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Cucurbita
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==Taxonomy== ''Cucurbita'' was formally described in a way that meets the requirements of modern [[botanical nomenclature]] by [[Carl Linnaeus|Linnaeus]] in his ''[[Genera Plantarum]]'',<ref name="generaname">{{cite book |chapter=Cucurbita |title=Genera Plantarum |author-link=Carl Linnaeus |first=Carl |last=Linnaeus |year=1754 |page=441 |volume=1 |publisher=Impensis Laurentii Salvii via Biodiversity Heritage Library |location=Stockholm |chapter-url=https://www.biodiversitylibrary.org/item/14678#page/476/mode/1up |access-date=2018-02-20 |archive-date=2017-10-01 |archive-url=https://web.archive.org/web/20171001122929/http://www.biodiversitylibrary.org/item/14678#page/476/mode/1up |url-status=live }}</ref> the fifth edition of 1754 in conjunction with the 1753 first edition of ''[[Species Plantarum]]''.<ref name="typename">{{cite book |chapter=Cucurbita |title=Species Plantarum |last=Linnaeus |first=Carl |year=1753 |publisher=Impensis Laurentii Salvii via Biodiversity Heritage Library |location=Stockholm |volume=2 |page=1010 |chapter-url=https://www.biodiversitylibrary.org/page/359031#page/452/mode/1up |access-date=2018-02-20 |archive-date=2017-07-02 |archive-url=https://web.archive.org/web/20170702060402/http://www.biodiversitylibrary.org/page/359031#page/452/mode/1up |url-status=live }}</ref> ''Cucurbita pepo'' is the [[type species]] of the genus.<ref name="typename" /><ref name="typespeciesnhm">{{cite journal |title=Cucurbita |journal=The Linnaean Plant Name Typification Project |year=2022 |url=http://www.nhm.ac.uk/research-curation/research/projects/linnaean-typification/database/list.dsml?Genusqtype=starts+with&Genus=Cucurbita&Speciesqtype=starts+with&Species=pepo&Varqtype=starts+with&Var=&Ref=&CGenusqtype=starts+with&CGenus=&CSpeciesqtype=starts+with&CSpecies=&CSspqtype=starts+with&CSsp=&CVarqtype=starts+with&CVar=&Family=&sort=Genus%2CSpecies |publisher=Natural History Museum |doi=10.5519/qwv6u7j5 |access-date=November 4, 2013 |author1=Natural History Museum |archive-date=November 29, 2014 |archive-url=https://web.archive.org/web/20141129044110/http://www.nhm.ac.uk/research-curation/research/projects/linnaean-typification/database/list.dsml?Genusqtype=starts+with&Genus=Cucurbita&Speciesqtype=starts+with&Species=pepo&Varqtype=starts+with&Var=&Ref=&CGenusqtype=starts+with&CGenus=&CSpeciesqtype=starts+with&CSpecies=&CSspqtype=starts+with&CSsp=&CVarqtype=starts+with&CVar=&Family=&sort=Genus%2CSpecies |url-status=live }}</ref> Linnaeus initially included the species ''C. pepo'', ''C. verrucosa'' and ''C. melopepo'' (both now included in ''C. pepo''), as well as ''C. citrullus'' (watermelon, now ''[[Citrullus lanatus]]'') and ''C. lagenaria'' (now ''[[Lagenaria siceraria]]'') (both are not ''Cucurbita'' but are in the family Cucurbitaceae.<ref>{{cite web |url=http://www.theplantlist.org/tpl1.1/search?q=cucurbita |work=The Plant List |title=Cucurbita |access-date=1 January 2015 |archive-date=17 March 2019 |archive-url=https://web.archive.org/web/20190317222125/http://www.theplantlist.org/tpl1.1/search?q=Cucurbita |url-status=live }}</ref> The ''Cucurbita digitata'', ''C. foetidissima'', ''C. galeotti'', and ''C. pedatifolia'' [[species group]]s are [[xerophytes]], arid zone perennials with storage roots; the remainder, including the five domesticated species, are all mesophytic annuals or short-life perennials with no storage roots.