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Cucurbita

January 19, 2024

Cucurbita (Latin for 'gourd')[3][4] is a genus of herbaceous fruits in the gourd family, Cucurbitaceae (also known as cucurbits or cucurbi), native to the Andes and Mesoamerica. Five edible species are grown and consumed for their flesh and seeds. They are variously known as squash, pumpkin, or gourd, depending on species, variety, and local parlance.[a] 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 the Cucurbita species.

Squash
Cucurbita fruits come in an assortment of colors and sizes.
Cross section of Cultivated Cucurbita of Canada
Scientific classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Cucurbitales
Family: Cucurbitaceae
Tribe: Cucurbiteae
Genus: Cucurbita
L.
Synonyms[1]

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. Although botanical fruits, Cucurbita gourds such as squash are typically cooked and eaten as vegetables. Pumpkins see more varied use, and are eaten both as vegetables and as desserts such as pumpkin pie.

Description edit

 
C. pepo pumpkins – 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 5 to 15 meters (15 to 50 feet) 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.[6] The morphological variation in the species C. pepo[7] and C. maxima[8] is so vast that its various subspecies and cultivars have been misidentified as totally separate species.[7]

 
The leaves of Cucurbita moschata often have white spots near the veins.

The 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.[9]

The species are monoecious, with unisexual male (staminate) and female (pistillate) flowers on a single plant 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 so that there appears to be one.[10][11] Female flowers have thick pedicels, and an inferior ovary with 3–5 stigmas that each have two lobes.[9][12] The female flowers of C. argyrosperma and C. ficifolia have larger corollas than the male flowers.[9] Female flowers of C. pepo have a small calyx, but the calyx of C. moschata male flowers is comparatively short.[9]

 
A variety of fruits displayed at the Real Jardín Botánico de Madrid in 2016

Cucurbita fruits are large and fleshy.[10] 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.[13][14] 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.[12] Fruit size varies considerably: wild fruit specimens can be as small as 4 centimeters (1+1⁄2 inches) and some domesticated specimens can weigh well over 300 kilograms (660 pounds).[9] The current world record was set in 2014 by Beni Meier of Switzerland with a 1,054 kg (2,323.7 lb) pumpkin.[15]

Reproductive biology edit

 
Cucurbita female flower with pollinating squash bees

All species of Cucurbita have 20 pairs of chromosomes.[16] 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.[6][17][18]

 
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,[19] 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.[20][21] Seedlessness is known to occur in certain cultivars of C. pepo.[22][23]

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.[24] The plant hormones ethylene and auxin are key in fruit set and development.[25] 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",[24] and male flowers are more frequent, an effect that appears due to reduced natural ethylene production within the plant stem.[26] 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.[20][27]

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.[28] Gibberellin is also involved in other developmental processes of plants such as seed and stem growth.[29]

Germination and seedling growth edit

 
Kabocha seedling seven days after being sown

Seeds with maximum germination potential develop (in C. moschata) by 45 days after anthesis, and seed weight reaches its maximum 70 days after anthesis.[30] Some varieties of C. pepo germinate best with eight hours of sunlight daily and a planting depth of 1.2 centimeters (1⁄2 in). Seeds planted deeper than 12.5 centimeters (5 in) are not likely to germinate.[31] 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.[32]

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.[19] 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.[33] Heavy metal contamination, including cadmium, has a significant negative impact on plant growth.[34] Cucurbita plants grown in the spring tend to grow larger than those grown in the autumn.[35]

Taxonomy edit

Cucurbita was formally described in a way that meets the requirements of modern botanical nomenclature by Linnaeus in his Genera Plantarum,[36] the fifth edition of 1754 in conjunction with the 1753 first edition of Species Plantarum.[37] Cucurbita pepo is the type species of the genus.[37][38] 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.[39]

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.[6][40] The five domesticated species are mostly isolated from each other by sterility barriers and have different physiological characteristics.[40] 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.[41] The buffalo gourd (C. foetidissima) has been used as an intermediary, as it can be crossed with all the common Cucurbita.[12]

 
An assortment of fruits of C. maxima and C. pepo

Various taxonomic treatments have been proposed for Cucurbita, ranging from 13 to 30 species.[3] 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:[6][42][43][44]

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:[16][47]

  • 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 edit

The full phylogeny of this genus is unknown, and research was ongoing in 2014.[48][49] The following cladogram of Cucurbita phylogeny is based upon a 2002 study of mitochondrial DNA by Sanjur and colleagues.[50]

Distribution and habitat edit

The ancestral species of the genus Cucurbita were present in the Americas before the arrival of humans,[51][52] and are native to the Americas. 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.[51] 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.[53] Recent genomic studies support the idea that the Cucurbita genus underwent a whole-genome duplication event, increasing their number of chromosomes and accelerating the rate at which their genomes evolve compared to other cucurbits.[54][55][56] 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.[40]

 
A festival-winning pumpkin in 2009 weighing 742 kilograms (1,636 lb)

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.[6] 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.[9]

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.[57] 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.[57]

 
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.[9] 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 1,800 meters (5,900 ft) 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.[9]

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 elevations ranging from 1,000 to 3,000 meters (3,300 to 9,800 ft) in areas with heavy rainfall. It does not hybridize well with the other cultivated species as it has significantly different enzymes and chromosomes.[9]

