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Cucurbita
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{{short description|A genus of herbaceous vines in the gourd family, Cucurbitaceae, also known as cucurbits, native to the Andes and Mesoamerica}}{{featured article}}{{italic title}}{{automatic taxobox|name = Squash|image = Squashes at Kew Gardens IncrEdibles 2013.jpg|image_caption = Cucurbita fruits come in an assortment of colors and sizes.|image_alt = Various sizes, shapes, and colors of Cucurbita|taxon = Cucurbita
Carl Linnaeus>L.Mellonia Guglielmo Gasparrini>Gasp. CUCURBITA L. >PUBLISHER=TROPICOS, MISSOURI BOTANICAL GARDEN, January 12, 2014,
}Cucurbita (Latin for gourd)BOOK, Gledhill, David, 2008, The Names of Plants, Cambridge University Press, Cambridge, 9780521866453, 127, is a genus of herbaceous vines in the gourd family, Cucurbitaceae, also known as cucurbits, native to the Andes and Mesoamerica. Five species are grown worldwide for their edible fruit, variously known as squash, pumpkin, or gourd depending on species, variety, and local parlance,{{efn|Due to wide variation in how the terms squash, pumpkin, and gourd are used, even among academics, in this article, the term squash can refer to any member of the genus Cucurbita. Pumpkin and gourd are used to refer to species, varieties, and cultivars commonly referred to by those terms.BOOK, Ferriol, María, Picó, Belén, 2007, Handbook of Plant Breeding: Vegetables I, Springer, New York, 3, 978-0-387-72291-7, 317, The common terms "pumpkin", "squash", "gourd", "cushaw", "ayote", "zapallo", "calabaza", etc. are often applied indiscriminately to different cultivated species of the New World genus Cucurbita L. (Cucurbitaceae): C. pepo L., C.  maxima Duchesne, C. moschata Duchesne, C. argyrosperma C. Huber and C. ficifolia Bouché., }} and for their seeds. Other kinds of gourd, also called bottle-gourds, are native to Africa and belong to the genus Lagenaria, which is in the same family and subfamily as Cucurbita but in a different tribe. These other gourds are used as utensils or vessels, and their young fruits are eaten much like those of Cucurbita species.Most Cucurbita species are herbaceous vines that grow several meters in length and have tendrils, but non-vining "bush" cultivars of C. pepo and C. maxima have also been developed. The yellow or orange flowers on a Cucurbita plant are of two types: female and male. The female flowers produce the fruit and the male flowers produce pollen. Many North and Central American species are visited by specialist bee pollinators, but other insects with more general feeding habits, such as honey bees, also visit.There is debate about the taxonomy of the genus, as the number of accepted species varies from 13 to 30. The five domesticated species are Cucurbita argyrosperma, C. ficifolia, C. maxima, C. moschata, and C. pepo. All of these can be treated as winter squash because the full-grown fruits can be stored for months; however, C. pepo includes some cultivars that are better used only as summer squash.The fruits of the genus Cucurbita are good sources of nutrients, such as vitamin A and vitamin C, among other nutrients according to species. The fruits have many culinary uses including pumpkin pie, biscuits, bread, desserts, puddings, beverages, and soups.

Description

(File:Cucurbita 2011 G1.jpg|thumb|C. pepo pumpkins â€“ the two bright orange ones in center right, and squashes C. maxima, all others)Cucurbita species fall into two main groups. The first group are annual or short-lived perennial vines and are mesophytic, i.e. they require a more or less continuous water supply. The second group are perennials growing in arid zones and so are xerophytic, tolerating dry conditions. Cultivated Cucurbita species were derived from the first group. Growing {{convert|5|to|15|m|ft|sp=us}} in height or length, the plant stem produces tendrils to help it climb adjacent plants and structures or extend along the ground. Most species do not readily root from the nodes; a notable exception is C. ficifolia, and the four other cultivated mesophytes do this to a lesser extent. The vine of the perennial Cucurbita can become semiwoody if left to grow. There is wide variation in size, shape, and color among Cucurbita fruits, and even within a single species. C. ficifolia is an exception, being highly uniform in appearance.JOURNAL, Nee, Michael, 1990, The Domestication of Cucurbita (Cucurbitaceae), Economic Botany, 44, 3, Supplement: New Perspectives on the Origin and Evolution of New World Domesticated Plants, 56–68, New York Botanical Gardens Press, New York, 4255271, 10.1007/BF02860475, The morphological variation in the species C. pepoJOURNAL, Decker-Walters, Deena S., Staub, Jack E., Chung, Sang-Min, Nakata, Eijiro, Quemada, Hector D., 2002, Diversity in Free-Living Populations of Cucurbita pepo (Cucurbitaceae) as Assessed by Random Amplified Polymorphic DNA, Systematic Botany, 27, 1, 19–28, American Society of Plant Taxonomists, 3093892, and C. maximaJOURNAL, Millán, R., Variaciones del Zapallito Amargo Cucurbita andreana y el Origen de Cucurbita maxima, Revista Argentina de Agronomía, 1945, 12, 86–93, Spanish, is so vast that its various subspecies and cultivars have been misidentified as totally separate species.File:Cucurbita moschata leaves.jpg|thumb|left|alt=Green Cucurbita moschata leaves with white spots|The leaves of Cucurbita moschataCucurbita moschataThe typical cultivated Cucurbita species has five-lobed or palmately divided leaves with long petioles, with the leaves alternately arranged on the stem. The stems in some species are angular. All of the above-ground parts may be hairy with various types of trichomes, which are often hardened and sharp. Spring-like tendrils grow from each node and are branching in some species. C. argyrosperma has ovate-cordate (egg-shaped to heart-shaped) leaves. The shape of C. pepo leaves varies widely. C. moschata plants can have light or dense pubescence. C. ficifolia leaves are slightly angular and have light pubescence. The leaves of all four of these species may or may not have white spots.There are male (staminate) and female (pistillate) flowers (unisexual flowers) on a single plant (monoecious), and these grow singly, appearing from the leaf axils. Flowers have five fused yellow to orange petals (the corolla) and a green bell-shaped calyx. Male flowers in Cucurbitaceae generally have five stamens, but in Cucurbita there are only three, and their anthers are joined together so that there appears to be one.BOOK, Mabberley, D. J., 2008, The Plant Book: A Portable Dictionary of the Vascular Plants, Cambridge University Press, Cambridge, 978-0-521-82071-4, 235, WEB, Lu, Anmin, Jeffrey, Charles, Flora of China, Cucurbita Linnaeus,weblink February 21, 2015, Female flowers have thick pedicels, and an inferior ovary with 3–5 stigmas that each have two lobes. The female flowers of C. argyrosperma and C. ficifolia have larger corollas than the male flowers. Female flowers of C. pepo have a small calyx, but the calyx of C. moschata male flowers is comparatively short.File:2016-11-10 Cucurbits (Cucurbitaceae) Madrid Botanico.jpg|thumb|alt=a variety of fruits displayed in a garden, some in a sack, some in a basket|A variety of fruits displayed at the Real Jardín Botánico de MadridReal Jardín Botánico de MadridCucurbita fruits are large and fleshy. Botanists classify the Cucurbita fruit as a pepo, which is a special type of berry derived from an inferior ovary, with a thick outer wall or rind with hypanthium tissue forming an exocarp around the ovary, and a fleshy interior composed of mesocarp and endocarp. The term "pepo" is used primarily for Cucurbitaceae fruits, where this fruit type is common, but the fruits of Passiflora and Carica are sometimes also pepos.WEB,weblink A Systematic Treatment of Fruit Types, Worldbotanical, October 10, 2013, BOOK, Schrager, Victor, The Compleat Squash: A Passionate Grower's Guide to Pumpkins, Squash, and Gourds, Artisan, New York, 2004, 25, 978-1-57965-251-7, The seeds, which are attached to the ovary wall (parietal placentation) and not to the center, are large and fairly flat with a large embryo that consists almost entirely of two cotyledons. Fruit size varies considerably: wild fruit specimens can be as small as {{convert|4|cm|in|sp=us}} and some domesticated specimens can weigh well over {{convert|300|kg|lb|sp=us}}. The current world record was set in 2014 by Beni Meier of Switzerland with a {{convert|2323.7|lb|kg|sp=us|adj=on}} pumpkin.NEWS, 2014 - Beni Meier and his 2323.7 pound World Record Giant Pumpkin!,weblink BigPumpkins.com, February 12, 2016, 2014,