<ref name="nee" /><ref name="whitaker">{{cite journal |author1=Whitaker, T.W. |author2=Bemis, W.P. |year=1975 |title=Origin and Evolution of the Cultivated Cucurbita |journal=Bulletin of the Torrey Botanical Club |volume=102 |issue=6 |pages=362β368 |jstor=2484762 |doi=10.2307/2484762}}</ref> The five domesticated species are mostly isolated from each other by [[Reproductive isolation|sterility barriers]] and have different physiological characteristics.<ref name="whitaker" /> Some [[Allogamy|cross pollinations]] can occur: ''C. pepo'' with ''C. argyrosperma'' and ''C. moschata''; and ''C. maxima'' with ''C. moschata''. Cross pollination does occur readily within the family Cucurbitaceae.<ref name="janssen">{{cite web |url=http://lancaster.unl.edu/hort/articles/2006/curbits.shtml |title=Curbit Family & Cross-Pollination |publisher=University of Nebraska β Lincoln |last=Janssen |first=Don |date=August 14, 2006 |access-date=January 14, 2015 |archive-date=February 13, 2015 |archive-url=https://web.archive.org/web/20150213065650/http://lancaster.unl.edu/hort/articles/2006/curbits.shtml |url-status=live }}</ref> The buffalo gourd (''C. foetidissima'') has been used as an intermediary, as it can be crossed with all the common ''Cucurbita''.<ref name="ucla" /> [[File:2006-10-18Cucurbita pepo06.jpg|thumb|alt=Several types and colors of ''Cucurbita''|An assortment of fruits of ''C. maxima'' and ''C. pepo'']] Various [[taxonomy (biology)|taxonomic]] treatments have been proposed for ''Cucurbita'', ranging from 13 to 30 species.<ref name="burrows">{{cite book|last1=Burrows|first1=George E.|last2=Tyrl|first2=Ronald J.|title=Toxic Plants of North America|publisher=Wiley-Blackwell|year=2013|location=Oxford|pages=389β391|isbn=978-0-8138-2034-7}}</ref> In 1990, ''Cucurbita'' expert Michael Nee classified them into the following oft-cited 13 species groups (27 species total), listed by group and alphabetically, with geographic origin:<ref name="nee" /><ref name="grin">{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/splist.pl?3164 |title=GRIN Species Records of Genus ''Cucurbita'' |author=GRIN |work=Taxonomy for Plants |publisher=[[United States Department of Agriculture|USDA]], [[Agricultural Research Service|ARS]], National Genetic Resources Program |location=National Germplasm Resources Laboratory, [[Beltsville, Maryland]] |access-date=September 1, 2013 |url-status=dead |archive-url=https://web.archive.org/web/20150924115025/http://www.ars-grin.gov/cgi-bin/npgs/html/splist.pl?3164 |archive-date=September 24, 2015 |author-link=Germplasm Resources Information Network }}</ref><ref name="itis">{{ITIS |id=22365 |taxon=Cucurbita |access-date=September 1, 2013}}</ref><ref>{{cite book|last=Goldman|first=Amy|title=The Compleat Squash: A Passionate Grower's Guide to Pumpkins, Squash, and Gourds|year=2004|publisher=Artisan|location=New York|page=77|isbn=978-1-57965-251-7}}</ref> * ''[[Cucurbita