C. maxima originated in South America over 4,000 years ago,[50] 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.[58] 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.[6] Types of C. maxima include triloba,[59] zapallito,[60] zipinka,[61] Banana, Delicious, Hubbard, Marrow (C. maxima Marrow), Show, and Turban.[62]

 
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.[63] 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 20 to 30 centimeters (8 to 12 in) wide. It generally grows at low elevations in hot climates with heavy rainfall, but some varieties have been found above 2,200 meters (7,200 ft).[9] Groups of C. moschata include Cheese, Crookneck (C. moschata), and Bell.[62]

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.[6][9][64][65] Debates about the origin of C. pepo have been on-going since at least 1857.[66] 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.[6] A more recent theory by botanist Thomas Andres in 1987 is that descendants of C. fraterna hybridized with C. texana,[67] 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.[9][50][67][68] C. pepo may have appeared in the Old World before moving from Mexico into South America.[9] It is found from sea level to slightly above 2,000 meters (6,600 ft). Leaves have 3–5 lobes and are 20–35 centimeters (8–14 in) wide. All the subspecies, varieties, and cultivars are interfertile.[7] In 1986 Paris proposed a revised taxonomy of the edible cultivated C. pepo based primarily on the shape of the fruit, with eight groups .[57][69] All but a few C. pepo cultivars can be included in these groups.[9][69][70][71] There is one non-edible cultivated variety: C. pepo var. ovifera.[72]

A classification of cultivated C. pepo varieties based on Paris' eight groups and the one non-edible variety
Cultivar group Botanical name Image Description
Acorn C. pepo var. turbinata   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,[69] ex: Acorn squash[9][70][71]
Cocozzelle C. pepo var. Ionga   Summer squash, long round slender fruit that is slightly bulbous at the apex,[69] similar to fastigata, ex: Cocozelle von tripolis[9][70][71]
Crookneck C. pepo var. torticollia (also torticollis)   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,[69] ex: Crookneck squash[9][70][71]
Pumpkin C. pepo var. pepo
 
 Winter squash, creeping plant, round, oblate, or oval shape and round or flat on the ends,[69] ex: Pumpkin;[9][70][71] includes C. pepo subsp. pepo var. styriaca, used for Styrian pumpkin seed oil[73]
Scallop C. pepo var. clypeata; called C. melopepo by Linnaeus[7]   Summer squash, prefers half-shrubby habitat, flattened or slightly discoidal shape, with undulations or equatorial edges,[69] ex: Pattypan squash[9][70][71]
Straightneck C. pepo var. recticollis   Summer squash, shrubby plant, with yellow or golden fruit and verrucose rind, similar to var. torticollia but a stem end that narrows,[69] ex: Straightneck squash[9][70][71]
Vegetable marrow C. pepo var. fastigata   Summer and winter squashes, creeper traits and a semi-shrub, cream to dark green color, short round fruit with a slightly broad apex,[69] ex: Spaghetti squash (a winter variety)[9][70][71]
Zucchini/Courgette C. pepo var. cylindrica   Summer squash, presently the most common group of cultivars, origin is recent (19th century), semi-shrubby, cylindrical fruit with a mostly consistent diameter,[69] similar to fastigata, ex: Zucchini[9][70][71]
Ornamental gourds C. pepo var. ovifera   Non-edible,[72] 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.[64]

Ecology edit

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).[74] Cucurbita can be susceptible to the pest Bemisia argentifolii (silverleaf whitefly)[75] 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).[76] The squash bug causes major damage to plants because of its very toxic saliva.[77] The red pumpkin beetle (Aulacophora foveicollis) is a serious pest of cucurbits, especially the pumpkin, which it can defoliate.[78]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.).[76] Defensive responses to viral, fungal, and bacterial leaf pathogens do not involve cucurbitacin.[79]

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),[80] watermelon mosaic virus (WMV), and zucchini yellow mosaic virus (ZYMV).[81][82][83][84] PRSV is the only one of these viruses that does not affect all cucurbits.[81][85] SqMV and CMV are the most common viruses among cucurbits.[86][87] 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.[80]

Cultivation edit

History edit

 
Cucurbita pepo subsp. texana, from the Grandes Heures of Anne of Brittany, 1503–1508, f. 161, earliest depiction of cucurbits in Europe

The genus was part of the culture of almost every native peoples group from southern South America to southern Canada.[52] Modern-day cultivated Cucurbita are not found in the wild.[6] 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.[50] Species native to North America include C. digitata (calabazilla),[88] and C. foetidissima (buffalo gourd),[89] C. palmata (coyote melon), and C. pepo.[6] 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.[90]

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.[6][64][65][91] 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.[92][93] 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 10 millimeters (3⁄8 in) 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.[12][94][95] During the same time frame, average rind thickness increased from 0.84–1.15 millimeters (1⁄32–3⁄64 in).[96] Recent genomic studies suggest that Cucurbita argyrosperma was domesticated in Mexico, in the region that is currently known as the state of Jalisco.[97]

Squash was domesticated first, followed by maize and then beans, becoming part of the Three Sisters agricultural system of companion planting.[98][99] 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.[100] Similar words for squash exist in related languages of the Algonquian family.[57][101]

Production edit

Squash and gourd* production (2021)
Country Production
(millions of tonnes)
  China 7.4
  Ukraine 1.3
  Russia 1.2
  United States 1.1
  Turkey 0.8
  Mexico 0.7
  Italy 0.6
  Indonesia 0.5
  Egypt 0.4
World 23.4
*includes pumpkins
Source: FAOSTAT of the United Nations[102]

In 2021, world production of squashes (including gourds and pumpkins) was 23.4 million tonnes, led by China with 32% of the total (table). Ukraine, Russia, and the United States were secondary producers.

