]].<ref name="Austin">{{Cite journal | doi=10.1080/11250008409439463|title = Evolution of the copulatory apparatus| journal=Bolletino di Zoologia| volume=51| issue=1–2| pages=249–269|year = 1984| vauthors = Austin CR | doi-access=free}}</ref><ref name="Diamond">{{Cite book| vauthors = Diamond J |author-link = Jared Diamond| title = The Rise and Fall of the Third Chimpanzee| year = 1991 | publisher =  Radius | pages = 360 pages<!-- Various pages in the book discuss all of these matters.-->|isbn = 978-0091742683|url=https://books.google.com/books?id=PQVFAQAAIAAJ}}</ref>
 * [[Cloacal kiss]], which consists in that the two animals touch their cloacae together in order to transfer the [[sperm]] of the male to the female. It is used in most birds and in the [[tuatara]], that do not have an intromittent organ.<ref name=VB>{{cite book | vauthors = Romer AS, Parsons TS |year=1977 |title=The Vertebrate Body |publisher=Holt-Saunders International |location= Philadelphia, PA|pages= 396–399|isbn= 978-0-03-910284-5}}</ref>
-* Via [[spermatophore]], a sperm-containing cap placed by the male in the female's cloaca. Usually, the sperm is stored in spermathecae on the roof of the cloaca until it is needed at the time of oviposition. It is used by some [[salamander|salamander and newt species]], by the [[arachnida]], some insects and some [[mollusks]].<ref name="wedell">{{cite journal | vauthors = Wedell N, Tregenza T, Simmons LW | title = Nuptial gifts fail to resolve a sexual conflict in an insect | journal = BMC Evolutionary Biology | volume = 8 | pages = 204 | date = July 2008 | pmid = 18627603 | pmc = 2491630 | doi = 10.1186/1471-2148-8-204 }}</ref><ref>{{cite journal | vauthors = Sozou PD, Seymour RM | title = Costly but worthless gifts facilitate courtship | journal = Proceedings. Biological Sciences | volume = 272 | issue = 1575 | pages = 1877–84 | date = September 2005 | pmid = 16191592 | pmc = 1559891 | doi = 10.1098/rspb.2005.3152 }}</ref>
+* Via [[spermatophore]], a sperm-containing cap placed by the male in the female's cloaca. Usually, the sperm is stored in spermathecae on the roof of the cloaca until it is needed at the time of oviposition. It is used by some [[salamander|salamander and newt species]], by the [[Arachnida]], some insects and some [[mollusks]].<ref name="wedell">{{cite journal | vauthors = Wedell N, Tregenza T, Simmons LW | title = Nuptial gifts fail to resolve a sexual conflict in an insect | journal = BMC Evolutionary Biology | volume = 8 | pages = 204 | date = July 2008 | issue = 1 | pmid = 18627603 | pmc = 2491630 | doi = 10.1186/1471-2148-8-204 | doi-access = free | bibcode = 2008BMCEE...8..204W }}</ref><ref>{{cite journal | vauthors = Sozou PD, Seymour RM | title = Costly but worthless gifts facilitate courtship | journal = Proceedings. Biological Sciences | volume = 272 | issue = 1575 | pages = 1877–84 | date = September 2005 | pmid = 16191592 | pmc = 1559891 | doi = 10.1098/rspb.2005.3152 }}</ref>
-* In [[sponges]], sperm cells are released into the water to fertilize ova that in some species are also released (external fertilization) and in others are retained by the "mother" (internal fertilization).<ref>{{cite book | vauthors = Bergquist PR | date = 1978 | title = Sponges | location = London | publisher = Hutchinson | isbn = 9780520036581 | url = https://books.google.com/books?id=MWtPaEo398YC }}</ref>
+* In [[sponges]], sperm cells are released into the water to fertilize ova that are retained by the female. Some species of sponge participate in external fertilization where the ova is released.<ref>{{cite book | vauthors = Bergquist PR | date = 1978 | title = Sponges | location = London | publisher = Hutchinson | isbn = 9780520036581 | url = https://books.google.com/books?id=MWtPaEo398YC }}</ref>
 ==Expulsion==
@@ Line 15: / Line 23: @@