argyrosperma|C. argyrosperma]]'' ([[synonym (biology)|synonym]] ''C. mixta'') β cushaw pumpkin; origin: Mexico ** ''[[Cucurbita kellyana|C. kellyana]]'', origin: Pacific coast of western Mexico ** ''[[Cucurbita palmeri|C. palmeri]]'', origin: Pacific coast of northwestern Mexico ** ''[[Cucurbita sororia|C. sororia]]'', origin: Pacific coast Mexico to Nicaragua, northeastern Mexico * ''[[Cucurbita digitata|C. digitata]]'' β fingerleaf gourd; origin: southwestern United States (USA), northwestern Mexico ** ''[[Cucurbita californica|C. californica]]'' ** ''[[Cucurbita cordata|C. cordata]]'' ** ''[[Cucurbita cylindrata|C. cylindrata]]'' ** ''[[Cucurbita palmata|C. palmata]]'' * ''[[Cucurbita ecuadorensis|C. ecuadorensis]]'', origin: Ecuador's Pacific coast * ''[[Cucurbita ficifolia|C. ficifolia]]'' β figleaf gourd, chilacayote, alcayota; origin: Mexico, Panama, northern Chile and Argentina * ''[[Cucurbita foetidissima|C. foetidissima]]'' β stinking gourd, buffalo gourd; origin: Mexico ** ''[[Cucurbita scabridifolia|C. scabridifolia]]'', likely a [[natural hybrid]] of ''C. foetidissima'' and ''C. pedatifolia''<ref name="andres">{{cite journal |last=Andres |first=Thomas C. |year=1987 |title=Relationship of ''Cucurbita scabridifolia'' to ''C. foetidissima'' and ''C. pedatifolia'': A Case of Natural Interspecific Hybridization |url=http://cuke.hort.ncsu.edu/cgc/cgc10/cgc10-38.html |journal=Cucurbit Genetics Cooperative Report |volume=10 |pages=74β75 |access-date=2013-09-02 |archive-date=2013-09-21 |archive-url=https://web.archive.org/web/20130921053838/http://cuke.hort.ncsu.edu/cgc/cgc10/cgc10-38.html |url-status=live }}</ref><ref name="bailey">{{cite journal |last=Bailey |first=Liberty Hyde |author-link=Liberty Hyde Bailey |journal=Gentes Herbarum |volume=6 |pages=267β322 |year= 1943 |title=Species of ''Cucurbita''}}</ref> * ''[[Cucurbita galeottii|C. galeottii]]'', little known; origin: [[Oaxaca]], Mexico * ''[[Cucurbita lundelliana|C. lundelliana]]'', origin: Mexico, Guatemala, Belize * ''[[Cucurbita maxima|C. maxima]]'' β winter squash, pumpkin; origin: Argentina, Bolivia, Ecuador ** ''[[Cucurbita andreana|C. andreana]]'', origin β Argentina * ''[[Cucurbita moschata|C. moschata]]'' β butternut squash, 'Dickinson' pumpkin, golden cushaw; origin: Bolivia, Colombia, Ecuador, Mexico, Panama, Puerto Rico, Venezuela * ''[[Cucurbita okeechobeensis|C. okeechobeensis]]'', origin: Florida ** ''[[Cucurbita martinezii|C. martinezii]]'', origin: Mexican Gulf Coast and foothills * ''[[Cucurbita pedatifolia|C. pedatifolia]]'', origin: [[QuerΓ©taro]], Mexico ** ''[[Cucurbita moorei|C. moorei]]'' * ''[[Cucurbita pepo|C. pepo]]'' β field pumpkin, summer squash, zucchini, vegetable marrow, courgette, acorn squash; origin: Mexico, US ** ''[[Cucurbita fraterna|C. fraterna]]'', origin: [[Tamaulipas]] and [[Nuevo LeΓ³n]], Mexico ** ''[[Cucurbita texana|C. texana]]'', origin: [[Texas]], US * ''[[Cucurbita radicans|C. radicans]]'' β calabacilla, calabaza de coyote; origin: Central Mexico ** ''[[Cucurbita gracilior|C. gracilior]]'' The taxonomy by Nee closely matches the species groupings reported in a pair of studies by a botanical team led by Rhodes and Bemis in 1968 and 