Toxicity edit

Cucurbitin is an amino acid and a carboxypyrrolidine that is found in raw Cucurbita seeds.[103][104] 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.[105]

Cucurmosin is a ribosome inactivating protein found in the flesh and seed of Cucurbita,[106][107] notably Cucurbita moschata.

Cucurbitacin is a plant steroid present in wild Cucurbita and in each member of the family Cucurbitaceae. Poisonous to mammals,[79] 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.[3][67][108] Ingesting too much cucurbitacin can cause stomach cramps, diarrhea and even collapse.[109] 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.[110] While the process of domestication has largely removed the bitterness from cultivated varieties,[3] there are occasional reports of cucurbitacin causing illness in humans.[3] Cucurbitacin is also used as a lure in insect traps.[108]

Uses edit

Nutrition edit

Summer squash, all varieties, raw
Nutritional value per 100 g (3.5 oz)
Energy69 kJ (16 kcal)
3.4 g
Sugars2.2 g
Dietary fiber1.1 g
0.2 g
1.2 g
VitaminsQuantity
%DV†
Vitamin A equiv.
1%
10 μg
1%
120 μg
2125 μg
Thiamine (B1)
4%
0.048 mg
Riboflavin (B2)
12%
0.142 mg
Niacin (B3)
3%
0.487 mg
Pantothenic acid (B5)
3%
0.155 mg
Vitamin B6
17%
0.218 mg
Folate (B9)
7%
29 μg
Vitamin C
20%
17 mg
Vitamin K
3%
3 μg
MineralsQuantity
%DV†
Iron
3%
0.35 mg
Magnesium
5%
17 mg
Manganese
8%
0.175 mg
Phosphorus
5%
38 mg
Potassium
6%
262 mg
Zinc
3%
0.29 mg
Other constituentsQuantity
Water95 g
Link to USDA Database entry, for comparison, see
†Percentages are roughly approximated using US recommendations for adults.
Source: USDA FoodData Central

As an example of Cucurbita, raw summer squash is 94% water, 3% carbohydrates, and 1% protein, with negligible fat content (table). In a 100-gram reference serving, raw squash supplies 69 kilojoules (16 kcal) of food energy 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.[111]

Pumpkin seeds contain vitamin E, crude protein, B vitamins and several dietary minerals (see nutrition table at pepita).[112] Also present in pumpkin seeds are unsaturated and saturated oils, palmitic, oleic and linoleic fatty acids,[113] as well as carotenoids.[114]

Culinary edit

The family Cucurbitaceae has many species used as human food.[9] Cucurbita species are some 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,[109][115] 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.[116] The seeds and fruits of most varieties can be stored for long periods of time,[6] particularly the sweet-tasting winter varieties with their thick, inedible skins.[117] Summer squash have a thin, edible skin. The seeds of both types can be roasted, eaten raw, made into pumpkin seed oil,[73] ground into a flour or meal,[118] or otherwise prepared.

Squashes are primarily grown for the fresh food market.[119]

 
Pumpkin custard made from kabocha, a cultivated variant of C. maxima

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.[12] 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.[120] 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,[121] salads, soups, and stuffing.[122] Squash soup is a dish in African cuisine.[123] The xerophytic species are proving useful in the search for nutritious foods that grow well in arid regions.[124] C. ficifolia is used to make soft and mildly alcoholic drinks.[9]

In India, squashes (ghiya) are cooked with seafood such as prawns.[125] 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,[126] 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.[127] 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.[128]

In culture edit

Art, music, and literature edit

 
Moche squash ceramic. 300 C.E. Larco Museum.

Along 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.[129][130] For example, cucurbits are often represented in Moche ceramics.[129][131]

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.[132][133] 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.[134] 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.[135]

In 1952, Stanley Smith Master, using the pen name Edrich Siebert, wrote "The Marrow Song (Oh what a beauty!)" to a tune in 6
8 time. It became a popular hit in Australia in 1973,[136] and was revived by the Wurzels in Britain on their 2003 album Cutler of the West.[137][138] John Greenleaf Whittier wrote a poem entitled The Pumpkin in 1850.[139] "The Great Pumpkin" is a fictional holiday figure in the comic strip Peanuts by Charles M. Schulz.[140]

Cleansing and personal care uses edit

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.[141][142] Pumpkin is also used in cosmetics.[143]

Folk remedies edit

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.[144] In southeastern Europe, seeds of C. pepo were used to treat irritable bladder and benign prostatic hyperplasia.[145] 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.[146] 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.[147]