 At some point, the growing [[egg (biology)|egg]] or [[offspring]] must be expelled. There are several possible [[modes of reproduction]]. These are traditionally classified as follows:
-* [[Oviparity]], as in most [[insect]]s and reptiles, [[monotreme]]s, dinosaurs and all birds which lay eggs that continue to develop after being laid, and hatch later.<ref>{{cite book | author-link1 = Thierry Lodé | vauthors = Lodé T | date = 2001 | title = Les stratégies de reproduction des animaux | trans-title = Reproduction Strategies in the Animal Kingdom | language = French | publisher = Dunod Sciences | location = Paris }}</ref>
+* [[Oviparity]], as in most [[invertebrate]]s and reptiles, [[monotreme]]s, dinosaurs and all birds which lay eggs that continue to develop after being laid, and hatch later.<ref>{{cite book | author-link1 = Thierry Lodé | vauthors = Lodé T | date = 2001 | title = Les stratégies de reproduction des animaux | trans-title = Reproduction Strategies in the Animal Kingdom | language = French | publisher = Dunod Sciences | location = Paris }}</ref>
-* [[Viviparity]], as in almost all mammals (such as [[whale]]s, [[kangaroo]]s and [[human]]s) which bear their young live. The developing young spend proportionately more time within the female's reproductive tract.  The young are later released to survive on their own, with varying amounts of help from the [[parent]] (s) of the [[species]].<ref>{{cite journal | vauthors = Blackburn DG | title = Classification of the reproductive patterns of amniotes. | journal = Herpetological Monographs | date = January 2000 | volume = 14 | pages = 371–7 | doi = 10.2307/1467051 | jstor = 1467051 | url = https://www.researchgate.net/publication/233720323 }}</ref>
+* [[Viviparity]], as in almost all mammals (such as [[whale]]s, [[kangaroo]]s and [[human]]s) which bear their young live. The developing young spend proportionately more time within the female's reproductive tract.  The young are later released to survive on their own, with varying amounts of help from the [[parent]](s) of the [[species]].<ref>{{cite journal | vauthors = Blackburn DG | title = Classification of the reproductive patterns of amniotes. | journal = Herpetological Monographs | date = January 2000 | volume = 14 | pages = 371–7 | doi = 10.2307/1467051 | jstor = 1467051 | url = https://www.researchgate.net/publication/233720323 }}</ref>
 * [[Ovoviviparity]], as in the [[garter snake]], most [[Viperidae|vipers]], and the [[Madagascar hissing cockroach]], which have eggs (with [[Eggshell|shells]]) that hatch as they are laid, making it resemble live birth.<ref>{{cite book |title=Biology of Sharks and Their Relatives | veditors = Carrier JC, Musick JA, Heithaus MR |year=2012 |publisher=CRC Press |isbn=978-1439839249 |pages=296–301}}</ref>
-== Advantages from Internal Fertilization ==
+== Advantages to internal fertilization ==
 Internal fertilization allows for:
 * Female [[mate choice]], which gives the female the ability to choose her partner before and after mating. The female cannot do this with external fertilization because she may have limited control of who is fertilizing her eggs, and when they are being fertilized.<ref name="Alonzo_2016" />
-* Making a decision for the conditions of reproduction, like location and time.<ref>{{cite journal | vauthors = Yokoe M, Takayama-Watanabe E, Saito Y, Kutsuzawa M, Fujita K, Ochi H, Nakauchi Y, Watanabe A | display-authors = 6 | title = A Novel Cysteine Knot Protein for Enhancing Sperm Motility That Might Facilitate the Evolution of Internal Fertilization in Amphibians | journal = PLOS ONE | volume = 11 | issue = 8 | pages = e0160445 | date = 2016-08-31 | pmid = 27579691 | pmc = 5007030 | doi = 10.1371/journal.pone.0160445 | bibcode = 2016PLoSO..1160445Y | veditors = Klymkowsky M }}</ref> In external fertilization a female can only choose the time in which she releases her eggs, but not when they are fertilized. This is similar, in ways, to [[cryptic female choice]].