Taxonomy

Cucurbita was formally described in a way that meets the requirements of modern botanical nomenclature by Linnaeus in his Genera Plantarum,BOOK, Cucurbita, Genera Plantarum, Carl Linnaeus, Carl, Linnaeus, 1754, 441, 1, Impensis Laurentii Salvii via Biodiversity Heritage Library, Stockholm,weblink the fifth edition of 1754 in conjunction with the 1753 first edition of Species Plantarum.BOOK, Cucurbita, Species Plantarum, Linnaeus, Carl, 1753, Impensis Laurentii Salvii via Biodiversity Heritage Library, Stockholm, 2, 1010,weblink Cucurbita pepo is the type species of the genus.WEB, Cucurbita, The Linnaean Plant Name Typification Project,weblink Natural History Museum, November 4, 2013, 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.WEB,weblink The Plant List, Cucurbita, 1 January 2015, The Cucurbita digitata, C. foetidissima, C. galeotti, and C. pedatifolia species groups 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.JOURNAL, Whitaker, T.W., Bemis, W.P., 1975, Origin and Evolution of the Cultivated Cucurbita, Bulletin of the Torrey Botanical Club, 102, 6, 362–368, 2484762, 10.2307/2484762, The five domesticated species are mostly isolated from each other by sterility barriers and have different physiological characteristics. Some 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.WEB,weblink Curbit Family & Cross-Pollination, University of Nebraska - Lincoln, Janssen, Don, August 14, 2006, January 14, 2015, JOURNAL, World Applied Sciences Journal, 20, 10, 2012, 1366–1370, Production of Cucurbita Interspecific Hybrids ThroughCross Pollination and Embryo Rescue Technique, Rakha, M. T., Metwally, E. I., Moustafa, E. A., Etman, A. A., Dewir, Y. H.,weblink The buffalo gourd (C. foetidissima), which; according to some, does not taste good, has been used as an intermediary as it can be crossed with all the common Cucurbita.(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 taxonomic treatments have been proposed for Cucurbita, ranging from 13–30 species.BOOK, Burrows, George E., Tyrl, Ronald J., Toxic Plants of North America, Wiley-Blackwell, 2013, Oxford, 389–391, 978-0-8138-2034-7, 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:WEB,weblink GRIN Species Records of Genus Cucurbita, GRIN, Taxonomy for Plants, United States Department of Agriculture, USDA, Agricultural Research Service, ARS, National Genetic Resources Program, National Germplasm Resources Laboratory, Beltsville, Maryland, September 1, 2013, yes,weblink" title="web.archive.org/web/20150924115025weblink">weblink September 24, 2015, Germplasm Resources Information Network, {{ITIS |id=22365 |taxon=Cucurbita |accessdate=September 1, 2013}}BOOK, Goldman, Amy, The Compleat Squash: A Passionate Grower's Guide to Pumpkins, Squash, and Gourds, 2004, Artisan, New York, 77, 978-1-57965-251-7, 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 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:JOURNAL, Bemis, W. P., Rhodes, A. M., Whitaker, Thomas W., Carmer, S. G., 1970, Numerical Taxonomy Applied to Cucurbita Relationships, American Journal of Botany, 57, 4, 404–412, 2440868, 10.2307/2440868,
  • 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 phylogeny of this genus is unknown, and research was ongoing in 2014.JOURNAL, Gon g, L., Pachner, M., Kalai, K., Lelley, T., Genome, SSR-based Genetic Linkage Map of Cucurbita moschata and its Synteny With Cucurbita pepo, November 2008, 51, 11, 878–887, 18956020, 10.1139/G08-072, JOURNAL, Gong, L., Stift, G., Koffler, R., Pachner, M., Lelley, T., Theoretical and Applied Genetics, Microsatellites for the Genus Cucurbita and an SSR-based Genetic Linkage Map of Cucurbita pepo L., June 2008, 117, 1, 37–48, 18379753, 10.1007/s00122-008-0750-2, 2413107, The following cladogram of Cucurbita phylogeny is based upon a 2002 study of mitochondrial DNA by Sanjur and colleagues.{{clade| style=font-size:85%;line-height:85%;
|1={{clade
|1= Sechium edule
|2={{clade
|1=C. ficifolia
|2={{clade
|1=C. foetidissima
|2={{clade
|1={{clade
|1= C. maxima and C. andreana
|2= C. ecuadorensis
}}
|2={{clade
|1 = C. martinezii
|2= {{clade
|1=C. pepo subspp. fraterna and ovifera
|2= {{clade
|1 = C. pepo subsp. pepo
|2= {{clade
|1 = C. sororia, in part
|2= {{clade
|1 = C. moschata
|2 = C. sororia, in part and C. argyrosperma
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}

Reproductive biology

File:Peponapis pruinosaCane-12.JPG|thumb|alt=Bee pollinating female Cucurbita flower|Cucurbita female flower with pollinating squash beesquash beeAll species of Cucurbita have 20 pairs of chromosomes.JOURNAL, Rhodes, A. M., Bemis, W. P., Whitaker, Thomas W., Carmer, S. G., 1968, A Numerical Taxonomic Study of Cucurbita, Brittonia, 20, 3, 251–266, New York Botanical Garden Press, 2805450, 10.2307/2805450, Many North and Central American species are visited by specialist pollinators in the apid tribe Eucerini, especially the genera Peponapis and Xenoglossa, and these squash bees can be crucial to the flowers producing fruit after pollination.JOURNAL, Hurd, Paul D., Linsley, E. Gorton, 1971, Squash and Gourd Bees (Peponapis, Xenoglossa) and the Origin of the Cultivated Cucurbita, Evolution (journal), Evolution, 25, 1, 218–234, Society for the Study of Evolution, St. Louis, MO, 2406514, 10.2307/2406514, JOURNAL, Whitaker, Thomas W., Bemis, W. P., Evolution in the Genus Cucurbita, Evolution, 18, 4, 553–559, 1964, 2406209, 10.2307/2406209, (File:Cucurbita maxima Zapallo Plomo semillería Costanzi - flowers detail (masculine) - male flower, some petals and 1 filament removed.jpg|thumb|120px|left|alt=Male Cucurbita flower|Male flower, part of the perianth and one filament removed)When there is more pollen applied to the stigma, more seeds are produced in the fruits and the fruits are larger with greater likelihood of maturation, an effect called xenia. Competitively grown specimens are therefore often hand-pollinated to maximize the number of seeds in the fruit, which increases the fruit size; this pollination requires skilled technique.JOURNAL, Journal of New Seeds, 1, 3–4, 2000, 1–47, Rationale and Methods for Producing Hybrid Cucurbit Seed, Robinson, Richard W., 10.1300/J153v01n03_01, JOURNAL, Stephenson, Andrew G., Devlin, B., Horton, J. Brian, The Effects of Seed Number and Prior Fruit Dominance on the Pattern of Fruit Production in Cucurbita pepo (Zucchini Squash),weblink Annals of Botany, 1988, 62, 6, 653–661, 10.1093/oxfordjournals.aob.a087705, Seedlessness is known to occur in certain cultivars of C. pepo.JOURNAL, Robinson, R. W., Reiners, Stephen, Parthenocarpy in Summer Squash,weblink HortScience, July 1999, 34, 4, 715–717, 10.21273/HORTSCI.34.4.715, JOURNAL, Menezes, C. B., Maluf, W. R., Azevedo, S. M., Faria, M. V., Nascimento, I. R., Gomez, L. A., Bearzoti, E., Inheritance of Parthenocarpy in Summer Squash (Cucurbita pepo L.)., Genetics and Molecular Research, March 2005, 4, 1, 39–46, 15841434, The most critical factors in flowering and fruit set are physiological, having to do with the age of the plant and whether it already has developing fruit.JOURNAL, Stapleton, Suzanne Cady, Wien, H. Chris, Morse, Roger A., 2000, Flowering and Fruit Set of Pumpkin Cultivars under Field Conditions, HortScience, 35, 6, 1074–1077,weblink 10.21273/HORTSCI.35.6.1074, The plant hormones ethylene and auxin are key in fruit set and development.JOURNAL, Martínez, Cecelia, Manzano, Susana, Megías, Zoraida, Garrido, Dolores, Picó, Belén, Jamilena, Manuel, 2013, Involvement of Ethylene Biosynthesis and Signalling in Fruit Set and Early Fruit Development in Zucchini Squash (Cucurbita pepo L.), BMC Plant Biology, 13, 139, 139, 10.1186/1471-2229-13-139, 24053311, Ethylene promotes the production of female flowers. When a plant already has a fruit developing, subsequent female flowers on the plant are less likely to mature, a phenomenon called "first-fruit dominance", and male flowers are more frequent, an effect that appears due to reduced natural ethylene production within the plant stem.JOURNAL, Krupnick, Gary A., Brown, Kathleen M., Stephenson, Andrew G., 1999, The Influence of Fruit on the Regulation of Internal Ethylene Concentrations and Sex Expression in Cucurbita texana, International Journal of Plant Sciences, 160, 2, 321–330, 10.1086/314120, Ethephon, a plant growth regulator product that is converted to ethylene after metabolism by the plant, can be used to increase fruit and seed production.JOURNAL, Murray, M., 1987, Field Applications Of Ethephon For Hybrid And Open-Pollinated Squash (Cucurbita Spp) Seed Production, Acta Horticulturae, 201, 201, 149–156, 10.17660/ActaHortic.1987.201.15, The plant hormone gibberellin, produced in the stamens, is essential for the development of all parts of the male flowers. The development of female flowers is not yet understood.JOURNAL, Pimenta Lange, Maria João, Knop, Nicole, Lange, Theo, 2012, Stamen-derived Bioactive Gibberellin is Essential for Male Flower Development of Cucurbita maxima L., Journal of Experimental Botany, 63, 7, 2681–2691, 10.1093/jxb/err448, 3346225, 22268154, Gibberellin is also involved in other developmental processes of plants such as seed and stem growth.WEB, Plant Hormones, Charles Sturt University,weblink January 15, 2014, yes,weblink" title="web.archive.org/web/20140116140049weblink">weblink January 16, 2014, {{Clear}}

