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''Cucurbita'' (
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally spoken in the area around Rome, known as Latium. Through the power of the Roman Republic, it became the dominant la ...

Latin
for
gourd Gourds include the fruits of some flowering plant The flowering plants, also known as Angiospermae (), or Magnoliophyta (), are the most diverse group of Embryophyte, land plants, with 64 Order(biology), orders, 416 Family (biology), families, ...

gourd
) is a
genus Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank In biological classification In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure ...
of
herbaceous Herbaceous plants are vascular plant Vascular plants (from Latin ''vasculum'': duct), also known as Tracheophyta (the tracheophytes , from the Greek ''trācheia''), form a large group of plants ( 300,000 accepted known species) that are defined ...
vegetables Vegetables are parts of plants that are consumed by humans or other animals as food. The original meaning is still commonly used and is applied to plants collectively to refer to all edible plant matter, including the flowers, fruit In ...

vegetables
in the gourd
family In , family (from la, familia) is a of people related either by (by recognized birth) or (by marriage or other relationship). The purpose of families is to maintain the well-being of its members and of society. Ideally, families would off ...
,
Cucurbitaceae The Cucurbitaceae, also called cucurbits or the gourd family, are a plant family (biology), family consisting of about 965 species in around 95 genera, of which the most important to humans are: *''Cucurbita'' – Squash (plant), squash, pumpkin, ...

Cucurbitaceae
(also known as ''cucurbits'' or ''cucurbi'') native to the
Andes The Andes, Andes Mountains or Andean Mountains ( es, Cordillera de los Andes) are the List of mountain ranges#Mountain ranges by length, longest continental mountain range in the world, forming a continuous highland along the western edge of Sout ...

Andes
and
Mesoamerica Mesoamerica is a historical and important region In geography, regions are areas that are broadly divided by physical characteristics ( physical geography), human impact characteristics ( human geography), and the interaction of humanity and th ...
. Five species are grown worldwide for their edible
vegetable Vegetables are parts of plants that are consumed by humans or other animals as food. The original meaning is still commonly used and is applied to plants collectively to refer to all edible plant matter, including the flowers, fruits, stems ...

vegetable
, variously known as squash,
pumpkin A pumpkin is a cultivar of winter squash that is round with smooth, slightly ribbed skin, and is most often deep yellow to orange in coloration. The thick shell contains the seeds and pulp. The name is most commonly used for cultivars of ''Cucu ...

pumpkin
, or gourd, depending on species,
variety Variety may refer to: Science and technology Mathematics * Algebraic variety, the set of solutions of a system of polynomial equations * Variety (universal algebra), classes of algebraic structures defined by equations in universal algebra Hort ...
, and local parlance, and for their seeds. Other kinds of gourd, also called bottle-gourds, are native to Africa and belong to the genus ''
Lagenaria ''Lagenaria'' is a genus of gourd-bearing vines in the squash family (Cucurbitaceae). ''Lagenaria'' contains six species, all of which are indigenous to tropical Africa.tribe The term tribe is used in many different contexts to refer to a category of human Humans (''Homo sapiens'') are the most populous and widespread species of primates, characterized by bipedality, opposable thumbs, hairlessness, and intellig ...
. 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
tendril upA curling tendril In botany, a tendril is a specialized Plant stem, stem, leaf or Petiole (botany), petiole with a threadlike shape used by climbing plants for support and attachment, as well as cellular invasion by parasitic plants such as '' ...

tendril
s, 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 Pollen Tube Diagram Pollen is a powdery substance consisting of pollen grains which are microsporophytes of seed plants The spermatophytes, also known as phanerogams (taxon Phanerogamae) or phaenogams (taxon Phaenogamae), comprise those plan ...

pollen
. Many North and Central American species are visited by specialist
bee Bees are flying insect Insects or Insecta (from Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken in the area around Rome, known ...
pollinator A pollinator is an animal that moves pollen File:Pollen Tube.svg, Pollen Tube Diagram Pollen is a powdery substance consisting of pollen grains which are Sporophyte, microsporophytes of spermatophyta, seed plants, which produce male gametes ...

pollinator
s, but other insects with more general feeding habits, such as
honey bee A honey bee (also spelled honeybee) is a eusocial Eusociality (from Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Sou ...