+* Making a decision for the conditions of reproduction, like location and time.<ref name="cysteine knot">{{cite journal | vauthors = Yokoe M, Takayama-Watanabe E, Saito Y, Kutsuzawa M, Fujita K, Ochi H, Nakauchi Y, Watanabe A | display-authors = 6 | title = A Novel Cysteine Knot Protein for Enhancing Sperm Motility That Might Facilitate the Evolution of Internal Fertilization in Amphibians | journal = PLOS ONE | volume = 11 | issue = 8 | pages = e0160445 | date = 2016-08-31 | pmid = 27579691 | pmc = 5007030 | doi = 10.1371/journal.pone.0160445 | bibcode = 2016PLoSO..1160445Y | veditors = Klymkowsky M | doi-access = free }}</ref> In external fertilization a female can only choose the time in which she releases her eggs, but not when they are fertilized. This is similar, in ways, to [[cryptic female choice]].
 * Egg protection on dry land.<ref>{{cite journal | vauthors = Altig R, McDiarmid RW | title = Morphological diversity and evolution of egg and clutch structure in amphibians. | journal = Herpetological Monographs | date = December 2007 | volume = 21 | issue = 1 | pages = 1–32 | doi = 10.1655/06-005.1 | s2cid = 55728625 }}</ref> While oviparous animals either have a jelly like ovum or a hard shell enclosing their egg, internally fertilizing animals grow their eggs and offspring inside themselves. This offers protection from predators and from dehydration on land.<ref>{{Cite web|date=2018-07-17|title=43.2A: External and Internal Fertilization |url= https://bio.libretexts.org/Bookshelves/Introductory_and_General_Biology/Book%3A_General_Biology_(Boundless)/43%3A_Animal_Reproduction_and_Development/43.2%3A_Fertilization/43.2A%3A_External_and_Internal_Fertilization|access-date=2020-11-09|website=Biology LibreTexts|language=en}}</ref> This allows for a higher chance of survival when there is a regulated temperature and protected area within the mother.
-== Disadvantages to Internal Fertilization ==
+== Disadvantages to internal fertilization ==
-* [[Gestation]] can and will add additional risks for the mother.<ref name=":1">{{cite journal | vauthors = Wallen K, Zehr JL | title = Hormones and history: the evolution and development of primate female sexuality | journal = Journal of Sex Research | volume = 41 | issue = 1 | pages = 101–12 | date = February 2004 | pmid = 15216429 | pmc = 1255935 | doi = 10.1080/00224490409552218 }}</ref> The additional risks from gestation come from extra energy demands.
+* [[Gestation]] can and will add additional risks for the mother.<ref name=":1">{{cite journal | vauthors = Wallen K, Zehr JL | title = Hormones and history: the evolution and development of primate female sexuality | journal = Journal of Sex Research | volume = 41 | issue = 1 | pages = 101–12 | date = February 2004 | pmid = 15216429 | pmc = 1255935 | doi = 10.1080/00224490409552218 }}</ref> The additional risks from gestation come from extra energy demands.