Germination and seedling growth

File:Kabocha (GH) 21June2005 sown 14June.JPG|thumb|alt=Kabocha seedling at seven days age|KabochaKabochaSeeds with maximum germination potential develop (in C. moschata) by 45 days after anthesis, and seed weight reaches its maximum 70 days after anthesis.JOURNAL, Wilson, Mack A., Splittstoesser, Walter E., 1980, The Relationship Between Embryo Axis Weight and Reserve Protein During Development and Pumpkin Seed Germination, Journal of Seed Technology, 5, 2, 35–41, 23432821, Some varieties of C. pepo germinate best with eight hours of sunlight daily and a planting depth of {{convert|1.2|cm|in|sp=us}}. Seeds planted deeper than {{convert|12.5|cm|in|sp=us}} are not likely to germinate.JOURNAL, Oliver, Lawrence R., Harrison, Steve A., McClelland, Marilyn, 1983, Germination of Texas Gourd (Cucurbita texana) and Its Control in Soybeans (Glycine max), Weed Science, 31, 5, 700–706, 4043694, 10.1017/S0043174500070211, In C. foetidissima, a weedy species, plants younger than 19 days old are not able to sprout from the roots after removing the shoots. In a seed batch with 90 percent germination rate, over 90 percent of the plants had sprouted after 29 days from planting.JOURNAL, Horak, Michael J., Sweat, Jonathan K., 1994, Germination, Emergence, and Seedling Establishment of Buffalo Gourd (Cucurbita foetidissima), Weed Science, 42, 3, 358–363, 4045510, 10.1017/S0043174500076621, Experiments have shown that when more pollen is applied to the stigma, as well as the fruit containing more seeds and being larger (the xenia effect mentioned above), the germination of the seeds is also faster and more likely, and the seedlings are larger.JOURNAL, Winsor, J. A., Davis, L. E., Stephenson, A. G., 1987, The Relationship Between Pollen Load and Fruit Maturation and the Effect of Pollen Load on Offspring Vigor in Cucurbita pepo, The American Naturalist, 129, 5, 643–656, 2461727, 10.1086/284664, Various combinations of mineral nutrients and light have a significant effect during the various stages of plant growth. These effects vary significantly between the different species of Cucurbita. A type of stored phosphorus called phytate forms in seed tissues as spherical crystalline intrusions in protein bodies called globoids. Along with other nutrients, phytate is used completely during seedling growth.JOURNAL, Beecroft, Penny, Lott, John N. A., Changes in the Element Composition of Globoids From Cucurbita maxima and Cucurbita andreana Cotyledons During Early Seedling Growth, Canadian Journal of Botany, 74, 6, 1996, 838–847, 10.1139/b96-104, Heavy metal contamination, including cadmium, has a significant negative impact on plant growth.JOURNAL, Subin, M. P., Francis, Steffy, Phytotoxic Effects of Cadmium on Seed Germination, Early Seedling Growth and Antioxidant Enzyme Activities in Cucurbita maxima Duchesne, International Research Journal of Biological Sciences, 2, 9, 2013, 40–47, 10.1139/b96-104, Cucurbita plants grown in the spring tend to grow larger than those grown in the autumn.JOURNAL, Fenner, G. P., Patteron, G. W., Lusby, W. R., Developmental Regulation of Sterol Biosynthesis in Cucurbita maxima L., Lipids, 24, 4, 1989, 271–277, 10.1007/BF02535162,