honey bee
s, also visit. There is debate about the
taxonomy Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term may also refer to a specific classification scheme. Originally used only about biological ...
of the genus, as the number of accepted species varies from 13 to 30. The five
domesticated Domestication is a sustained multi-generational relationship in which one group of organisms assumes a significant degree of influence over the reproduction and care of another group to secure a more predictable supply of resources from that seco ...

domesticated
species are ''
Cucurbita argyrosperma ''Cucurbita argyrosperma'', also the Japanese pie pumpkin or cushaw pumpkin, and silver-seed gourd, is a species of winter squash originally from the south of Mexico. This annual plant, annual herbaceous plant is cultivated in the Americas for its ...
'', '' C. ficifolia'', '''', '''', and '' C. pepo''. All of these can be treated as
winter squash Winter squash is an annual vegetable representing several squash (plant), squash species within the genus ''Cucurbita''. It differs from summer squash in that it is harvested and eaten in the mature stage when the seeds within have matured fully an ...
because the full-grown fruits can be stored for months; however, ''C. pepo'' includes some cultivars that are better used only as
summer squash 240px, 'Zephyr', a variety of straightnecked summer squash Summer squash are Cucurbita, squashes that are harvested when immature, while the rind is still tender and edible. Nearly all summer squashes are varieties of ''Cucurbita pepo'', though no ...
. The fruits of the genus ''Cucurbita'' are good sources of
nutrient A nutrient is a substance Substance may refer to: * Substance (Jainism), a term in Jain ontology to denote the base or owner of attributes * Chemical substance, a material with a definite chemical composition * Matter, anything that has mass and t ...
s, such as
vitamin A Vitamin A is a group of unsaturated nutritional organic compounds , CH4; is among the simplest organic compounds. In chemistry, organic compounds are generally any chemical compounds that contain carbon-hydrogen chemical bond, bonds. Due to ...

vitamin A
and
vitamin C Vitamin C (also known as ascorbic acid and ascorbate) is a vitamin found in various foods and sold as a dietary supplement. It is used to prevent and treat scurvy. Vitamin C is an Nutrient#Essential nutrients, essential nutrient involved in t ...

vitamin C
, among other nutrients according to species. The fruits have many culinary uses including
pumpkin pie Pumpkin pie is a dessert Dessert () is a course (food), course that concludes a meal. The course consists of sweet foods, such as confections, and possibly a beverage such as dessert wine and liqueur. In some parts of the world, such as much o ...

pumpkin pie
, biscuits, bread, desserts, puddings, beverages, and soups.


Description

''Cucurbita'' species fall into two main groups. The first group are
annual Annual may refer to: *Annual publication, periodical publications appearing regularly once per year **Yearbook **Literary annual *Annual plant *Annual report *Annual giving *Annual, Morocco, a settlement in northeastern Morocco *Annuals (band), a ...
or short-lived
perennial A perennial plant or simply perennial is a plant Plants are predominantly photosynthetic eukaryotes of the Kingdom (biology), kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and incl ...
vines and are
mesophyticMesophytes are terrestrial plants which are neither adapted to particularly dry nor particularly wet environments. An example of a mesophytic habitat would be a rural temperate meadow, which might contain Solidago, goldenrod, Trifolium, clover, Leuca ...
, i.e. they require a more or less continuous water supply. The second group are
perennials A perennial plant or simply perennial is a plant that lives more than two years. The term (''wikt:per-#Prefix, per-'' + ''wikt:-ennial#Suffix, -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annual pla ...
growing in arid zones and so are
xerophytic A xerophyte (from Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is approxima ...

xerophytic
, tolerating dry conditions. Cultivated ''Cucurbita'' species were derived from the first group. Growing in height or length, the plant stem produces
tendril upA curling tendril In botany, a tendril is a specialized Plant stem, stem, leaf or Petiole (botany), petiole with a threadlike shape used by climbing plants for support and attachment, as well as cellular invasion by parasitic plants such as '' ...

tendril
s 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. The morphological variation in the species ''C. pepo'' and ''C. maxima'' is so vast that its various
subspecies In biological classification In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactio ...
and cultivars have been misidentified as totally separate species. The typical cultivated ''Cucurbita'' species has five-lobed or 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
trichome Image:Cap1033-botao1.jpg, Flower bud of a ''Capsicum pubescens'' plant, with many trichomes Trichomes ( or ), from the Greek language, Greek τρίχωμα (trichōma) meaning "hair", are fine outgrowths or appendages on plants, algae, lichens, a ...
s, which are often hardened and sharp. Spring-like tendrils grow from each
node In general, a node is a localized swelling (a "knot A knot is an intentional complication in Rope, cordage which may be practical or decorative, or both. Practical knots are classified by function, including hitches, bends, loop knots, and splic ...