-* Along with internal fertilization comes [[sexual reproduction]] , in most cases. Sexual reproduction comes with some risks as well. The risks with sexual reproduction are with intercourse, it is infrequent and only works well during peak [[fertility]]. While animals which externally fertilize are able to release egg and sperm, usually into the water, not needing a specific partner to reproduce.<ref name=":1" />
+* Along with internal fertilization comes [[sexual reproduction]], in most cases. Sexual reproduction comes with some risks as well. The risks with sexual reproduction are with intercourse, it is infrequent and only works well during peak [[fertility]]. While animals which externally fertilize are able to release egg and sperm, usually into the water, not needing a specific partner to reproduce.<ref name=":1" />
-*Fewer offspring are produced through internal fertilization in comparison to external fertilization. This is both because the mother cannot hold and grow as many offspring as eggs, and the mother cannot provide and obtain enough resources for a larger amount of offspring.<ref>{{Cite journal|last=Parker|first=G. A.|date=1970|title=Sperm Competition and Its Evolutionary Consequences in the Insects|url=https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1469-185X.1970.tb01176.x|journal=Biological Reviews|language=en|volume=45|issue=4|pages=525–567|doi=10.1111/j.1469-185X.1970.tb01176.x|issn=1469-185X}}</ref>
+*Fewer offspring are produced through internal fertilization in comparison to external fertilization. This is both because the mother cannot hold and grow as many offspring as eggs, and the mother cannot provide and obtain enough resources for a larger amount of offspring.<ref>{{Cite journal|last=Parker|first=G. A.|date=1970|title=Sperm Competition and Its Evolutionary Consequences in the Insects|url=https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1469-185X.1970.tb01176.x|journal=Biological Reviews|language=en|volume=45|issue=4|pages=525–567|doi=10.1111/j.1469-185X.1970.tb01176.x|s2cid=85156929|issn=1469-185X}}</ref>
 == Fish ==
-Some species of fish like guppies have the ability to internally fertilize, this process happens by the male inserting a tubular fin into the female’s reproductive opening and then will deposit sperm into her reproductive tract. There are other species of fish that are mouthbrooders which means that one fish puts the eggs in its mouth for incubation. A certain type of fish that is a mouthbrooder is called cichlids and many of them are maternal mouthbrooders. The process for this is the female would lay the egg and pick it up in her mouth. Then the males will encourage the female to open her mouth so they can fertilize the eggs while it’s in the female’s mouth.<ref name=":0">{{Cite book|last=Cotner|first=Sehoya|url=https://open.lib.umn.edu/evolutionbiology/|title=The Evolution and Biology of Sex|last2=Wassenberg|first2=Deena|language=en}}</ref> Internal fertilization in cartilaginous fishes contains the same evolutionary origin as reptiles, birds, and mammals that internally fertilize. Also in these internally fertilizing fish while the sperm is transferred to the reproductive tract there is no noticeable change in tonality. <ref>Engel, Kathrin M, Dzyuba, Viktoriya, Ninhaus-Silveira, Alexandre, Veríssimo-Silveira, Rosicleire, Dannenberger, Dirk, Schiller, Jürgen, . . . Dzyuba, Borys. (2020). Sperm Lipid Composition in Early Diverged Fish Species: Internal vs. External Mode of Fertilization. ''Biomolecules (Basel, Switzerland),'' ''10''(2), 172.</ref>
+Some species of fish like guppies have the ability to internally fertilize, this process happens by the male inserting a tubular fin into the female's reproductive opening and then will deposit sperm into her reproductive tract. There are other species of fish that are mouthbrooders which means that one fish puts the eggs in its mouth for incubation. A certain type of fish that is a mouthbrooder is called cichlids and many of them are maternal mouthbrooders. The process for this is the female would lay the egg and pick it up in her mouth. Then the males will encourage the female to open her mouth so they can fertilize the eggs while it is in the female's mouth.<ref name=":0">{{Cite book|last1=Cotner|first1=Sehoya|url=https://open.lib.umn.edu/evolutionbiology/|title=The Evolution and Biology of Sex|last2=Wassenberg|first2=Deena|year=2020|language=en}}</ref> Internal fertilization in cartilaginous fishes contains the same evolutionary origin as reptiles, birds, and mammals that internally fertilize. Also in these internally fertilizing fish while the sperm is transferred to the reproductive tract there is no noticeable change in tonality.<ref>{{Cite journal |last1=Engel |first1=Kathrin M |last2=Dzyuba |first2=Viktoriya |last3=Ninhaus-Silveira |first3=Alexandre |last4=Veríssimo-Silveira |first4=Rosicleire |last5=Dannenberger |first5=Dirk |last6=Schiller |first6=Jürgen |last7=Steinbach |first7=Christoph |last8=Dzyuba |first8=Borys |date=February 2020 |title=Sperm Lipid Composition in Early Diverged Fish Species: Internal vs. External Mode of Fertilization |journal=[[Biomolecules (journal)|Biomolecules]] |volume=10 |issue=2 |pages=172 |doi=10.3390/biom10020172 |pmid=31979037 |pmc=7072473 |doi-access=free}}</ref>