Distribution and habitat

(File:2009 Circleville Pumpkin Show champions.jpg|thumb|left|alt=Very large orange pumpkins| A festival-winning pumpkin in 2009 weighing {{convert|742|kg|lb|sp=us}})Archaeological investigations have found evidence of domestication of Cucurbita going back over 8,000 years from the very southern parts of Canada down to Argentina and Chile. Centers of domestication stretch from the Mississippi River watershed and Texas down through Mexico and Central America to northern and western South America. Of the 27 species that Nee delineates, five are domesticated. Four of them, C. argyrosperma, C. ficifolia, C. moschata, and C. pepo, originated and were domesticated in Mesoamerica; for the fifth, C. maxima, these events occurred in South America.WEB, Saade, R. Lira, Hernández, S. Montes, Cucurbits,weblink Purdue Horticulture, September 2, 2013, Within C. pepo, the pumpkins, the scallops, and possibly the crooknecks are ancient and were domesticated at different times and places. The domesticated forms of C. pepo have larger fruits than non-domesticated forms and seeds that are bigger but fewer in number. In a 1989 study on the origins and development of C. pepo, botanist Harry Paris suggested that the original wild specimen had a small round fruit and that the modern pumpkin is its direct descendant. He suggested that the crookneck, ornamental gourd, and scallop are early variants and that the acorn is a cross between the scallop and the pumpkin.(File:Cucurbita moschata Butternut 2012 G2.jpg|thumb|alt=Sliced butternut squash|C. moschata 'Butternut')C. argyrosperma is not as widespread as the other species. The wild form C. a. subsp. sororia is found from Mexico to Nicaragua, and cultivated forms are used in a somewhat wider area stretching from Panama to the southeastern United States. It was probably bred for its seeds, which are large and high in oil and protein, but its flesh is of poorer quality than that of C. moschata and C. pepo. It is grown in a wide altitudinal range: from sea level to as high as {{convert|1800|m|ft|sp=us}} in dry areas, usually with the use of irrigation, or in areas with a defined rainy season, where seeds are sown in May and June.C. ficifolia and C. moschata were originally thought to be Asiatic in origin, but this has been disproven. The origin of C. ficifolia is Latin America, most likely southern Mexico, Central America, or the Andes. It grows at altitudes ranging from {{convert|1000|m|ft|sp=us}} to {{convert|3000|m|ft|sp=us}} in areas with heavy rainfall. It does not hybridize well with the other cultivated species as it has significantly different enzymes and chromosomes.C. maxima originated in South America over 4,000 years ago, probably in Argentina and Uruguay. The plants are sensitive to frost, and they prefer both bright sunlight and soil with a pH of 6.0 to 7.0.WEB, Cucurbita maxima Origin/ Habitat,weblink University of Wisconsin, 2007, September 2, 2013, C. maxima did not start to spread into North America until after the arrival of Columbus. Varieties were in use by native peoples of the United States by the 16th century. Types of C. maxima include triloba,WEB, Holotype of Cucurbita maxima Duchesne var. triloba Millán [family CUCURBITACEAE],weblink October 3, 2013, {{subscription required}} zapallito,JOURNAL, López-Anido, F., Cravero, V., Asprelli, P., Cointry, E., Firpo, I., García, S. M., Inheritance of Immature Fruit Color in Cucurbita maxima var. zapallito (Carrière) Millán,weblink Cucurbit Genetics Cooperative Report, 26, 48–50, 2003, zipinka,WEB, Holotype of Cucurbita maxima Duchesne var. zipinka Millán [family CUCURBITACEAE],weblink October 3, 2013, {{subscription required}} Banana, Delicious, Hubbard, Marrow (C. maxima Marrow), Show, and Turban.BOOK, Robinson, Richard Warren, Decker-Walters, D. S., Cucurbits, 1997, CAB International, Oxfordshire, UK, 71–83, 978-0-85199-133-7, File:Cucurbita moschata 'Tromboncino'.jpg|thumb|left|alt=Curved green squashes|Fruit of the 'Tromboncino' cultivar of the Crookneck (C. moschata) Group are eaten either when very young, or as mature winter squash.]]C. moschata is native to Latin America, but the precise location of origin is uncertain.JOURNAL, Wessel-Beaver, Linda, 2000, Evidence for the Center of Diversity of Cucurbita moschata in Colombia,weblink Cucurbit Genetics Cooperative Report, 23, 54–55, It has been present in Mexico, Belize, Guatemala, and Peru for 4,000–6,000 years and has spread to Bolivia, Ecuador, Panama, Puerto Rico, and Venezuela. This species is closely related to C. argyrosperma. A variety known as the Seminole Pumpkin has been cultivated in Florida since before the arrival of Columbus. Its leaves are {{convert|20|to|30|cm|in|sp=us|0}} wide. It generally grows at low altitudes in hot climates with heavy rainfall, but some varieties have been found above {{convert|2200|m|ft|sp=us}}. Groups of C. moschata include Cheese, Crookneck (C. moschata), and Bell.C. pepo is one of the oldest, if not the oldest, domesticated species with the oldest known locations being Oaxaca, Mexico, 8,000–10,000 years ago, and Ocampo, Tamaulipas, Mexico, about 7,000 years ago. It is known to have appeared in Missouri, United States, at least 4,000 years ago.WEB, Wilson, Hugh D., Texas A&M Bioinformatics Working Group,weblink Free-living Cucurbita pepo in the United States Viral Resistance, Gene Flow, and Risk Assessment, What is Cucurbita texana?, September 8, 2013, Debates about the origin of C. pepo have been on-going since at least 1857.JOURNAL, Kirkpatrick, Kurt J., Wilson, Hugh D., 1988, Interspecific Gene Flow in Cucurbita: C. texana vs. C. pepo, American Journal of Botany, 75, 4, 519–527, Botanical Society of America, 10.2307/2444217, 2444217, There have traditionally been two opposing theories about its origin: 1) that it is a direct descendant of C. texana and 2) that C. texana is merely feral C. pepo. A more recent theory by botanist Thomas Andres in 1987 is that descendants of C. fraterna hybridized with C. texana, resulting in two distinct domestication events in two different areas: one in Mexico and one in the eastern United States, with C. fraterna and C. texana, respectively, as the ancestral species.BOOK, Soltis, Douglas E., Soltis, Pamela S., Pamela S. Soltis, Isozymes in Plant Biology, 176, London, Dioscorodes Press, 1990, 978-0-412-36500-3, C. pepo may have appeared in the Old World before moving from Mexico into South America. It is found from sea level to slightly above {{convert|2000|m|ft|sp=us}}. Leaves have 3–5 lobes and are {{convert|20–35|cm|in|sp=us|0}} wide. All the subspecies, varieties, and cultivars are interfertile. In 1986 Paris proposed a revised taxonomy of the edible cultivated C. pepo based primarily on the shape of the fruit, with eight groups . All but a few C. pepo cultivars can be included in these groups.JOURNAL, Paris, Harry S., Phytologia, A Proposed Subspecific Classification for Cucurbita pepo,weblink 1986, 61, 3, 133–138, WEB,weblink Missouri Botanical Garden, Cucurbita pepo, September 2, 2013, BOOK, Heistinger, Andrea, The Manual of Seed Saving: Harvesting, Storing, and Sowing Techniques for Vegetables, Herbs, and Fruits, Timber Press, Portland, OR, 2013, 278, 978-1-60469-382-9, There is one non-edible cultivated variety: C. pepo var. ovifera.JOURNAL, Decker, Deena S., Wilson, Hugh D., 1987, Allozyme Variation in the Cucurbita pepo Complex: C. pepo var. ovifera vs. C. texana, Systematic Botany, 12, 2, 263–273, American Society of Plant Taxonomists, 2419320, 10.2307/2419320, {{br}}{| class="wikitable sortable"|+A classification of cultivated C. pepo varieties based on Paris' eight groups and the one non-edible variety! Cultivar group !! Botanical name !! Image !! DescriptionC. pepo
var. turbinata >100px| Winter squash, both a shrubby and creeping plant, obovoid or conical shape, pointed at the apex and with longitudinal grooves, thus resembling a spinning top, ex: Acorn squashC. pepo var. Ionga >100px| Summer squash, long round slender fruit that is slightly bulbous at the apex, similar to fastigata, ex: Cocozelle von tripolisC. pepo var. torticollia (also torticollis) >100px| Summer squash, shrubby plant, with yellow, golden, or white fruit which is long and curved at the end and generally has a verrucose (wart-covered) rind, ex: Crookneck squashC. pepo var. pepo >75pxPumpkin; includes C. pepo subsp. pepo var. styriaca, used for Styrian pumpkin seed oilFüRNKRANZ LAST2 = LUKESCH LAST3 = MüLLER LAST4 = HUSS LAST5 = GRUBE LAST6 = BERG YEAR = 2012 JOURNAL = MICROBIAL ECOLOGY (JOURNAL) > VOLUME = 63 PAGES = 418–428 JSTOR = 41412429 PMID = 21947430, C. pepo var. clypeata; called C. melopepo by Carl Linnaeus>Linnaeus (File:Pattypan squash J1.jpgalt=Whitish round squash) Summer squash, prefers half-shrubby habitat, flattened or slightly discoidal shape, with undulations or equatorial edges, ex: Pattypan squashC. pepo var. recticollis >100px| Summer squash, shrubby plant, with yellow or golden fruit and verrucose rind, similar to var. torticollia but a stem end that narrows, ex: Straightneck squashC. pepo var. fastigata >100px| Summer and winter squashes, creeper traits and a semi-shrub, cream to dark green color, short round fruit with a slightly broad apex, ex: Spaghetti squash (a winter variety)C. pepo var. cylindrica >100px| Summer squash, presently the most common group of cultivars, origin is recent (19th century), semi-shrubby, cylindrical fruit with a mostly consistent diameter, similar to fastigata, ex: ZucchiniC. pepo var. ovifera >100px| Non-edible, field squash closely related to C. texana, vine habitat, thin stems, small leaves, three sub-groups: C. pepo var. ovifera (egg-shaped, pear-shaped), C. pepo var. aurantia (orange color), and C. pepo var. verrucosa (round warty gourds), ornamental gourds found in Texas and called var. texana and ornamental gourds found outside of Texas (Illinois, Missouri, Arkansas, Oklahoma, and Louisiana) are called var. ozarkana.