In China, C. moschata seeds were also used in traditional Chinese medicine for the treatment of the parasitic disease schistosomiasis[148] and for the expulsion of tape worms.[149]

In Mexico, herbalists use C. ficifolia in the belief that it reduces blood sugar levels.[150]

Festivals edit

 
A pyramid of squashes in the Waterlily House, Kew Gardens, 2013

Cucurbita fruits including pumpkins and marrows are celebrated in festivals in countries such as Argentina, Austria,[151] Bolivia,[152] Britain, Canada,[153] Croatia,[154] France,[155][156] Germany, India, Italy,[157][158][159][160] Japan,[161] Peru,[162] Portugal, Spain,[163] Switzerland,[164] and the United States. Argentina holds an annual nationwide pumpkin festival Fiesta Nacional del Zapallo (lit. 'Squashes and Pumpkins National Festival'), in Ceres, Santa Fe,[165] on the last day of which a Reina Nacional del Zapallo (lit. 'National Queen of the Pumpkin') is chosen.[166][167][168] 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").[169] Ludwigsburg, Germany annually hosts the world's largest pumpkin festival.[170] In Britain a giant marrow (zucchini) weighing 54.3177 kilograms (119 lb 12 oz) was displayed in the Harrogate Autumn Flower Show in 2012.[171] 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.[172][173] 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.[174]

Halloween is widely celebrated with jack-o-lanterns made of large orange pumpkins carved with ghoulish faces and illuminated from inside with candles.[175] The pumpkins used for jack-o-lanterns are C. pepo,[176][177] not to be confused with the ones typically used for pumpkin pie in the United States, which are C. moschata.[120] 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.[178]

See also edit

Notes edit

  1. ^ 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.[5]

References edit

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External links edit

  •   The dictionary definition of Cucurbita at Wiktionary
  •   Media related to Cucurbita at Wikimedia Commons
  •   Data related to Cucurbita at Wikispecies