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 Puberty is the process of physical changes through which a child's Human body, body matures into an adult body capable of sexual reproduction. It is initiated by hormone, hormonal signals from the brain to the gonads: the ovary, ovaries in a gi ...

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. The species are
monoecious Monoecy (; adjective form: monoecious ) is a sexual system in seed plants The spermatophytes (; ), also known as phanerogams (taxon Phanerogamae) or phaenogams (taxon Phaenogamae), comprise those plant Plants are predominantly photosynthet ...
, with unisexual male (
staminate The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament an ...
) and female (
pistil '' stigmas and style Gynoecium (, from Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided in ...
late) flowers on a single plant and these grow singly, appearing from the leaf axils. Flowers have five fused yellow to orange petals (the
corolla Corolla may refer to: *Corolla (botany) upright=1.4, Diagram showing the parts of a mature flower. In this example the perianth is separated into a calyx (sepals) and corolla (petals) The perianth (perigonium, perigon or perigone in monocots) is ...

corolla
) and a green bell-shaped calyx. Male flowers in Cucurbitaceae generally have five stamens, but in ''Cucurbita'' there are only three, and their
anther The stamen (plural The plural (sometimes list of glossing abbreviations, abbreviated ), in many languages, is one of the values of the grammatical number, grammatical category of number. The plural of a noun typically denotes a quantity greate ...
s are joined together so that there appears to be one. Female flowers have thick
pedicels Image:Delphinium nuttallianum 15498.JPG, The inflorescence of ''Delphinium nuttallianum''. Each flower is held on a pedicel from one to several centimeters long. A pedicel is a stem that attaches a single flower to the inflorescence. Such inflores ...
, and an
inferior ovary In the flowering plant Flowering plants include multiple members of the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or r ...
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. ''Cucurbita'' fruits are large and fleshy. Botanists classify the ''Cucurbita'' fruit as a pepo, which is a special type of
berry A berry is a small, pulpy, and often edible fruit In botany Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who speci ...
derived from an inferior ovary, with a thick outer wall or rind with
hypanthium In angiosperms, a hypanthium or floral cup is a structure where basal portions of the calyx, the corolla, and the stamens form a cup-shaped tube. It is sometimes called a floral tube, a term that is also used for corolla tube and calyx tube. I ...
tissue forming an
exocarp Fruit anatomy is the plant anatomy Plant anatomy or phytotomy is the general term for the study of the internal structure A structure is an arrangement and organization of interrelated elements in a material object or system A system is a ...
around the ovary, and a fleshy interior composed of
mesocarp Fruit anatomy is the plant anatomy Plant anatomy or phytotomy is the general term for the study of the internal structure A structure is an arrangement and organization of interrelated elements in a material object or system A system is a ...
and
endocarp Fruit anatomy is the plant anatomy Plant anatomy or phytotomy is the general term for the study of the internal structure A structure is an arrangement and organization of interrelated elements in a material object or system A system is a ...
. The term "pepo" is used primarily for Cucurbitaceae fruits, where this fruit type is common, but the fruits of ''
Passiflora ''Passiflora'', known also as the passion flowers or passion vines, is a genus Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as we ...

Passiflora
'' and ''
Carica ''Carica'' is a genus of flowering plant Flowering plants include multiple members of the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek Greek may refer to: Greece Anything of, from, or ...

Carica
'' are sometimes also pepos. 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
cotyledon File:Germination-en.svg, 335px, Schematic of epigeal vs hypogeal germination A cotyledon (; ; ; , gen. (), ) is a significant part of the embryo within the seed of a plant, and is defined as "the embryonic leaf in seed-bearing plants, one o ...
s. Fruit size varies considerably: wild fruit specimens can be as small as and some domesticated specimens can weigh well over . The current world record was set in 2014 by Beni Meier of
Switzerland , french: Suisse(sse), it, svizzero/svizzera or , rm, Svizzer/Svizra , government_type = under an , leader_title1 = , leader_name1 = , leader_title2 = , leader_name2 = , legislatur ...

Switzerland
with a pumpkin.


