Editing Gynandromorphism

{{Short description|Organism with both male and female characteristics}}
[[File:Open wing basking position of Gynandromorph of Athyma inara Westwood, 1850 - Colour Sergeant butterfly MG 0820 WLB.jpg|thumb|Gynandromorph of ''[[Athyma inara]] inara'']]
[[File:ZwitterHauhechelblaeuling.jpg|thumb|Gynandromorph of the [[common blue]] butterfly (''Polyommatus icarus'')]]

[[File:Heteropteryx dilatata 0034b L.D.jpg|right|thumb|Gynandromorph of ''[[Heteropteryx dilatata]]'']]
[[File:Gynandromorphism in Crocothemis servilia.jpg|thumb|Gynandromorph of ''[[Crocothemis servilia]]'']]

'''Gynandromorphism''' is the phenomenon that occurs when an individual organism possesses both [[male]] and [[female]] [[Phenotype|phenotypes]] due to [[Chimera (genetics)|genetic chimera]] of [[Sex chromosome|sex chromosomes]] in cells across the body and is most easily recognized in [[species]] that display [[sexual dimorphism]]<ref name=":0">{{Cite journal |last1=Jahner |first1=Joshua P. |last2=Lucas |first2=Lauren K. |last3=Wilson |first3=Joseph S. |last4=Forister |first4=Matthew L. |date=5 April 2015 |title=Morphological Outcomes of Gynandromorphism in Lycaeides Butterflies (Lepidoptera: Lycaenidae) |url=https://academic.oup.com/jinsectscience/article/15/1/38/2583133#202765122 |journal=Journal of Insect Science |volume=15 |issue=1 |pages=38 |doi=10.1093/jisesa/iev020 |pmid=25843591 |via=Oxford Academic|pmc=7175718 }}</ref>'''.''' An individual who displays this characteristic is called a '''gynandromorph.''' The term comes from the Greek γυνή (''gynē'') 'female', ἀνήρ (''anēr'') 'male', and μορφή (''morphē'') 'form', and is most commonly documented the field of [[entomology]].<ref name=":0" /> The definition of “gynandromorphism” is distinct from both [[Intersex|intersexuality]] and [[Hermaphrodite|hermaphroditism]], although they are sometimes used interchangeably.<ref name=":1">{{Cite journal |last1=Fusco |first1=Giuseppe |last2=Minelli |first2=Alessandro |date=12 January 2023 |title=Descriptive versus causal morphology: gynandromorphism and intersexuality |url=https://link.springer.com/article/10.1007/s12064-023-00385-1#Abs1 |journal=Theory in Bioscience |volume=142 |pages=1–11 |doi=10.1007/s12064-023-00385-1 |pmid=36633802 |via=Springer Nature Link|hdl=11577/3466453 |hdl-access=free }}</ref>