History and domestication

File:Bnf Anne f161.jpg|thumb|upright|alt=Early 1500s painting of squash plants and fruits|Cucurbita pepoCucurbita pepoThe ancestral species of the genus Cucurbita were present in the Americas before the arrival of humans,JOURNAL, Bemis, W. P., Whitaker, Thomas W., Thomas W. Whitaker, April 1969, The Xerophytic Cucurbita of Northwestern Mexico and Southwestern United States, Madroño (journal), Madroño, 20, 2, 33–41, California Botanical Society, 41423342, JOURNAL, Smith, Bruce D.,weblink Eastern North America as an Independent Center of Plant Domestication, Proceedings of the National Academy of Sciences of the United States of America, 15 August 2006, 103, 33, 12223–12228, 10.1073/pnas.0604335103, 16894156, 1567861, 2006PNAS..10312223S, and are native to the New World. The likely center of origin is southern Mexico, spreading south through what is now known as Mesoamerica, on into South America, and north to what is now the southwestern United States. Evolutionarily speaking, the genus is relatively recent in origin, dating back only to the Holocene, whereas the family Cucurbitaceae, in the shape of seeds similar to Bryonia, dates to the Paleocene.BOOK, Kubitzki, Klaus, Flowering Plants. Eudicots: Sapindales, Cucurbitales, Myrtaceae,weblink 2011, Springer Science & Business Media, Heidelberg, 978-3-642-14397-7, 120–122, The fossil record of Cucurbitaceae and indeed of the order Cucurbitales is sparse.. The oldest fossils are seeds from the Uppermost Paleocene and Lower Eocene London Clay (65MA).. Bryonia-like seeds from fossil beda at Tambov, Western Siberia date to the Lower Sarmat, 15-13 MA ago. Subfossil records of Cucurbita pepo have been dated to 8,000-7,000 B.C. at Guila Naquitz ..., those of C. moschata in the northern Peruvian Andes to up to 9,200 B.P., No species within the genus is entirely genetically isolated. C. moschata can intercross with all the others, though the hybrid offspring may not themselves be fertile unless they become polyploid. The genus was part of the culture of almost every native peoples group from southern South America to southern Canada. Modern-day cultivated Cucurbita are not found in the wild. Genetic studies of the mitochondrial gene nad1 show there were at least six independent domestication events of Cucurbita separating domestic species from their wild ancestors.JOURNAL, Sanjur, Oris I., Piperno, Dolores R., Andres, Thomas C., Wessel-Beaver, Linda, 2002, Phylogenetic Relationships among Domesticated and Wild Species of Cucurbita (Cucurbitaceae) Inferred from a Mitochondrial Gene: Implications for Crop Plant Evolution and Areas of Origin, Proceedings of the National Academy of Sciences of the United States of America, 99, 1, 535–540, National Academy of Sciences, Washington, DC, 3057572,weblink 10.1073/pnas.012577299, 2002PNAS...99..535S, 11782554, 117595, Species native to North America include C. digitata (calabazilla),WEB, Cucurbita digitata A. Gray,weblink United States Department of Agriculture, September 1, 2013, and C. foetidissima (buffalo gourd),WEB, Cucurbita ficifolia Bouché,weblink United States Department of Agriculture, September 1, 2013, C. palmata (coyote melon), and C. pepo. Some species, such as C. digitata and C. ficifolia, are referred to as gourds. Gourds, also called bottle-gourds, which are used as utensils or vessels, belong to the genus Lagenaria and are native to Africa. Lagenaria are in the same family and subfamily as Cucurbita but in a different tribe.WEB, Roberts, Katherine M., Cucurbita spp. and Lagenaria siceraria (Molina) - Standley Squash, Gourd, and Pumpkin; Bottle Gourd: Cucurbitaceae,weblink Washington University in St. Louis, March 27, 2012, December 31, 2014, The earliest known evidence of the domestication of Cucurbita dates back at least 8,000 years ago, predating the domestication of other crops such as maize and beans in the region by about 4,000 years.BOOK, Gibbon, Guy E., Ames, Kenneth M., Archaeology of Prehistoric Native America: An Encyclopedia, Routledge, New York, 1998, 238, 978-0-8153-0725-9, JOURNAL, Roush, Wade,weblink Archaeobiology: Squash Seeds Yield New View of Early American Farming, Science, 9 May 1997, 276, 5314, 894–895, American Association For the Advancement of Science, 10.1126/science.276.5314.894, This evidence was found in the Guilá Naquitz cave in Oaxaca, Mexico, during a series of excavations in the 1960s and 1970s, possibly beginning in 1959.JOURNAL, Schoenwetter, James, April 1974, Pollen Records of Guila Naquitz Cave, American Antiquity, 39, 2, 292–303, Society for American Archaeology, 279589, 10.2307/279589, JOURNAL, Benz, Bruce F., Archaeological Evidence of Teosinte Domestication From Guilá Naquitz, Oaxaca, 2005, Proceedings of the National Academy of Sciences, 98, 4, 2104–2106, 10.1073/pnas.98.4.2104, 11172083, 29389, 2001PNAS...98.2104B, Solid evidence of domesticated C. pepo was found in the Guilá Naquitz cave in the form of increasing rind thickness and larger peduncles in the newer stratification layers of the cave. By c. 8,000 years BP the C. pepo peduncles found are consistently more than {{convert|10|mm|in|sp=us}} thick. Wild Cucurbita peduncles are always below this 10 mm barrier. Changes in fruit shape and color indicate that intentional breeding of C. pepo had occurred by no later than 8,000 years BP.WEB,weblink Cucurbitaceae--Fruits for Peons, Pilgrims, and Pharaohs, University of California at Los Angeles, September 2, 2013, yes,weblink" title="web.archive.org/web/20131016003715weblink">weblink October 16, 2013, JOURNAL, Smith, Bruce D., 22 December 1989, Origins of Agriculture in Eastern North America,weblink Science, Washington, DC, 1566–1571, 10.1126/science.246.4937.1566, 17834420, 246, 4937, 1989Sci...246.1566S, JOURNAL, Smith, Bruce D., May 1997, The Initial Domestication of Cucurbita pepo in the Americas 10,000 Years Ago,weblink Science, Washington, DC, 10.1126/science.276.5314.932, 276, 5314, 932–934, During the same time frame, average rind thickness increased from {{convert|0.84|mm|in|sp=us}} to {{convert|1.15|mm|in|sp=us}}.BOOK, Feinman, Gary M., Manzanilla, Linda, Cultural Evolution: Contemporary Viewpoints,weblink 2000, Kluwer Academic, New York, 31, 978-0-306-46240-5, Squash was domesticated first, followed by maize and then beans, becoming part of the Three Sisters agricultural system of companion planting.JOURNAL, Landon, Amanda J., The "How" of the Three Sisters: The Origins of Agriculture in Mesoamerica and the Human Niche, Nebraska Anthropologist,weblink 2008, University of Nebraska-Lincoln, Lincoln, NE, 110–124, JOURNAL, Bushnell, G. H. S., The Beginning and Growth of Agriculture in Mexico, Philosophical Transactions of the Royal Society of London, 1976, London, 275, 936, 117–120, 10.1098/rstb.1976.0074, 1976RSPTB.275..117B, The English word "squash" derives from askutasquash (a green thing eaten raw), a word from the Narragansett language, which was documented by Roger Williams, the founder of Rhode Island, in his 1643 publication A Key Into the Language of America.WEB,weblink How Did the Squash Get its Name?, Library of Congress, October 10, 2013, Similar words for squash exist in related languages of the Algonquian family.JOURNAL, Paris, Harry S., 1989, Historical Records, Origins, and Development of the Edible Cultivar Groups of Cucurbita pepo (Cucurbitaceae), Economic Botany, 43, 4, 423–443, New York Botanical Garden Press, 4255187, 10.1007/bf02935916, BOOK, Charles L., Cutler, O Brave New Words: Native American Loanwords in Current English, University of Oklahoma Press, Norman, OK, 2000, 39–42, 978-0-8061-3246-4,weblink

Production

The family Cucurbitaceae has many species used as human food. Cucurbita species are some one of the most important of those, with the various species being prepared and eaten in many ways. Although the stems and skins tend to be more bitter than the flesh,WEB, Glover, Tony,weblink Bitter Cucumbers and Squash, Alabama Cooperative Extension System, November 23, 2014, the fruits and seeds of cultivated varieties are quite edible and need little or no preparation. The flowers and young leaves and shoot tips can also be consumed.BOOK, Lim, Tong Kwee, 2012, 283, Edible Medicinal And Non-Medicinal Plants: Volume 2, Fruits, Springer, New York, 978-94-007-1763-3, The seeds and fruits of most varieties can be stored for long periods of time, particularly the sweet-tasting winter varieties with their thick, inedible skins. Summer squash have a thin, edible skin. The seeds of both types can be roasted, eaten raw, made into pumpkin seed oil, ground into a flour or meal,JOURNAL, Lazos, E. S., Certain Functional Properties of Defatted Pumpkin Seed Flour, Plant Foods for Human Nutrition, 42, 3, July 1992, 257–273, 1502127, 10.1007/bf02193934, or otherwise prepared.Squashes are primarily grown for the fresh food market.WEB, Geisler, Malinda, May 2012, Squash,weblink Agricultural Marketing Resource Center, Iowa State University, October 13, 2013, The Food and Agriculture Organization of the United Nations (FAO) reported that the ranking of the top five squash-producing countries was stable between 2005 and 2009. Those countries are: China, India, Russia, the United States, and Egypt. By 2012, Iran had moved into the 5th slot, with Egypt falling to 6th. The top 10 countries in terms of metric tons of squashes produced are:WEB, Pumpkins, Squash, and Gourds,weblink Food and Agriculture Organization of the United Nations, 2012, October 13, 2013, (File:Sangkhaya fak thong.JPG|thumb|right|upright|alt=Slice of yellowish pumpkin custard with brown shell|Pumpkin custard made from kabocha, a cultivated variant of C. maxima){| class="wikitable sortable" style="margin: 1em auto 1em auto;"|+ Top ten squash producers â€” 2012! Country! Production(metric tons)PRC}} China6,140,840IND}} India4,424,200RUS}} Russia988,180USA}} USA778,630IRN}} Iran695,600EGY}} Egypt658,234MEX}} Mexico522,388UKR}} Ukraine516,900ITA}} Italy508,075TUR}} Turkey430,402 bgcolor=#eeeeee class="sortbottom"|Top 10 total15,663,449The only additional countries that rank in the top 20 where squashes are native are Cuba, which ranks 14th with 347,082 metric tons, and Argentina, which ranks 17th, with 326,900 metric tons. In addition to being the 4th largest producer of squashes in the world, the United States is the world's largest importer of squashes, importing 271,614 metric tons in 2011, 95 percent of that from Mexico. Within the United States, the states producing the largest amounts are Florida, New York, California, and North Carolina.{{clear}}