Taxonomy

''Cucurbita'' was formally described in a way that meets the requirements of modern
botanical nomenclature Botanical nomenclature is the formal, scientific naming of plants. It is related to, but distinct from taxonomy Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such ...
by
Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his Nobility#Ennoblement, ennoblement as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomia ...

Linnaeus
in his ''
Genera Plantarum ''Genera Plantarum'' is a publication of Swedish naturalist Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his Nobility#Ennoblement, ennoblement as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a S ...
'', the fifth edition of 1754 in conjunction with the 1753 first edition of ''
Species Plantarum ' (Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken in the area around Rome, known as Latium. Through the power of the Roman Republic, i ...
''. ''Cucurbita pepo'' is the
type species In zoological nomenclature The International Code of Zoological Nomenclature (ICZN) is a widely accepted Convention (norm), convention in zoology that rules the formal scientific name, scientific naming of organisms treated as animals. It is als ...
of the genus. 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 ''Watermelon'' (''Citrullus lanatus'') is a flowering plant The flowering plants, also known as Angiospermae (), or Magnoliophyta (), are the most diverse group of Embryophyte, land plants, with 64 Order(biology), orders, 416 Family (biolo ...

Citrullus lanatus
'') and ''C. lagenaria'' (now ''
Lagenaria siceraria Calabash (''Lagenaria siceraria''), also known as bottle gourd, white-flowered gourd, long melon, New Guinea bean and Tasmania bean, is a vine grown for its fruit. It can be either harvested young to be consumed as a vegetable, or harvested mat ...

Lagenaria siceraria
'') (both are not ''Cucurbita'' but are in the family Cucurbitaceae. The ''Cucurbita digitata'', ''C. foetidissima'', ''C. galeotti'', and ''C. pedatifolia''
species group In biology, a species complex is a group of closely related organisms that are so similar in appearance that the boundaries between them are often unclear. Terms that are sometimes used synonymously but have more precise meanings are cryptic speci ...
s are xerophytes, arid zone perennials with storage roots; the remainder, including the five domesticated species, are all mesophytic annuals or short-life perennials with no storage roots. 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. 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''. Various
taxonomic Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term may also refer to a specific classification scheme. Originally used only about biological ...
treatments have been proposed for ''Cucurbita'', ranging from 13 to 30 species. 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: * '' C. argyrosperma'' (
synonym A synonym is a word, morpheme A morpheme is the smallest meaningful lexical item in a language. A morpheme is not a word. The difference between a morpheme and a word is that a morpheme bound and free morphemes, sometimes does not stand alone, ...
''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 In biology, a hybrid is the offspring resulting from combining the qualities of two organisms of different breeds, varieties, species or genera through sexual reproduction. Hybrids are not always intermediates between their parents (such as in b ...
of ''C. foetidissima'' and ''C. pedatifolia'' * '' C. galeottii'' is little known; origin:
Oaxaca ) , population_note = , population_rank = 10th , timezone1 = CST CST or Cst may refer to: Time zones * Central Standard Time, North America's Central Time Zone: UTC−06:00 * China Standard Time: UTC+08:00 * Cuba ...

Oaxaca
, Mexico * '' C. lundelliana'', origin: Mexico, Guatemala, Belize * '''' – winter squash, pumpkin; origin: Argentina, Bolivia, Ecuador ** '' C. andreana'', origin – Argentina * '''' – 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:
Querétaro Querétaro (), officially the Free and Sovereign State of Querétaro ( es, Estado Libre y Soberano de Querétaro, links=no; Otomi language, Otomi: ''Hyodi Ndämxei''), is one of the Political divisions of Mexico, 32 federal entities of Mexico. I ...

Querétaro
, Mexico ** '' C. moorei'' * '' C. pepo'' – field pumpkin, summer squash, zucchini, vegetable marrow, courgette, acorn squash; origin: Mexico, USA ** '' C. fraterna'', origin:
Tamaulipas Tamaulipas (), officially the Free and Sovereign State of Tamaulipas ( es, Estado Libre y Soberano de Tamaulipas), is one of the 31 states which, along with Mexico City ) , blank1_name_sec1 = Human Development Index, HDI , blank1_ ...

Tamaulipas
and
Nuevo León Nuevo León () is a state State may refer to: Arts, entertainment, and media Literature * ''State Magazine'', a monthly magazine published by the U.S. Department of State * The State (newspaper), ''The State'' (newspaper), a daily newspaper in ...
, Mexico ** '' C. texana'', origin:
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2020, ...