== Occurrence ==
Gynandromorphism has been noted in [[Lepidoptera]] (butterflies and moths) since the 1700s.<ref name="Rudolphi">{{cite journal |last1=Rudolphi |first1=Karl Asmund |title=Beschreibung einer seltenen menschlichen Zwitterbildung nebst vorangeschickten allgemeinen Bemerkungen über Zwitter-Thiere |journal=Abhandlungen der Königlichen Akademie der Wissenschaften zu Berlin |date=1828 |pages=45–69 |url=https://books.google.com/books?id=2WFDAQAAMAAJ&pg=PA45 |language=de}}</ref><ref name="Packard1875">{{cite journal |last1=Packard |first1=Alpheus Spring |title=On Gynandromorphism in the Lepidoptera |journal=Memoirs Read Before the Boston Society of Natural History |date=1875 |volume=2 |pages=409–412 |url=https://archive.org/details/memoirsreadbefor02bost/page/n443/mode/2up}}</ref><ref name="Pavid">{{cite web |last1=Pavid |first1=Katie |title=Beauty of the dual-gender butterfly |url=https://www.nhm.ac.uk/discover/beauty-dual-gender-butterfly.html |publisher=Natural History Museum |access-date=11 May 2021}}</ref> It has also been observed in [[crustaceans]], such as [[lobsters]] and [[crabs]], in [[spider]]s,<ref>{{Cite journal|last1=Suzuki|first1=Yuya|last2=Kuramitsu|first2=Kazumu|last3=Yokoi|first3=Tomoyuki|date=2019-06-14|title=Morphology and sex-specific behavior of a gynandromorphic ''Myrmarachne formicaria'' (Araneae: Salticidae) spider|journal=The Science of Nature|language=en|volume=106|issue=7|pages=34|doi=10.1007/s00114-019-1625-x|pmid=31201570|bibcode=2019SciNa.106...34S|s2cid=189819156|issn=1432-1904|hdl=2241/00159248|url=https://tsukuba.repo.nii.ac.jp/?action=repository_uri&item_id=53830|hdl-access=free}}</ref> [[tick]]s,<ref name="Labruna">{{cite journal |last1=Labruna |first1=M. B. |last2=Ribeiro |first2=A. F. |last3=Cruz |first3=M. V. |last4=Camargo |first4=L. M A. |last5=Camargo |first5=E. P. |title=Gynandromorphism in ''Amblyomma cajennense'' and ''Rhipicephalus sanguineus'' (Acari: Ixodidae) |journal=Journal of Parasitology |date=August 2002 |volume=88 |issue=4 |pages=810–811 |doi=10.1645/0022-3395(2002)088[0810:GIACAR]2.0.CO;2|pmid=12197141 |s2cid=26299983 }}</ref> [[fly|flies]],<ref name="Morgan1914">{{cite journal |last1=Morgan |first1=T. H. |title=Mosaics and gynandromorphs in Drosophila |journal=Experimental Biology and Medicine |date=1 June 1914 |volume=11 |issue=6 |pages=171–172 |doi=10.3181/00379727-11-105|s2cid=87401593 }}</ref> [[locust]]s,<ref name="Maeno">{{cite journal |last1=Maeno |first1=Koutaro |last2=Tanaka |first2=Seiji |title=Morphological and behavioural characteristics of a gynandromorph of the desert locust, Schistocerca gregaria |journal=Physiological Entomology |date=September 2007 |volume=32 |issue=3 |pages=294–299 |doi=10.1111/j.1365-3032.2007.00573.x|s2cid=85317122 }}</ref> [[Cricket (insect)|crickets]],<ref name="Taniyama">{{cite journal |last1=Taniyama |first1=Katsuya |last2=Onodera |first2=Kaori |last3=Tanaka |first3=Kazuhiro |title=Sexual identity and sexual attractiveness of a gynandromorph of the lawn ground cricket, Polionemobius mikado (Orthoptera: Trigonidiidae): Gynandromorph of Polionemobius mikado |journal=Entomological Science |date=December 2018 |volume=21 |issue=4 |pages=423–427 |doi=10.1111/ens.12321|s2cid=91381517 }}</ref> [[dragonfly|dragonflies]],<ref name="Renjith2020">{{cite journal |last1=Renjith |first1=R. V. |last2=Chandran |first2=A. Vivek |title=A record of gynandromorphism in the libellulid dragonfly Crocothemis servilia (Insecta: Odonata) from India |journal=Journal of Threatened Taxa |date=26 June 2020 |volume=12 |issue=9 |pages=16183–16186 |doi=10.11609/jott.5322.12.9.16183-16186|doi-access=free }}</ref> [[ant]]s,<ref name="Donisthorpe">{{cite journal |last1=Donisthorpe |first1=Horace |title=Gynandromorphism in ants |journal=Zoologischer Anzeiger |date=1929 |volume=82 |pages=92–96 |url=http://antbase.org/ants/publications/5762/5762.pdf}}</ref><ref name="Cokendolpher">{{cite journal |last1=Cokendolpher |first1=James C. |last2=Francke |first2=Oscar F. |title=Gynandromorphic Desert Fire Ant, ''Solenopsis aurea'' Wheeler (Hymenoptera: Formicidae) |journal=Journal of the New York Entomological Society |date=1983 |volume=91 |issue=3 |pages=242–245 |jstor=25009362 |url=https://www.jstor.org/stable/25009362 |issn=0028-7199}}</ref> [[termite]]s,<ref name="Miyaguni2017">{{cite journal |last1=Miyaguni |first1=Yasushi |last2=Nozaki |first2=Tomonari |last3=Yashiro |first3=Toshihisa |title=The first report of gynandromorphy in termites (Isoptera; Kalotermitidae; Neotermes koshunensis) |journal=The Science of Nature |date=August 2017 |volume=104 |issue=7–8 |pages=60 |doi=10.1007/s00114-017-1478-0|pmid=28676938 |bibcode=2017SciNa.104...60M |s2cid=21170853 }}</ref> [[bee]]s,<ref name="Lucia2013">{{cite journal |last1=Lucia |first1=Mariano |last2=Gonzalez |first2=Victor. H. |title=A New Gynandromorph of ''Xylocopa frontalis'' with a Review of Gynandromorphism in ''Xylocopa'' (Hymenoptera: Apidae: Xylocopini) |journal=Annals of