Nutrients

{{nutritional value | name=Summer squash, all varieties, raw| water=95 g| kJ=69| protein=1.2 g| fat=0.2 g| carbs=3.4 g| fiber=1.1 g| sugars=2.2 g| iron_mg=0.35| magnesium_mg=17| phosphorus_mg=38| potassium_mg=262| zinc_mg=0.29| manganese_mg=0.175| vitC_mg=17| thiamin_mg=0.048| riboflavin_mg=0.142| niacin_mg=0.487| pantothenic_mg=0.155| vitB6_mg=0.218| folate_ug=29| vitA_ug=10| betacarotene_ug=120| lutein_ug=2125| vitK_ug=3| source_usda = 1| note=Link to USDA Database entry, for comparison, see values for raw pumpkin}}As an example of Cucurbita, raw summer squash is 94% water, 3% carbohydrates, and 1% protein, with negligible fat content (table). In 100 grams, raw squash supplies 16 calories and is rich in vitamin C (20% of the Daily Value, DV), moderate in vitamin B6 and riboflavin (12–17% DV), but otherwise devoid of appreciable nutrient content (table), although the nutrient content of different Curcubita species may vary somewhat.WEB, What's So Great About Winter Squash?,weblink University of the District of Columbia, January 14, 2015, Pumpkin seeds contain vitamin E, crude protein, B vitamins and several dietary minerals (see nutrition table at pepita).JOURNAL, Mansour, Esam H., Dworschák, Erno, Lugasi, Andrea, Barna, Barna, Gergely, Anna, Nutritive Value of Pumpkin (Cucurbita Pepo Kakai 35) Seed Products, Journal of the Science of Food and Agriculture, 1993, 61, 1, 73–78, 10.1002/jsfa.2740610112, Also present in pumpkin seeds are unsaturated and saturated oils, palmitic, oleic and linoleic fatty acids,JOURNAL, Oil and tocopherol content and composition of pumpkin seed oil in 12 cultivars, Stevenson DG, Eller FJ, Wang L, Jane JL, Wang T, Inglett GE, J Agric Food Chem, 2007, 55, 10, 4005–16, 17439238,weblink 10.1021/jf0706979, as well as carotenoids.JOURNAL, 4013658, Supercritical carbon dioxide extraction of carotenoids from pumpkin (Cucurbita spp.): a review., Durante M, Lenucci MS, Mita G, Int J Mol Sci, 2014, 15, 4, 6725–40, 10.3390/ijms15046725, 24756094,

Toxins

Cucurbitin is an amino acid and a carboxypyrrolidine that is found in raw Cucurbita seeds.BOOK, Peirce, Andrea, The American Pharmaceutical Association Practical Guide to Natural Medicines, Stonesong Press, William Morrow & Company, New York, 1999, 212–214, 978-0-688-16151-4,weblink JOURNAL, Mihranian, Valentine H., Abou-Chaar, Charles I., Extraction, Detection, and Estimation of Cucurbitin in Cucurbita Seeds, Lloydia, American Society of Pharmacognosy, 31, 1, 1968, 23–29, It retards the development of parasitic flukes when administered to infected host mice, although the effect is only seen if administration begins immediately after infection.REPORT,weblink Assessment report on Cucurbita pepo L., pdf, Committee on Herbal Medicinal Products (HMPC), European Medicines Agency, 13 September 2011, 25–26, 21 November 2015, Cucurmosin is a ribosome inactivating protein found in the flesh and seed of Cucurbita,BOOK, Preedy, Victor R., Watson, Ronald Ross, Patel, Vinwood B., Nuts and Seeds in Health and Disease Prevention, 2011, Academic Press, London, 936, 978-0-12-375688-6, JOURNAL, Barbieri, L., Polito, L., Bolognesi, A., Ciani, M., Pelosi, E., Farini, V., Jha, A. K., Sharma, N., Vivanco, J. M., Chambery, A., Parente, A., Stirpe, F., Ribosome-inactivating Proteins in Edible Plants and Purification and Characterization of a New Ribosome-inactivating Protein From Cucurbita moschata, Biochimica et Biophysica Acta, 1760, 5, 783–792, May 2006, 16564632, 10.1016/j.bbagen.2006.01.002, notably Cucurbita moschata. Cucurmosin is more toxic to cancer cells than healthy cells.JOURNAL, Hou, Xiaomin, Meeha n, Edward J., Xie, Jieming, Huang, Mingdong, Chen, Minghuang, Chen, Liqing, Atomic Resolution Structure of Cucurmosin, a Novel Type 1 ribosome-inactivating Protein From the Sarcocarp of Cucurbita moschata, Journal of Structural Biology, 164, 1, 81–87, October 2008, 10.1016/j.jsb.2008.06.011, 18652900, Cucurbitacin is a plant steroid present in wild Cucurbita and in each member of the family Cucurbitaceae. Poisonous to mammals,JOURNAL, Tallamy, Douglas W., Krischik, Vera A., 1989, Variation and Function of Cucurbitacins in Cucurbita: An Examination of Current Hypotheses, The American Naturalist, 133, 6, 766–786, The University of Chicago Press, 2462036, 10.1086/284952, it is found in quantities sufficient to discourage herbivores. It makes wild Cucurbita and most ornamental gourds, with the exception of an occasional C. fraterna and C. sororia, bitter to taste.JOURNAL, Andres, Thomas C., 1987, Cucurbita fraterna, the Closest Wild Relative and Progenitor of C. pepo,weblink Cucurbit Genetics Cooperative Report, 10, 69–71, BOOK, Western Corn Rootworm: Ecology and Management, 67–71,weblink 2005, 4, Vidal, S., CAB International, Wallingford, UK, 978-0-85199-817-6, 10.1079/9780851998176.0000, yes,weblink" title="web.archive.org/web/20141129042743weblink">weblink 2014-11-29, Ingesting too much cucurbitacin can cause stomach cramps, diarrhea and even collapse.JOURNAL, Chen, Jian Chao, Chiu, Ming Hua, Nie, Rui Lin, Cordell, Geoffrey A., Qui, Samuel X., 2005, Cucurbitacins and Cucurbitane Glycosides: Structures and Biological Activities, Natural Product Reports, 22, 5, 386–399, 16010347, 10.1039/B418841C, This bitterness is especially prevalent in wild Cucurbita; in parts of Mexico the flesh of the fruits is rubbed on a woman's breast to wean children.WEB, Natural Hybridization of Wild Cucurbita sororia Group and Domesticated C. mixta in Southern Sonora, Mexico,weblink Merrick, Laura C., North Carolina State University, November 24, 2014, While the process of domestication has largely removed the bitterness from cultivated varieties, there are occasional reports of cucurbitacin causing illness in humans. Cucurbitacin is also used as a lure in insect traps.

Pests and diseases

Cucurbita species are used as food plants by the larvae of some Lepidoptera species, including the cabbage moth (Mamestra brassicae), Hypercompe indecisa, and the turnip moth (Agrotis segetum).WEB,weblink Pumpkin, Drugs.com, 2009, September 1, 2013, Cucurbita can be susceptible to the pest Bemisia argentifolii (silverleaf whitefly)JOURNAL, McAuslane, Heather J., Webb, Susan E., Elmstrom, Gary W., June 1996, Resistance in Germplasm of Cucurbita pepo to Silverleaf, a Disorder Associated with Bemisia argentifolii (Homoptera: Aleyrodidae), The Florida Entomologist, 79, 2, 206–221, Florida Entomological Society, Lutz, FL, 3495818, 10.2307/3495818,weblink as well as aphids (Aphididae), cucumber beetles (Acalymma vittatum and Diabrotica undecimpunctata howardi), squash bug (Anasa tristis), the squash vine borer (Melittia cucurbitae), and the two-spotted spidermite (Tetranychus urticae).WEB, Vegetable Pumpkin,weblink University of Illinois Extension, October 20, 2013, The squash bug causes major damage to plants because of its very toxic saliva.WEB, Common Name: Squash Bug,weblink University of Florida, October 26, 2013, The red pumpkin beetle (Raphidopalpa foveicollis) is a serious pest of cucurbits, especially the pumpkin, which it can defoliate.WEB,weblink Pumpkin beetle, Pests of Cucurbits, IndiaAgroNet.com, 20 May 2016, Cucurbits are susceptible to diseases such as bacterial wilt (Erwinia tracheiphila), anthracnose (Colletotrichum spp.), fusarium wilt (Fusarium spp.), phytophthora blight (Phytophthora spp. water molds), and powdery mildew (Erysiphe spp.). Defensive responses to viral, fungal, and bacterial leaf pathogens do not involve cucurbitacin.Species in the genus Cucurbita are susceptible to some types of mosaic virus including: cucumber mosaic virus (CMV), papaya ringspot virus-cucurbit strain (PRSV), squash mosaic virus (SqMV), tobacco ringspot virus (TRSV),WEB, Mosaic Diseases of Cucurbits,weblink University of Illinois, October 26, 2013, watermelon mosaic virus (WMV), and zucchini yellow mosaic virus (ZYMV).WEB, Virus Diseases of Cucurbit Crops,weblink Department of Agriculture, Government of Western Australia, December 31, 2014,weblink" title="web.archive.org/web/20141112044744weblink">weblink 2014-11-12, yes, JOURNAL, Roossinck, Marilyn J., Palukaitis, Peter, Rapid Induction and Severity of Symptoms in Zucchini Squash (Cucurbita pepo) Map to RNA 1 of Cucumber Mosaic Virus, Molecular Plant-Microbe Interactions, 1990, 3, 3, 188–192,weblink 10.1094/mpmi-3-188, JOURNAL, Havelda, Zoltan, Maule, Andrew J., Plant Cell, Complex Spatial Responses to Cucumber Mosaic Virus Infection in Susceptible Cucurbita pepo Cotyledons, October 2000, 12, 10, 1975–1986, 149134, 10.1105/tpc.12.10.1975, 11041891, WEB, Virus Diseases of Cucurbits,weblink Cornell University, October 1984, October 26, 2013, PRSV is the only one of these viruses that does not affect all cucurbits.JOURNAL, Provvidenti, R., Gonsalves, D., Occurrence of Zucchini Yellow Mosaic Virus in Cucurbits from Connecticut, New York, Florida, and California,weblink 68, 5, May 1984, Plant Disease, 443–446, 0191-2917, 10.1094/pd-69-443, SqMV and CMV are the most common viruses among cucurbits.WEB, Squash,weblink Texas A&M University, October 26, 2013, JOURNAL, Salama, El-Sayed A., Sill Jr., W. H., 1968, Resistance to Kansas Squash Mosaic Virus Strains Among Cucurbita Species, Transactions of the Kansas Academy of Science, 71, 1, 62–68, 3627399, 10.2307/3627399, Symptoms of these viruses show a high degree of similarity, which often results in laboratory investigation being needed to differentiate which one is affecting plants.