January 19, 2024
cucurbita, latin, gourd, genus, herbaceous, fruits, gourd, family, ceae, also, known, cucurbits, cucurbi, native, andes, mesoamerica, five, edible, species, grown, consumed, their, flesh, seeds, they, variously, known, squash, pumpkin, gourd, depending, specie. Cucurbita Latin for gourd 3 4 is a genus of herbaceous fruits in the gourd family Cucurbitaceae also known as cucurbits or cucurbi native to the Andes and Mesoamerica Five edible species are grown and consumed for their flesh and seeds They are variously known as squash pumpkin or gourd depending on species variety and local parlance a 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 the Cucurbita species SquashCucurbita fruits come in an assortment of colors and sizes Cross section of Cultivated Cucurbita of CanadaScientific classificationKingdom PlantaeClade TracheophytesClade AngiospermsClade EudicotsClade RosidsOrder CucurbitalesFamily CucurbitaceaeTribe CucurbiteaeGenus CucurbitaL Synonyms 1 Mellonia Gasp Melopepo Mill Ozodycus Raf Pepo Mill Pileocalyx Gasp Sphenantha Schrad Tristemon Scheele 1848 illegitimate homonym not Raf 1819 Juncaginaceae nor Raf 1838 Juncaceae nor Klotzsch 1838 Ericaceae 2 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 Although botanical fruits Cucurbita gourds such as squash are typically cooked and eaten as vegetables Pumpkins see more varied use and are eaten both as vegetables and as desserts such as pumpkin pie Contents 1 Description 1 1 Reproductive biology 1 1 1 Germination and seedling growth 2 Taxonomy 2 1 Phylogeny 3 Distribution and habitat 4 Ecology 5 Cultivation 5 1 History 5 2 Production 6 Toxicity 7 Uses 7 1 Nutrition 7 2 Culinary 8 In culture 8 1 Art music and literature 8 2 Cleansing and personal care uses 8 3 Folk remedies 8 4 Festivals 9 See also 10 Notes 11 References 12 External linksDescription edit nbsp C pepo pumpkins the two bright orange ones in center right and squashes C maxima all othersCucurbita 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 5 to 15 meters 15 to 50 feet 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 6 The morphological variation in the species C pepo 7 and C maxima 8 is so vast that its various subspecies and cultivars have been misidentified as totally separate species 7 nbsp The leaves of Cucurbita moschata often have white spots near the veins The 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 9 The species are monoecious with unisexual male staminate and female pistillate flowers on a single plant 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 so that there appears to be one 10 11 Female flowers have thick pedicels and an inferior ovary with 3 5 stigmas that each have two lobes 9 12 The female flowers of C argyrosperma and C ficifolia have larger corollas than the male flowers 9 Female flowers of C pepo have a small calyx but the calyx of C moschata male flowers is comparatively short 9 nbsp A variety of fruits displayed at the Real Jardin Botanico de Madrid in 2016Cucurbita fruits are large and fleshy 10 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 13 14 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 12 Fruit size varies considerably wild fruit specimens can be as small as 4 centimeters 1 1 2 inches and some domesticated specimens can weigh well over 300 kilograms 660 pounds 9 The current world record was set in 2014 by Beni Meier of Switzerland with a 1 054 kg 2 323 7 lb pumpkin 15 Reproductive biology edit nbsp Cucurbita female flower with pollinating squash beesAll species of Cucurbita have 20 pairs of chromosomes 16 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 6 17 18 nbsp Male flower part of the perianth and one filament removedWhen 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 19 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 20 21 Seedlessness is known to occur in certain cultivars of C pepo 22 23 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 24 The plant hormones ethylene and auxin are key in fruit set and development 25 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 24 and male flowers are more frequent an effect that appears due to reduced natural ethylene production within the plant stem 26 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 20 27 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 28 Gibberellin is also involved in other developmental processes of plants such as seed and stem growth 29 Germination and seedling growth edit nbsp Kabocha seedling seven days after being sownSeeds with maximum germination potential develop in C moschata by 45 days after anthesis and seed weight reaches its maximum 70 days after anthesis 30 Some varieties of C pepo germinate best with eight hours of sunlight daily and a planting depth of 1 2 centimeters 1 2 in Seeds planted deeper than 12 5 centimeters 5 in are not likely to germinate 31 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 32 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 19 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 33 Heavy metal contamination including cadmium has a significant negative impact on plant growth 34 Cucurbita plants grown in the spring tend to grow larger than those grown in the autumn 35 Taxonomy editCucurbita was formally described in a way that meets the requirements of modern botanical nomenclature by Linnaeus in his Genera Plantarum 36 the fifth edition of 1754 in conjunction with the 1753 first edition of Species Plantarum 37 Cucurbita pepo is the type species of the genus 37 38 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 39 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 6 40 The five domesticated species are mostly isolated from each other by sterility barriers and have different physiological characteristics 40 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 41 The buffalo gourd C foetidissima has been used as an intermediary as it can be crossed with all the common Cucurbita 12 nbsp An assortment of fruits of C maxima and C pepoVarious taxonomic treatments have been proposed for Cucurbita ranging from 13 to 30 species 3 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 6 42 43 44 C argyrosperma synonym C mixta cushaw pumpkin origin Mexico C kellyana origin Pacific coast of western Mexico C palmeri origin Pacific coast of northwestern Mexico C sororia origin Pacific coast Mexico to Nicaragua northeastern Mexico C digitata fingerleaf gourd origin southwestern United States USA northwestern Mexico C californica C cordata C cylindrata C palmata C ecuadorensis origin Ecuador s Pacific coast C ficifolia figleaf gourd chilacayote alcayota origin Mexico Panama northern Chile and Argentina C foetidissima stinking gourd buffalo gourd origin Mexico C scabridifolia likely a natural hybrid of C foetidissima and C pedatifolia 45 46 C galeottii little known origin Oaxaca