the Entomological Society of America |date=1 November 2013 |volume=106 |issue=6 |pages=853–856 |doi=10.1603/AN13085|s2cid=84567180 |doi-access=free |hdl=11336/23238 |hdl-access=free }}</ref> [[lizard]]s,<ref name="Mitchell1978">{{cite journal |last1=Mitchell |first1=Joseph C. |last2=Fouquette |first2=M. J. |title=A Gynandromorphic Whiptail Lizard, Cnemidophorus inornatus, from Arizona |journal=Copeia |date=10 February 1978 |volume=1978 |issue=1 |pages=156 |doi=10.2307/1443840|jstor=1443840 }}</ref> [[snake]]s,<ref name="Krohmer1989">{{cite journal |last1=Krohmer |first1=Randolph W. |title=Reproductive Physiology and Behavior of a Gynandromorph Redsided Garter Snake, Thamnophis sirtalis parietalis, from Central Manitoba, Canada |journal=Copeia |date=27 December 1989 |volume=1989 |issue=4 |pages=1064–1068 |doi=10.2307/1446001|jstor=1446001 }}</ref> [[rodent]]s,<ref name="Asdell1942">{{cite journal |last1=Asdell |first1=S. A. |title=The Accessory Reproductive Tract in Mammalian True Hermaphrodites, an Effect of Position |journal=The American Naturalist |date=1942 |volume=76 |issue=762 |pages=75–84 |doi=10.1086/281015 |jstor=2457667 |bibcode=1942ANat...76...75A |s2cid=83563917 |url=https://www.jstor.org/stable/2457667 |issn=0003-0147|url-access=subscription }}</ref><ref name="Hollander1956">{{cite journal |last1=Hollander |first1=W. F. |last2=Gowen |first2=John W. |last3=Stadler |first3=Janice |title=A study of 25 gynandromorphic mice of the Bagg albino strain |journal=The Anatomical Record |date=February 1956 |volume=124 |issue=2 |pages=223–243 |doi=10.1002/ar.1091240207|pmid=13302819 |s2cid=21938150 }}</ref> and [[bird]]s.<ref>{{Cite web |last=National Geographic |title=Rare Half Male, Half Female Cardinal Spotted in Pennsylvania |website= [[National Geographic Society]]|url=https://www.nationalgeographic.com/animals/article/half-male-half-female-cardinal-pennsylvania |date=2019-07-01 |access-date=2025-03-19}}</ref><ref>{{cite journal |pages=7730–5 |doi=10.1073/pnas.0408350102 |title=Sexually dimorphic expression of trkB, a Z-linked gene, in early posthatch zebra finch brain |year=2005 |first1=Xuqi |last1=Chen |first2=Robert J. |last2=Agate |first3=Yuichiro |last3=Itoh |first4=Arthur P. |last4=Arnold |journal=Proceedings of the National Academy of Sciences |volume=102 |issue=21 |pmid=15894627 |pmc=1140405|bibcode=2005PNAS..102.7730C |doi-access=free}}
*{{cite magazine |author=Laura Wright |date=March 25, 2003 |title=Unique Bird Sheds Light on Sex Differences in the Brain |magazine=[[Scientific American]] |url=http://www.scientificamerican.com/article.cfm?id=unique-bird-sheds-light-o}}</ref><ref>[[Gouldian finch]]
''Erythrura gouldiae'' [http://users.skynet.be/fa398872/engoulm10.htm Gynandromorph] {{webarchive|url=https://web.archive.org/web/20060716013321/http://users.skynet.be/fa398872/engoulm10.htm |date=2006-07-16 }}</ref><ref>Powdermill Banding [http://www.westol.com/~banding/Pictorial_Highlights_090605.html Fall 2004] {{webarchive|url=https://web.archive.org/web/20061231235033/http://www.westol.com/~banding/Pictorial_Highlights_090605.html |date=2006-12-31 }}</ref><ref>A Gender-bender Colored Cardinal, by Tim Wall, Discovery News, 31 May 2011 [http://news.discovery.com/animals/bi-color-cardinal-mystery-110531.html] {{Webarchive|url=https://web.archive.org/web/20120930101733/http://news.discovery.com/animals/bi-color-cardinal-mystery-110531.html|date=2012-09-30}}</ref><ref>{{cite news|url=http://news.bbc.co.uk/2/hi/uk_news/scotland/edinburgh_and_east/8561814.stm|title=Half-cock chicken mystery solved|date=11 March 2010|work=BBC News}}</ref> Although it can be seen in a variety of species, gynandromorphism is notably uncommon.<ref>{{Cite journal |last1=Martens |first1=Andreas |last2=Wildermuth |first2=Hansruedi |date=1 June 2021 |title=Gynandromorphism and intersexuality in Odonata: a review |url=https://bioone.org/journals/odonatologica/volume-50/issue-1-2/zenodo.4746242/Gynandromorphism-and-intersexuality-in-Odonata-a-review/10.5281/zenodo.4746242.short |journal=Odonatologica |volume=50 |issue=1–2 |pages=65–80 |doi=10.5281/zenodo.4746242.short |doi-broken-date=27 April 2025 |via=BioOne Digital Library}}</ref> Reporting depends on ease of detecting the phenomenon (whether a species displays noticeable sexual dimorphism) and how well-studied a region or organism is. For example, up until 2023 gynandromorphism had been reported in more than 40 bird species, but the vast majority of these are from the [[Palearctic]] and [[Nearctic]]; meanwhile, incidents in species from other regions may go underreported due to lack of data collection.<ref name=Stepniewski2023>{{cite journal| last1=Stępniewski | first1=J. | last2=Surmacki | first2=A. | year=2023 | title=The first case of bilateral gynandromorphic plumage type in the bearded reedling Panurus biarmicus | journal=The European Zoological Journal | volume=90 | issue=2 | pages=643–647 | doi=10.1080/24750263.2023.2231000 | doi-access=free }}</ref>