Human culture

Culinary uses

Long before European contact, Cucurbita had been a major food source for the native peoples of the Americas, and the species became an important food for European settlers, including the Pilgrims, even featuring at the first Thanksgiving. Commercially produced pumpkin commonly used in pumpkin pie is most often varieties of C. moschata; Libby's, by far the largest producer of processed pumpkin, uses a proprietary strain of the Dickinson pumpkin variety of C. moschata for its canned pumpkin.WEB, Richardson, R. W.,weblink Squash and Pumpkin, United States Department of Agriculture, Agricultural Research Service, National Plant Germplasm System, September 2, 2013, yes,weblink" title="web.archive.org/web/20150924160527weblink">weblink September 24, 2015, Other foods that can be made using members of this genus include biscuits, bread, cheesecake, desserts, donuts, granola, ice cream, lasagna dishes, pancakes, pudding, pumpkin butter,WEB,weblink Spiced Pumpkin Butter, Better Homes and Gardens, January 12, 2014, salads, soups, and stuffing.NEWS, Lynch, Rene,weblink Pumpkin Bread and 18 Other Pumpkin Recipes You Must Make Now, Los Angeles Times, October 1, 2013, October 8, 2013, Squash soup is a dish in African cuisine.BOOK, Garratt, N., Mango and Mint: Arabian, Indian, and North African Inspired Vegan Cuisine, PM Press, Tofu Hound Press, 2013, 978-1-60486-323-9,weblink 46, The xerophytic species are proving useful in the search for nutritious foods that grow well in arid regions.JOURNAL, Bemis, W. P., 1978, The Versatility of the Feral Buffalo Gourd, Cucurbita foetidissima HBK,weblink Cucurbit Genetics Cooperative Report, 1, 25, C. ficifolia is used to make soft and mildly alcoholic drinks.In India, squashes (ghia) are cooked with seafood such as prawns.BOOK, Jaffrey, Madhur, Madhur Jaffrey's Indian Cookery, 1982, BBC Books, London, 978-0-563-16491-3, 90, In France, marrows (courges) are traditionally served as a gratin, sieved and cooked with butter, milk, and egg, and flavored with salt, pepper, and nutmeg,BOOK, David, Elizabeth, French Country Cooking, 1987, 1951, Dorling Kindersley, London, 978-0-86318-251-8, 179, and as soups. In Italy, zucchini and larger squashes are served in a variety of regional dishes, such as cocuzze alla puviredda cooked with olive oil, salt and herbs from Apulia; as torta di zucca from Liguria, or torta di zucca e riso from Emilia-Romagna, the squashes being made into a pie filling with butter, ricotta, parmesan, egg, and milk; and as a sauce for pasta in dishes like spaghetti alle zucchine from Sicily.BOOK, della Salda, Anna Gosetti, Le Ricette Regionali Italiane, Italian, 1993, 1967, Solares, 107, 439, 878, 987, In Japan, squashes such as small C. moschata pumpkins (kabocha) are eaten boiled with sesame sauce, fried as a tempura dish, or made into balls with sweet potato and Japanese mountain yam.BOOK, Yoneda, Soei, The Heart of Zen Cuisine, 1987, 1982, Kodansha America, New York, 978-0-87011-848-7, 131, 133, 154,

Art, music, and literature

File:Zapallomuseolarco.jpg|thumb|left|upright|alt=Squash carved into a teapot shape|Moche squash ceramic. 300 C.E. Larco MuseumLarco MuseumAlong with maize and beans, squash has been depicted in the art work of the native peoples of the Americas for at least 2,000 years.WEB, Moche Decorated Ceramics, Metropolitan Museum of Art,weblink October 10, 2013, JOURNAL, Benson, Elizabeth P., 1983, A Moche "Spatula", Metropolitan Museum Journal, 18, 39–52, The University of Chicago Press, 1512797, 10.2307/1512797, For example, cucurbits are often represented in Moche ceramics.BOOK, The Spirit of Ancient Peru: Treasures from the Museo Arqueológico Rafael Larco Herrera, Thames and Hudson, Berrin Larco Museum, Katherine, 1997, New York, 978-0-500-01802-6, Though native to the western hemisphere, Cucurbita began to spread to other parts of the world after Christopher Columbus's arrival in the New World in 1492.JOURNAL, Whitaker, Thomas W., American Origin of Cultivated Cucurbits, Annals of the Missouri Botanical Garden, 34, 2, 101–111, Missouri Botanical Garden Press, St. Louis, MO, 1947, 10.2307/2394459, 2394459, JOURNAL, Whitaker, Thomas W., The Origin of the Cultivated Cucurbita, The American Naturalist, 90, 852, University of Chicago Press, Chicago, IL, 171–176, 1956, 10.1086/281923, 2458406, Until recently, the earliest known depictions of this genus in Europe was of Cucurbita pepo in De Historia Stirpium Commentarii Insignes in 1542 by the German botanist Leonhart Fuchs, but in 1992, two paintings, one of C. pepo and one of C. maxima, painted between 1515 and 1518, were identified in festoons at Villa Farnesina in Rome.JOURNAL, Janick, Jules, Paris, Harry S., The Cucurbit Images (1515–1518) of the Villa Farnesina, Rome, Annals of Botany, February 2006, 97, 2, 165–176, 10.1093/aob/mcj025, 2803371, 16314340, Also, in 2001 depictions of this genus were identified in Grandes Heures of Anne of Brittany (Les Grandes Heures d'Anne de Bretagne), a French devotional book, an illuminated manuscript created between 1503 and 1508. This book contains an illustration known as Quegourdes de turquie, which was identified by cucurbit specialists as C. pepo subsp. texana in 2006.JOURNAL, Paris, Harry S., Daunay, Marie-Christine, Pitrat, Michel, Janick, Jules, First Known Image of Cucurbita in Europe, 1503–1508, Annals of Botany, July 2006, 98, 1, 41–47, 10.1093/aob/mcl082, 2803533, 16687431, In 1952, Stanley Smith Master, using the pen name Edrich Siebert, wrote "The Marrow Song (Oh what a beauty!)" to a tune in {{music|time|6|8}} time. It became a popular hit in Australia in 1973,WEB, The Marrow Song (Oh What A Beauty!) by Edrich Siebert,weblink Songfacts, November 20, 2014, and was revived by the Wurzels in Britain on their 2003 album Cutler of the West.WEB, The Marrow Song,weblink The Wurzels, November 20, 2014, WEB, The Wurzels: Cutler of the West,weblink Last.fm, November 20, 2014, John Greenleaf Whittier wrote a poem entitled The Pumpkin in 1850.WEB,weblink The Pumpkin, Poets.org, November 20, 2014, "The Great Pumpkin" is a fictional holiday figure in the comic strip Peanuts by Charles M. Schulz.WEB,weblink 'It's the Great Pumpkin, Charlie Brown': 7 Things You Don't Know About Tonight's 'Peanuts' Special, Washington Post, Cavna, Michael, October 27, 2011, November 20, 2014,

Cleansing uses

C. foetidissima contains a saponin that can be obtained from the fruit and root. This can be used as a soap, shampoo, and bleach. Prolonged contact can cause skin irritation.JOURNAL, Cutler, H.C., Whitaker, T.W., 1961, History and Distribution of the Cultivated Cucurbits in the Americas, American Antiquity, 26, 4, 469–485, 278735, 10.2307/278735, {{subscription required}}BOOK, Heiser, Charles B., 2016, The Gourd Book, University of Oklahoma Press, Norman, OK, 9, 33–34, 978-0806155678,