1970 based on statistical groupings of several [[phenotype|phenotypic]] traits of 21 species. Seeds for studying additional species members were not available. Sixteen of the 21 species were grouped into five clusters with the remaining five being classified separately:<ref name="rhodes" /><ref name="bemis1970">{{cite journal | last1 = Bemis | first1 = W. P. | last2 = Rhodes | first2 = A. M. | last3 = Whitaker | first3 = Thomas W. | last4 = Carmer | first4 = S. G. | year = 1970 | title = Numerical Taxonomy Applied to ''Cucurbita'' Relationships | journal = [[American Journal of Botany]] | volume = 57 | issue = 4 | pages = 404β412 | jstor = 2440868 | doi = 10.2307/2440868}}</ref> * ''C. digitata'', ''C. palmata'', ''C. californica'', ''C. cylindrata'', ''C. cordata'' * ''C. martinezii'', ''C. okeechobeensis'', ''C. lundelliana'' * ''C. sororia'', ''C. gracilior'', ''C. palmeri''; ''C. argyrosperma'' (reported as ''C. mixta'') was considered close to the three previous species * ''C. maxima'', ''C. andreana'' * ''C. pepo'', ''C. texana'' * ''C. moschata'', ''C. ficifolia'', ''C. pedatifolia'', ''C. foetidissima'', and ''C. ecuadorensis'' were placed in their own separate species groups as they were not considered significantly close to any of the other species studied. ===Phylogeny=== The full [[Phylogenetics|phylogeny]] of this genus is unknown, and research was ongoing in 2014.<ref name="gongpkl2">{{cite journal |last1=Gon g|first1=L. |last2=Pachner |first2=M. |last3=Kalai |first3=K. |last4=Lelley |first4=T. |journal=Genome |title=SSR-based Genetic Linkage Map of ''Cucurbita moschata'' and its Synteny With ''Cucurbita pepo'' |date=November 2008 |volume=51 |issue=11 |pages=878β887 |pmid=18956020 |doi=10.1139/G08-072}}</ref><ref name="gongskpl1">{{cite journal |last1=Gong |first1=L. |last2=Stift |first2= G. |last3=Koffler |first3=R. |last4=Pachner |first4=M. |last5=Lelley |first5=T. |journal=Theoretical and Applied Genetics |title=Microsatellites for the Genus ''Cucurbita'' and an SSR-based Genetic Linkage Map of ''Cucurbita pepo'' L. |date=June 2008 |volume=117 |issue=1 |pages=37β48 |pmid=18379753 |doi=10.1007/s00122-008-0750-2 |pmc=2413107}}</ref> The following [[cladogram]] of ''Cucurbita'' phylogeny is based upon a 2002 study of [[mitochondrial DNA]] by Sanjur and colleagues.<ref name="sanjur" /> {{clade| style=font-size:85%;line-height:85%; |1={{clade |1= ''[[Chayote|Sechium edule]]'' |2={{clade |1=''[[Cucurbita ficifolia|C. ficifolia]]'' |2={{clade |1=''[[Cucurbita foetidissima|C. foetidissima]]'' |2={{clade |1={{clade |1= ''[[Cucurbita maxima|C. maxima]]'' and ''[[Cucurbita andreana|C. andreana]]'' |2= ''[[Cucurbita ecuadorensis|C. ecuadorensis]]'' }} |2={{clade |1 = ''[[Cucurbita martinezii|C. martinezii]]'' |2= {{clade |1=''[[Cucurbita pepo|C. pepo]]'' subspp. ''fraterna'' and ''ovifera'' |2= {{clade |1 = ''[[Cucurbita pepo|C. pepo]]'' subsp. ''pepo'' |2= {{clade |1 = ''[[Cucurbita sororia|C. sororia]]'', in part |2= {{clade |1 = ''[[Cucurbita moschata|C. moschata]]'' |2 = ''[[Cucurbita sororia|C. sororia]]'', in part and ''[[Cucurbita argyrosperma|C. argyrosperma]]'' }} }} }} }} }} }} }} }} }} }}
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