== Pattern of distribution of male and female tissues in a single organism ==

Several patterns of tissue distribution occur amongst observed gynandromorphs. Patch-like patterns may arise, but commonly the phenotype presents in a symmetrical pattern, of which there are three main types: [[Anatomical terms of location#Medial and lateral|bilateral]], oblique, and transverse.<ref name=":1" /> Bilateral describes an organism that is split laterally with one side possessing female characteristics and the other having male characteristics. Oblique refers to a diagonal line across the [[sagittal plane]] that separates the male and female phenotypes. Lastly, transverse is a separation of male and female phenotypes along an axis that intersects with what would be considered the primary axis for the body of a given organism.<ref name=":1" /> 

A notable example in birds is the [[zebra finch]]. These birds have lateralized brain structures in the face of a common [[steroid]] signal, providing strong evidence for a non-hormonal primary sex mechanism regulating brain differentiation.<ref>{{cite journal |pages=701–8 |doi=10.1038/nrn1494 |title=Sex chromosomes and brain gender |year=2004 |last1=Arnold |first1=Arthur P. |journal=Nature Reviews Neuroscience |volume=5 |issue=9 |pmid=15322528|s2cid=7419814 }}</ref>

<gallery mode="packed">
Papilioandrogeusfemale.jpg|Normal female of ''[[Papilio androgeus]]''
Papilioandrogeusgynandromorph.jpg|Mosaic gynandromorph of ''Papilio androgeus''
Papilioandrogeosmale.jpg|Normal male of ''Papilio androgeus''
</gallery>