Mexico C lundelliana origin Mexico Guatemala Belize C maxima winter squash pumpkin origin Argentina Bolivia Ecuador C andreana origin Argentina C moschata butternut squash Dickinson pumpkin golden cushaw origin Bolivia Colombia Ecuador Mexico Panama Puerto Rico Venezuela C okeechobeensis origin Florida C martinezii origin Mexican Gulf Coast and foothills C pedatifolia origin Queretaro Mexico C moorei C pepo field pumpkin summer squash zucchini vegetable marrow courgette acorn squash origin Mexico USA C fraterna origin Tamaulipas and Nuevo Leon Mexico C texana origin Texas USA C radicans calabacilla calabaza de coyote origin Central Mexico C graciliorThe 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 16 47 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 edit The full phylogeny of this genus is unknown and research was ongoing in 2014 48 49 The following cladogram of Cucurbita phylogeny is based upon a 2002 study of mitochondrial DNA by Sanjur and colleagues 50 Sechium eduleC ficifoliaC foetidissimaC maxima and C andreanaC ecuadorensisC martineziiC pepo subspp fraterna and oviferaC pepo subsp pepoC sororia in partC moschataC sororia in part and C argyrospermaDistribution and habitat editThis section may require copy editing for redundancy You can assist by editing it December 2023 Learn how and when to remove this template message The ancestral species of the genus Cucurbita were present in the Americas before the arrival of humans 51 52 and are native to the Americas 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 51 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 53 Recent genomic studies support the idea that the Cucurbita genus underwent a whole genome duplication event increasing their number of chromosomes and accelerating the rate at which their genomes evolve compared to other cucurbits 54 55 56 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 40 nbsp A festival winning pumpkin in 2009 weighing 742 kilograms 1 636 lb 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 6 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 9 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 57 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 57 nbsp 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 9 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 1 800 meters 5 900 ft 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 9 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 elevations ranging from 1 000 to 3 000 meters 3 300 to 9 800 ft in areas with heavy rainfall It does not hybridize well with the other cultivated species as it has significantly different enzymes and chromosomes 9 C maxima originated in South America over 4 000 years ago 50 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 58 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 6 Types of C maxima include triloba 59 zapallito 60 zipinka 61 Banana Delicious Hubbard Marrow C maxima Marrow Show and Turban 62 nbsp 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 63 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 20 to 30 centimeters 8 to 12 in wide It generally grows at low elevations in hot climates with heavy rainfall but some varieties have been found above 2 200 meters 7 200 ft 9 Groups of C moschata include Cheese Crookneck C moschata and Bell 62 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 6 9 64 65 Debates about the origin of C pepo have been on going since at least 1857 66 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 6 A more recent theory by botanist Thomas Andres in 1987 is that descendants of C fraterna hybridized with C texana 67 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 9 50 67 68 C pepo may have appeared in the Old World before moving from Mexico into South America 9 It is found from sea level to slightly above 2 000 meters 6 600 ft Leaves have 3 5 lobes and are 20 35 centimeters 8 14 in wide All the subspecies varieties and cultivars are interfertile 7 In 1986 Paris proposed a revised taxonomy of the edible cultivated C pepo based primarily on the shape of the fruit with eight groups 57 69 All but a few C pepo cultivars can be included in these groups 9 69 70 71 There is one non edible cultivated variety C pepo var ovifera 72 A classification of cultivated C pepo varieties based on Paris eight groups and the one non edible variety Cultivar group Botanical name Image DescriptionAcorn C pepo var turbinata nbsp 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 69 ex Acorn squash 9 70 71 Cocozzelle C pepo var Ionga nbsp Summer squash long round slender fruit that is slightly bulbous at the apex 69 similar to fastigata ex Cocozelle von tripolis 9 70 71 Crookneck C pepo var torticollia also torticollis nbsp 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 69 ex Crookneck squash 9 70 71 Pumpkin C pepo var pepo nbsp Winter squash creeping plant round oblate or oval shape and round or flat on the ends 69 ex Pumpkin 9 70 71 includes C pepo subsp pepo var styriaca used for Styrian pumpkin seed oil 73 Scallop C pepo var clypeata called C melopepo by Linnaeus 7 nbsp Summer squash prefers half shrubby habitat flattened or slightly discoidal shape with undulations or equatorial edges 69 ex Pattypan squash 9 70 71 Straightneck C pepo var recticollis nbsp Summer squash shrubby plant with yellow or golden fruit and verrucose rind similar to var torticollia but a stem end that narrows 69 ex Straightneck squash 9 70 71 Vegetable marrow C pepo var fastigata nbsp Summer and winter squashes creeper traits and a semi shrub cream to dark green color short round fruit with a slightly broad apex 69 ex Spaghetti squash a winter variety 9 70 71 Zucchini Courgette C pepo var cylindrica nbsp Summer squash presently the most common group of cultivars origin is recent 19th century semi shrubby cylindrical fruit with a mostly consistent diameter 69 similar to fastigata ex Zucchini 9 70 71 Ornamental gourds C pepo var ovifera nbsp Non edible 72 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 64 Ecology editCucurbita 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 74 Cucurbita can be susceptible to the pest Bemisia argentifolii silverleaf whitefly 75 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 76 The squash bug causes major damage to plants because of its very toxic saliva 77 The red pumpkin beetle Aulacophora foveicollis is a serious pest of cucurbits especially the pumpkin which it can defoliate 78 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 76 Defensive responses to viral fungal and bacterial leaf pathogens do not involve cucurbitacin 79 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 80 watermelon mosaic virus WMV and zucchini yellow mosaic virus ZYMV 81 82 83 84 PRSV is the only one of these viruses that does not affect all cucurbits 81 85 SqMV and CMV are the