Texas
, USA * '' C. radicans'' – calabacilla, calabaza de coyote; origin: Central Mexico ** '' C. gracilior'' The taxonomy by Nee closely matches the species groupings reported in a pair of studies by a botanical team led by Rhodes and Bemis in 1968 and 1970 based on statistical groupings of several phenotype, phenotypic traits of 21 species. Seeds for studying additional species members were not available. Sixteen of the 21 species were grouped into five clusters with the remaining five being classified separately: * ''C. digitata'', ''C. palmata'', ''C. californica'', ''C. cylindrata'', ''C. cordata'' * ''C. martinezii'', ''C. okeechobeensis'', ''C. lundelliana'' * ''C. sororia'', ''C. gracilior'', ''C. palmeri''; ''C. argyrosperma'' (reported as ''C. mixta'') was considered close to the three previous species * ''C. maxima'', ''C. andreana'' * ''C. pepo'', ''C. texana'' * ''C. moschata'', ''C. ficifolia'', ''C. pedatifolia'', ''C. foetidissima'', and ''C. ecuadorensis'' were placed in their own separate species groups as they were not considered significantly close to any of the other species studied.


Phylogeny

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


Reproductive biology

All species of ''Cucurbita'' have 20 pairs of chromosomes. Many North and Central American species are visited by specialist
pollinator A pollinator is an animal that moves pollen File:Pollen Tube.svg, Pollen Tube Diagram Pollen is a powdery substance consisting of pollen grains which are Sporophyte, microsporophytes of spermatophyta, seed plants, which produce male gametes ...

pollinator
s in the Apidae, apid tribe Eucerini, especially the genera ''Peponapis'' and ''Xenoglossa'', and these squash bees can be crucial to the flowers producing fruit after pollination. 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 (plants), 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. Parthenocarpy, Seedlessness is known to occur in certain cultivars of ''C. pepo''. 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. The plant hormones ethylene and auxin are key in fruit set and development. 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. 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. 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. Gibberellin is also involved in other developmental processes of plants such as seed and stem growth.


Germination and seedling growth

Seeds with maximum germination potential develop (in ''C. moschata'') by 45 days after anthesis, and seed weight reaches its maximum 70 days after anthesis. Some varieties of ''C. pepo'' germinate best with eight hours of sunlight daily and a planting depth of . Seeds planted deeper than are not likely to germinate. 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. 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. 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 Phytic acid, phytate forms in seed tissues as spherical crystalline intrusions in protein bodies called Globoid (botany), globoids. Along with other nutrients, phytate is used completely during seedling growth. Heavy metal (chemistry), Heavy metal contamination, including cadmium, has a significant negative impact on plant growth. ''Cucurbita'' plants grown in the spring tend to grow larger than those grown in the autumn.


Distribution and habitat

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 Mesoamerica is a historical and important region In geography, regions are areas that are broadly divided by physical characteristics ( physical geography), human impact characteristics ( human geography), and the interaction of humanity and th ...
; for the fifth, ''C. maxima'', these events occurred in South America. 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. ''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#Organic oils, oil and Protein (nutrient), 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 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 to 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#Effect on plants, frost, and they prefer both bright sunlight and soil with a pH of 6.0 to 7.0. ''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'', ''zapallito'', ''zipinka'', Banana, Delicious, Hubbard, Marrow (''C. maxima'' Marrow), Show, and Turban. ''C. moschata'' is native to Latin America, but the precise location of origin is uncertain. 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 wide. It generally grows at low altitudes in hot climates with heavy rainfall, but some varieties have been found above . 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 ) , population_note = , population_rank = 10th , timezone1 = CST CST or Cst may refer to: Time zones * Central Standard Time, North America's Central Time Zone: UTC−06:00 * China Standard Time: UTC+08:00 * Cuba ...

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. Debates about the origin of ''C. pepo'' have been on-going since at least 1857. 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 organism, 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. ''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 . Leaves have 3–5 lobes and are wide. All the subspecies, varieties, and cultivars are Hybrid (biology), 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. There is one non-edible cultivated variety: ''C. pepo'' var. ''ovifera''.