== Causes ==
The exact cause for gynandromorphism is unknown, and appears to vary by species.<ref name=":0" /> One proposed cause for the phenotype in birds is a disruption that occurs in [[meiosis]] in female birds -- who possess [[ZW sex-determination system|ZW chromosomes]] -- in which [[Polar body|polar bodies]] are not forced out of the cell.<ref name=":2">{{Cite journal |last1=Morris |first1=Kirsten R |last2=Hirst |first2=Claire E. |last3=Major |first3=Andrew T. |last4=Ezaz |first4=Tariq |last5=Ford |first5=Mark |last6=Bibby |first6=Susan |last7=Doran |first7=Tim J. |last8=Smith |first8=Craig A. |date=15 August 2018 |title=Gonadal and Endocrine Analysis of a Gynandromorphic Chicken |url=https://academic.oup.com/endo/article/159/10/3492/5074254 |journal=Endocrinology |volume=159 |issue=10 |pages=3492–3502 |doi=10.1210/en.2018-00553 |pmid=30124802 |via=Oxford Academic}}</ref> This is then followed with [[Fertilisation|fertilization]] by two Z-bearing [[Sperm|sperm cells]] which creates cells in the embryo containing both ZZ and ZW chromosomes.<ref name=":2" /> The same phenomenon can be seen in other species, such as ''[[Drosophila melanogaster]]'', and creates a [[Mosaic (genetics)|genetic mosaic]] which gives rise to the mixed phenotype.

== As a research tool ==
Gynandromorphs occasionally afford a powerful tool in genetic, developmental, and behavioral analyses. In ''[[Drosophila melanogaster]]'', for instance, they provided evidence that male courtship behavior originates in the brain,<ref>{{Cite journal|last=Hotta|first=Y, and Benzer, S.|date=1972|title=Mapping of Behaviour in Drosophila mosaics|journal=Nature|volume=240|issue=5383|pages=527–535|doi=10.1038/240527a0|pmid=4568399|bibcode=1972Natur.240..527H|s2cid=4181921}}</ref> that males can distinguish conspecific females from males by the scent or some other characteristic of the posterior, dorsal, [[integument]] of females,<ref>{{Cite journal|last=Nissani|first=M.|date=1975|title=A new behavioral bioassay for an analysis of sexual attraction and pheromones in insects|journal=Journal of Experimental Zoology|volume=192 |issue=2|pages=271–5|doi=10.1002/jez.1401920217|pmid=805823|bibcode=1975JEZ...192..271N }}</ref><ref>{{Cite journal|last=Hotta|first=Y., Benzer, S.|date=1976|title=Courtship in Drosophila mosaics: sex-specific foci for sequential action patterns|journal=Proc Natl Acad Sci U S A|volume=73 |issue=11|pages=4154–4158|doi=10.1073/pnas.73.11.4154|pmid=825859|bibcode=1976PNAS...73.4154H|pmc=431365|doi-access=free}}</ref> that the [[germ cell]]s originate in the posterior-most region of the [[blastoderm]],<ref>{{Cite journal|date=1977|title=Cell lineage analysis of germ cells of Drosophila melanogaster|journal=Nature|volume=265|issue=5596|pages=729–731|doi=10.1038/265729a0|pmid=404558|bibcode=1977Natur.265..729N|last1=Nissani|first1=Moti|s2cid=4146956}}</ref> and that somatic components of the gonads originate in the mesodermal region of the fourth and fifth abdominal segment.<ref>{{Cite journal|last=Szabad|first=Janos, and Nothiger, Rolf|date=1992|title=Gynandromorphs of Drosophila suggest one common primordium for the somatic cells of the female and male gonads in the region of abdominal segments 4 and 5|url=http://dev.biologists.org/content/develop/115/2/527.full.pdf|journal=Development|volume=115|issue=2|pages=527–533|doi=10.1242/dev.115.2.527|pmid=1425338}}</ref>

==See also==
*[[Mosaic (genetics)|Mosaicism]] 
*[[Androgyny]]
*[[Chimerism]]
*[[Gynomorph]]
*[[Half-sider budgerigar]]
*[[Hermaphrodite]]

==References==
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==External links==
{{Commons category|Gynandromorphs}}
*"[https://www.livescience.com/14210-gynandromorphs-dual-sex-disorder-strange-birds-butterflies-gallery.html Stunning Dual-Sex Animals]" at [[Live Science]]
* Aayushi Pratap: [https://www.sciencenews.org/article/bird-male-female-grosbeak-gynandromorph This rare bird is male on one side and female on the other]; on: Sciencenews; October 6, 2020; about a gynandromorph [[rose-breasted grosbeak]].

[[Category:Insect physiology]]
[[Category:Sexual dimorphism]]