most common viruses among cucurbits 86 87 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 80 Cultivation editHistory edit nbsp Cucurbita pepo subsp texana from the Grandes Heures of Anne of Brittany 1503 1508 f 161 earliest depiction of cucurbits in EuropeThe genus was part of the culture of almost every native peoples group from southern South America to southern Canada 52 Modern day cultivated Cucurbita are not found in the wild 6 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 50 Species native to North America include C digitata calabazilla 88 and C foetidissima buffalo gourd 89 C palmata coyote melon and C pepo 6 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 90 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 6 64 65 91 This evidence was found in the Guila Naquitz cave in Oaxaca Mexico during a series of excavations in the 1960s and 1970s possibly beginning in 1959 92 93 Solid evidence of domesticated C pepo was found in the Guila 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 10 millimeters 3 8 in 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 12 94 95 During the same time frame average rind thickness increased from 0 84 1 15 millimeters 1 32 3 64 in 96 Recent genomic studies suggest that Cucurbita argyrosperma was domesticated in Mexico in the region that is currently known as the state of Jalisco 97 Squash was domesticated first followed by maize and then beans becoming part of the Three Sisters agricultural system of companion planting 98 99 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 100 Similar words for squash exist in related languages of the Algonquian family 57 101 Production edit Squash and gourd production 2021 Country Production millions of tonnes nbsp China 7 4 nbsp Ukraine 1 3 nbsp Russia 1 2 nbsp United States 1 1 nbsp Turkey 0 8 nbsp Mexico 0 7 nbsp Italy 0 6 nbsp Indonesia 0 5 nbsp Egypt 0 4World 23 4 includes pumpkinsSource FAOSTAT of the United Nations 102 In 2021 world production of squashes including gourds and pumpkins was 23 4 million tonnes led by China with 32 of the total table Ukraine Russia and the United States were secondary producers Toxicity editCucurbitin is an amino acid and a carboxypyrrolidine that is found in raw Cucurbita seeds 103 104 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 105 Cucurmosin is a ribosome inactivating protein found in the flesh and seed of Cucurbita 106 107 notably Cucurbita moschata Cucurbitacin is a plant steroid present in wild Cucurbita and in each member of the family Cucurbitaceae Poisonous to mammals 79 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 3 67 108 Ingesting too much cucurbitacin can cause stomach cramps diarrhea and even collapse 109 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 110 While the process of domestication has largely removed the bitterness from cultivated varieties 3 there are occasional reports of cucurbitacin causing illness in humans 3 Cucurbitacin is also used as a lure in insect traps 108 Uses editNutrition edit Summer squash all varieties rawNutritional value per 100 g 3 5 oz Energy69 kJ 16 kcal Carbohydrates3 4 gSugars2 2 gDietary fiber1 1 gFat0 2 gProtein1 2 gVitaminsQuantity DV Vitamin A equiv beta Carotenelutein zeaxanthin1 10 mg1 120 mg2125 mgThiamine B1 4 0 048 mgRiboflavin B2 12 0 142 mgNiacin B3 3 0 487 mgPantothenic acid B5 3 0 155 mgVitamin B617 0 218 mgFolate B9 7 29 mgVitamin C20 17 mgVitamin K3 3 mgMineralsQuantity DV Iron3 0 35 mgMagnesium5 17 mgManganese8 0 175 mgPhosphorus5 38 mgPotassium6 262 mgZinc3 0 29 mgOther constituentsQuantityWater95 gLink to USDA Database entry for comparison see values for raw pumpkinUnits mg micrograms mg milligrams IU International units Percentages are roughly approximated using US recommendations for adults Source USDA FoodData CentralAs an example of Cucurbita raw summer squash is 94 water 3 carbohydrates and 1 protein with negligible fat content table In a 100 gram reference serving raw squash supplies 69 kilojoules 16 kcal of food energy 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 111 Pumpkin seeds contain vitamin E crude protein B vitamins and several dietary minerals see nutrition table at pepita 112 Also present in pumpkin seeds are unsaturated and saturated oils palmitic oleic and linoleic fatty acids 113 as well as carotenoids 114 Culinary edit The family Cucurbitaceae has many species used as human food 9 Cucurbita species are some 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 109 115 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 116 The seeds and fruits of most varieties can be stored for long periods of time 6 particularly the sweet tasting winter varieties with their thick inedible skins 117 Summer squash have a thin edible skin The seeds of both types can be roasted eaten raw made into pumpkin seed oil 73 ground into a flour or meal 118 or otherwise prepared Squashes are primarily grown for the fresh food market 119 nbsp Pumpkin custard made from kabocha a cultivated variant of C maximaLong 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 12 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 120 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 121 salads soups and stuffing 122 Squash soup is a dish in African cuisine 123 The xerophytic species are proving useful in the search for nutritious foods that grow well in arid regions 124 C ficifolia is used to make soft and mildly alcoholic drinks 9 In India squashes ghiya are cooked with seafood such as prawns 125 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 126 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 127 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 128 In culture editArt music and literature edit nbsp Moche squash ceramic 300 C E Larco Museum Along 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 129 130 For example cucurbits are often represented in Moche ceramics 129 131 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 132 133 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 134 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 135 In 1952 Stanley Smith Master using the pen name Edrich Siebert wrote The Marrow Song Oh what a beauty to a tune in 68 time It became a popular hit in Australia in 1973 136 and was revived by the Wurzels in Britain on their 2003 album Cutler of the West 137 138 John Greenleaf Whittier wrote a poem entitled The Pumpkin in 1850 139 The Great Pumpkin is a fictional holiday figure in the comic strip Peanuts by Charles M Schulz 140 Cleansing and personal care uses edit 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 141 142 Pumpkin is also used in cosmetics 143 Folk remedies edit 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 144 In southeastern Europe seeds of C pepo were used to treat irritable bladder and benign prostatic hyperplasia 145 