 == Amphibians ==
-Most amphibians have external fertilization but there is an exception to some like salamanders which mostly have internal fertilization. Male salamanders don't have a penis to insert and deposit sperm inside females. Their alternate for that consists of an encased capsule of sperm and nutrients called a spermatophore. The male will deposit a spermatophore on the ground and the female will pick it up with her cloaca (a combined urinary and genital opening) and fertilize her eggs with it.<ref name=":0" /> Over time amphibians have been found evolving to increasing internal fertilization. Within amphibians, it is common for high vertebrates to internally fertilize because of the transition from water to land during vertebrate evolution. There is an advantage for the amphibians who are internally fertilizing allowing for the selection of a time and place for reproduction. <ref>Yokoe, Misato, Takayama-Watanabe, Eriko, Saito, Yoko, Kutsuzawa, Megumi, Fujita, Kosuke, Ochi, Haruki, . . . Watanabe, Akihiko. (2016). A Novel Cysteine Knot Protein for Enhancing Sperm Motility That Might Facilitate the Evolution of Internal Fertilization in Amphibians. PloS One, 11(8), E0160445.</ref>
+Most amphibians have external fertilization but there is an exception to some like salamanders which mostly have internal fertilization. Salamanders do not use intercourse for sexual reproduction due to their lack of external penis. Rather, the male salamander produces an encased capsule of sperm and nutrients called a [[spermatophore]]. The male deposits a spermatophore on the ground and the female will pick it up with her cloaca (a combined urinary and genital opening) and fertilize her eggs with it.<ref name=":0" /> Over time amphibians have been found evolving to increasing internal fertilization.{{Citation needed|date=October 2022}} Within amphibians, it is common for high vertebrates to internally fertilize because of the transition from water to land during vertebrate evolution. There is an advantage for the amphibians who are internally fertilizing allowing for the selection of a time and place for reproduction.<ref name="cysteine knot" />
 == Birds ==
-The male species tends to lack penises so they reproduce by internally fertilizing. Birds use their cloacal contact which is the birds kissing. The cloaca is their mouth so to start the process they connect their cloacas and the male transfer's the sperm to the female's mouth. Although most birds don't have a penis some bird species do like ducks and geese and use their penis for internal fertilization. <ref name=":0" /> Being the only vertebrate taxon with internal fertilization, the bird’s intromittent phallus has been lost in most species. The phallus is a penis and being an evolution without this makes them have to be internally fertilized. <ref>Brennan, Patricia L. R, Birkhead, Tim R, Zyskowski, Kristof, Van Der Waag, Jessica, & Prum, Richard O. (2008). Independent evolutionary reductions of the phallus in basal birds. Journal of Avian Biology, 39(5), 487-492.</ref>
+Most birds do not have penises, but achieve internal fertilization via cloacal contact (or "cloaca kiss"). In these birds, males and females contact their cloacas together, typically briefly, and transfer sperm to the female. However, water fowls such as ducks and geese have penises and are able to use them for internal fertilization. <ref name=":0" /> While birds have internal fertilization, most species no longer have phallus structures. This makes them the only vertebrate taxon to fall into both categories of lacking the phallus but participating in internal fertilization.<ref>{{Cite journal |last1=Brennan |first1=Patricia L. R. |last2=Birkhead |first2=Tim R. |last3=Zyskowski |first3=Kristof |last4=Van Der Waag |first4=Jessica |last5=Prum |first5=Richard O. |date=10 September 2008 |title=Independent evolutionary reductions of the phallus in basal birds |journal=[[Journal of Avian Biology]] |volume=39 |issue=5 |pages=487–492 |doi=10.1111/j.0908-8857.2008.04610.x |doi-access=free}}</ref>
 == See also ==

v t e Modes of reproduction
Modes	Oviparity Viviparity (Histotrophic, Hemotrophic) Ovoviviparity
Fertilisation	External fertilization Internal fertilization
Parental care	birds fish (mouthbrooding) humans mammals
Related topics	Adelphophagy Gastric-brooding frog Live-bearing aquarium fish Male pregnancy Matrotrophy Oophagy Pregnancy in fish Trophic egg