Folk remedies

Cucurbita have been used in various cultures as folk remedies. Pumpkins have been used by Native Americans to treat intestinal worms and urinary ailments. This Native American remedy was adopted by American doctors in the early nineteenth century as an anthelmintic for the expulsion of worms.BOOK,weblink Environmental History of the Hudson River, Robert E. Henshaw, Albany, NY, State University of New York Press, 2011, 978-1-4384-4026-2, In southeastern Europe, seeds of C. pepo were used to treat irritable bladder and benign prostatic hyperplasia.BOOK,weblink Rational Phytotherapy: A Reference Guide for Physicians and Pharmacists, Volker Schulz, Springer, Munich, 5th, 2004, 304–305, 978-3-540-40832-1, In Germany, pumpkin seed is approved for use by the Commission E, which assesses folk and herbal medicine, for irritated bladder conditions and micturition problems of prostatic hyperplasia stages 1 and 2, although the monograph published in 1985 noted a lack of pharmacological studies that could substantiate empirically found clinical activity.WEB,weblink Pumpkin seed (Cucurbitae peponis semen), Heilpflanzen-Welt Bibliothek, March 25, 2015, The FDA in the United States, on the other hand, banned the sale of all such non-prescription drugs for the treatment of prostate enlargement in 1990.BOOK,weblink Tyler's Honest Herbal: A Sensible Guide to the Use of Herbs and Related Remedies, Steven, Foster, Varro E., Tyler, 131–132, Routledge, Binghamton, NY, 4th, 1999, 9781136745010, In China, C. moschata seeds were also used in traditional Chinese medicine for the treatment of the parasitic disease schistosomiasisJOURNAL, Xiao, S. H., Keiser, J., Chen, M. G., Tanner, M., Utzinger, J., Research and Development of Antischistosomal Drugs in the People's Republic of China a 60-year review, Advances in Parasitology, 2010, 73, 231–295, 10.1016/S0065-308X(10)73009-8, 20627145, 9780123815149, and for the expulsion of tape worms.BOOK,weblink Practical Therapeutics of Traditional Chinese Medicine, Yan, Wu, Warren, Fischer, Paradigm Publications, Taos, NM, 1997, 282–283, 978-0-912111-39-1, In Mexico, herbalists use C. ficifolia in the belief that it reduces blood sugar levels.JOURNAL, Andrade-Cetto, A., Heinrich, M., Mexican Plants With Hypoglycaemic Effect Used in the Treatment of Diabetes, Journal of Ethnopharmacology, 99, 3, 325–348, July 2005, 15964161, 10.1016/j.jep.2005.04.019,

Festivals

File:Squash Pyramid Display.JPG|thumb|100px|upright|alt=White, green, and orange squashes built into a Christmas tree shape|A pyramid of squashes in the Waterlily House, Kew GardensKew GardensCucurbita fruits including pumpkins and marrows are celebrated in festivals in countries such as Argentina, Bolivia,WEB, Sabores de Bolivia,weblink Cristina Olmos, Spanish, November 22, 2014, Britain, Canada,WEB, Smashing Success! Crowd Watches as Pumpkin Dropped on Old Car ... All for a Gourd Cause,weblink Calgary Sun, October 11, 2014, McMurray, Jenna, November 20, 2014, yes,weblink" title="web.archive.org/web/20141018144126weblink">weblink October 18, 2014, Croatia,WEB, Pumpkin Festival,weblink Tourist Board of Ivanić-Grad, 3 February 2016, France,WEB, Caligny: le Village de l'Orne où le Potiron est Roi,weblink info.fr, French, December 31, 2014, WEB, Fête de la Citrouille et des Cucurbitacées de Saint Laurent,weblink France Bleu, French, November 22, 2014, Germany, Italy,WEB, Festa della Zucca,weblink Associazione Pro Loco di Venzone, Italian, November 22, 2014, yes,weblink" title="archive.is/20130413065928weblink">weblink April 13, 2013, WEB, 21º Festa della Zucca SALZANO,weblink Pro Loco Salzano, Italian, November 22, 2014, WEB, Tra Meno un Mese Torna la Fiera Regionale della Zucca di Piozzo, ecco il Programma Ufficiale della 21esima Edizione,weblink Pro Loco di Piozzo, Italian, November 22, 2014, WEB, Fiesta de la Calabaza en Gavirate (Varese),weblink SarayT, Spanish, November 22, 2014, Japan,WEB, Autumn Events Calendar,weblink Asahikawa Tourism, November 20, 2014, Peru,WEB, Festival del Zapallo y de la Trucha de Curibaya se Realizará en la Plaza Quiñonez,weblink Radio Uno, Spanish, November 22, 2014, Portugal, Spain,WEB, V feria de las Calabazas Gigantes de Navarra (Valtierra),weblink Bardeneras, Spanish, November 22, 2014, yes,weblink" title="web.archive.org/web/20141129143423weblink">weblink November 29, 2014, Switzerland,WEB, Festivals and Events in Switzerland,weblink Travelsignposts, November 22, 2014, and the United States. Argentina holds an annual nationwide pumpkin festival Fiesta Nacional del Zapallo ("Squashes and Pumpkins National Festival"), in Ceres, Santa Fe,WEB, Fiesta Nacional del Zapallo,weblink La Fiesta Nacional, Spanish, November 22, 2014, yes,weblink" title="web.archive.org/web/20130804165532weblink">weblink August 4, 2013, on the last day of which a Reina Nacional del Zapallo ("National Queen of the Pumpkin") is chosen.WEB, Esperanza: Rocío Damiano fue elegida Reina Nacional del Zapallo en Ceres,weblink Ente Cultura, Spanish, November 22, 2014, WEB, Ceres: Presentaron la Fiesta Nacional del Zapallo,weblink El Litoral, Spanish, November 22, 2014, WEB, Presentacion Oficial – 43º Fiesta Nacional del Zapallo,weblink Ceres Online, Spanish, November 22, 2014, In Portugal the Festival da Abóbora de Lourinhã e Atalaia ("Squashes and Pumpkins Festival in Lourinhã and Atalaia") is held in Lourinhã city, called the Capital Nacional da Abóbora (the "National Capital of Squashes and Pumpkins").WEB, Festival da Abóbora de Lourinhã e Atalaia,weblink Festival da Abobora, Portuguese, November 22, 2014, yes,weblink" title="web.archive.org/web/20141111011222weblink">weblink November 11, 2014, Ludwigsburg, Germany annually hosts the world's largest pumpkin festival.NEWS, The World's Largest Pumpkin Festival in Germany,weblink USA Today, November 20, 2014, In Britain a giant marrow (zucchini) weighing {{convert|54.3177|kg|lb|sp=us}} was displayed in the Harrogate Autumn Flower Show in 2012.WEB, Giant Vegetables from UK Festival,weblink November 20, 2014, CBS News, In the US, pumpkin chucking is practiced competitively, with machines such as trebuchets and air cannons designed to throw intact pumpkins as far as possible.WEB, Punkin Chunkin,weblink Science Channel, November 20, 2014, NEWS, Campbell, Andy, Punkin Chunkin 2013: Will Someone Finally Launch A Pumpkin One Mile?,weblink Huffington Post, November 20, 2014, November 26, 2013, The Keene Pumpkin Fest is held annually in New Hampshire; in 2013 it held the world record for the most jack-o-lanterns lit in one place, 30,581 on October 19, 2013.NEWS,weblink Dandrea, Alyssa, October 20, 2013, Smiles, Pumpkins Abound as Keene Breaks Jack-o'-lantern Record, The Keene Sentinel, November 20, 2013, Halloween is widely celebrated with jack-o-lanterns made of large orange pumpkins carved with ghoulish faces and illuminated from inside with candles.WEB, History of the Jack O' Lantern,weblink History.com, A&E Networks, November 20, 2014, The pumpkins used for jack-o-lanterns are C. pepo,WEB, Stephens, James M.,weblink Pumpkin — Cucurbita spp., University of Florida, November 23, 2014, WEB, Baggett, J. R.,weblink Attempts to Cross Cucurbita moschata (Duch.) Poir. 'Butternut' and C. pepo L. 'Delicata', North Carolina State University, November 23, 2014, not to be confused with the ones typically used for pumpkin pie in the United States, which are C. moschata. Kew Gardens marked Halloween in 2013 with a display of pumpkins, including a towering pyramid made of many varieties of squash, in the Waterlily House during its "IncrEdibles" festival.WEB, IncrEdible! Kew Gardens to Unveil Towering Pyramid of Pumpkins in London,weblink Country Life, November 20, 2014, September 12, 2013,

See also

Notes

{{notes}}

References

{{Reflist}}

External links

  • {{Wiktionary-inline|Cucurbita}}
  • {{Commons-inline}}
  • {{Wikispecies-inline}}
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