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 146 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 147 In China C moschata seeds were also used in traditional Chinese medicine for the treatment of the parasitic disease schistosomiasis 148 and for the expulsion of tape worms 149 In Mexico herbalists use C ficifolia in the belief that it reduces blood sugar levels 150 Festivals edit nbsp A pyramid of squashes in the Waterlily House Kew Gardens 2013Cucurbita fruits including pumpkins and marrows are celebrated in festivals in countries such as Argentina Austria 151 Bolivia 152 Britain Canada 153 Croatia 154 France 155 156 Germany India Italy 157 158 159 160 Japan 161 Peru 162 Portugal Spain 163 Switzerland 164 and the United States Argentina holds an annual nationwide pumpkin festival Fiesta Nacional del Zapallo lit Squashes and Pumpkins National Festival in Ceres Santa Fe 165 on the last day of which a Reina Nacional del Zapallo lit National Queen of the Pumpkin is chosen 166 167 168 In Portugal the Festival da Abobora de Lourinha e Atalaia Squashes and Pumpkins Festival in Lourinha and Atalaia is held in Lourinha city called the Capital Nacional da Abobora the National Capital of Squashes and Pumpkins 169 Ludwigsburg Germany annually hosts the world s largest pumpkin festival 170 In Britain a giant marrow zucchini weighing 54 3177 kilograms 119 lb 12 oz was displayed in the Harrogate Autumn Flower Show in 2012 171 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 172 173 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 174 Halloween is widely celebrated with jack o lanterns made of large orange pumpkins carved with ghoulish faces and illuminated from inside with candles 175 The pumpkins used for jack o lanterns are C pepo 176 177 not to be confused with the ones typically used for pumpkin pie in the United States which are C moschata 120 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 178 See also editList of gourds and squashes in the genus Cucurbita List of squash and pumpkin dishesNotes edit 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 5 References edit Cucurbita L Tropicos Missouri Botanical Garden Archived from the original on August 7 2019 Retrieved January 12 2014 Tristemon Tropicos Missouri Botanical Garden Archived from the original on February 20 2022 Retrieved January 14 2015 a b c d e Burrows George E Tyrl Ronald J 2013 Toxic Plants of North America Oxford Wiley Blackwell pp 389 391 ISBN 978 0 8138 2034 7 Gledhill David 2008 The Names of Plants Cambridge Cambridge University Press p 127 ISBN 9780521866453 Ferriol Maria Pico Belen 2007 3 Handbook of Plant Breeding Vegetables I New York Springer p 317 ISBN 978 0 387 72291 7 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 Bouche a b c d e f g h i j k l Nee Michael 1990 The Domestication of 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February 2016 Caligny le Village de l Orne ou le Potiron est Roi in French info fr 4 October 2014 Retrieved December 31 2014 Fete de la Citrouille et des Cucurbitacees de Saint Laurent in French France Bleu 2 October 2014 Retrieved November 22 2014 Festa della Zucca in Italian Associazione Pro Loco di Venzone Archived from the original on April 13 2013 Retrieved November 22 2014 21º Festa della Zucca SALZANO in Italian Pro Loco Salzano Archived from the original on November 29 2014 Retrieved November 22 2014 Tra Meno un Mese Torna la Fiera Regionale della Zucca di Piozzo ecco il Programma Ufficiale della 21esima Edizione in Italian Pro Loco di Piozzo Archived from the original on November 29 2014 Retrieved November 22 2014 Fiesta de la Calabaza en Gavirate Varese in Spanish SarayT 8 October 2013 Archived from the original on 25 December 2014 Retrieved November 22 2014 Autumn Events Calendar Asahikawa Tourism Archived from the original on November 3 2014 Retrieved November 20 2014 Festival del Zapallo y de la Trucha de Curibaya se Realizara en la Plaza Quinonez in Spanish Radio Uno 10 April 2014 Archived from the original on 29 November 2014 Retrieved November 22 2014 V feria de las Calabazas Gigantes de Navarra Valtierra in Spanish Bardeneras Archived from the original on November 29 2014 Retrieved November 22 2014 Festivals and Events in Switzerland Travelsignposts 15 February 2009 Archived from the original on 2 December 2014 Retrieved November 22 2014 Fiesta Nacional del Zapallo in Spanish La Fiesta Nacional Archived from the original on August 4 2013 Retrieved November 22 2014 Esperanza Rocio Damiano fue elegida Reina Nacional del Zapallo en Ceres in Spanish Ente Cultura Archived from the original on November 29 2014 Retrieved November 22 2014 Ceres Presentaron la Fiesta Nacional del Zapallo in Spanish El Litoral Archived from the original on November 29 2014 Retrieved November 22 2014 Presentacion Oficial 43º Fiesta Nacional del Zapallo in Spanish Ceres Online 23 May 2014 Archived from the original on 29 November 2014 Retrieved November 22 2014 Festival da Abobora de Lourinha e Atalaia in Portuguese Festival da Abobora Archived from the original on November 11 2014 Retrieved November 22 2014 The World s Largest Pumpkin Festival in Germany USA Today Archived from the original on November 29 2014 Retrieved November 20 2014 Giant Vegetables from UK Festival CBS News CBS News 14 September 2012 Archived from the original on November 29 2014 Retrieved November 20 2014 Punkin Chunkin Science Channel Archived from the original on November 13 2014 Retrieved November 20 2014 Campbell Andy November 26 2013 Punkin Chunkin 2013 Will Someone Finally Launch A Pumpkin One Mile Huffington Post Archived from the original on November 29 2014 Retrieved November 20 2014 Dandrea Alyssa October 20 2013 Smiles Pumpkins Abound as Keene Breaks Jack o lantern Record The Keene Sentinel Archived from the original on November 23 2013 Retrieved November 20 2013 History of the Jack O Lantern History com A amp E Networks Archived from the original on November 18 2014 Retrieved November 20 2014 Stephens James M Pumpkin Cucurbita spp University of Florida Archived from the original on October 31 2014 Retrieved November 23 2014 Baggett J R Attempts to Cross Cucurbita moschata Duch Poir Butternut and C pepo L Delicata North Carolina State University Archived from the original on February 6 2014 Retrieved November 23 2014 IncrEdible Kew Gardens to Unveil Towering Pyramid of Pumpkins in London Country Life September 12 2013 Archived from the original on November 29 2014 Retrieved November 20 2014 External links edit nbsp The dictionary definition of Cucurbita at Wiktionary nbsp Media related to Cucurbita at Wikimedia Commons nbsp Data related to Cucurbita at Wikispecies Retrieved from https en wikipedia org w index php title Cucurbita amp oldid 1192788444, wikipedia, wiki, book, books, library,

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