History and domestication

The ancestral species of the genus ''Cucurbita'' were present in the Americas before the Prehistoric migration and settlement of the Americas from Asia, arrival of humans, 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. 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. 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. Species native to North America include '' C. digitata'' (calabazilla), and ''C. foetidissima'' (buffalo gourd), '' 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 ''Lagenaria'' is a genus of gourd-bearing vines in the squash family (Cucurbitaceae). ''Lagenaria'' contains six species, all of which are indigenous to tropical Africa.tribe The term tribe is used in many different contexts to refer to a category of human Humans (''Homo sapiens'') are the most populous and widespread species of primates, characterized by bipedality, opposable thumbs, hairlessness, and intellig ...
. 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. 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. Solid evidence of domesticated ''C. pepo'' was found in the Guilá Naquitz cave in the form of increasing Peel (fruit), rind thickness and larger Peduncle (botany), peduncles in the newer Stratigraphy (archaeology), stratification layers of the cave. By c. 8,000 years Before Present, BP the ''C. pepo'' peduncles found are consistently more than 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. During the same time frame, average rind thickness increased from to . Recent genomic studies suggest that ''
Cucurbita argyrosperma ''Cucurbita argyrosperma'', also the Japanese pie pumpkin or cushaw pumpkin, and silver-seed gourd, is a species of winter squash originally from the south of Mexico. This annual plant, annual herbaceous plant is cultivated in the Americas for its ...
'' was domesticated in Mexico, in the region that is currently known as the state of Jalisco. Squash was domesticated first, followed by maize and then beans, becoming part of the Three Sisters (agriculture), Three Sisters agricultural system of companion planting. The English language, English word "squash" derives from ''askutasquash'' (a green thing eaten raw), a word from the Massachusett language, Narragansett language, which was documented by Roger Williams (theologian), Roger Williams, the founder of Rhode Island, in his 1643 publication ''A Key Into the Language of America''. Similar words for squash exist in related languages of the Algonquian languages, Algonquian Language family, family.


Production

The family Cucurbitaceae has many species used as human food. ''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, the fruits and pepita, 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. 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, or otherwise prepared. Squashes are primarily grown for the fresh food market. The Food and Agriculture Organization, 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 Tonne, metric tons of squashes produced are: The 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.


Nutrients

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 Vitamin C (also known as ascorbic acid and ascorbate) is a vitamin found in various foods and sold as a dietary supplement. It is used to prevent and treat scurvy. Vitamin C is an Nutrient#Essential nutrients, essential nutrient involved in t ...

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. Pumpkin seeds contain vitamin E, Protein (nutrient)#Testing in foods, crude protein, B vitamins and several dietary minerals (see nutrition table at pepita). Also present in pumpkin seeds are unsaturated fatty acid, unsaturated and saturated fatty acid, saturated oils, palmitic acid, palmitic, oleic acid, oleic and linoleic acid, linoleic fatty acids, as well as carotenoids.


Toxins

Cucurbitin is an amino acid and a carboxypyrrolidine that is found in raw ''Cucurbita'' seeds. It retards the development of parasitic Trematoda, flukes when administered to infected host mice, although the effect is only seen if administration begins immediately after infection. Cucurmosin is a ribosome inactivating protein found in the flesh and seed of ''Cucurbita'', notably ''Cucurbita moschata''. Cucurmosin is more toxic to cancer cells than healthy cells. Cucurbitacin is a steroid, plant steroid present in wild ''Cucurbita'' and in each member of the family ''Cucurbitaceae''. Poisonous to mammals, 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. Ingesting too much cucurbitacin can cause stomach cramps, diarrhea and even collapse. 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. 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''). ''Cucurbita'' can be susceptible to the pest ''Bemisia argentifolii'' (silverleaf whitefly) 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''). The squash bug causes major damage to plants because of its very toxic saliva. The red pumpkin beetle (''Raphidopalpa foveicollis'') is a serious pest of cucurbits, especially the pumpkin, which it can defoliate. Cucurbitaceae, Cucurbits are susceptible to diseases such as bacterial wilt (''Erwinia tracheiphila''), anthracnose (''Colletotrichum'' spp.), fusarium wilt (''Fusarium'' spp.), phytophthora blight (''Phytophthora'' spp. Oomycete, water molds), and powdery mildew (''Erysiphe'' spp.). Plant defense against herbivory, 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), watermelon mosaic virus (WMV), and zucchini yellow mosaic virus (ZYMV). PRSV is the only one of these viruses that does not affect all cucurbits. SqMV and CMV are the most common viruses among cucurbits. 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 (Plymouth Colony), Pilgrims, even featuring at the first Thanksgiving (United States), Thanksgiving. Commercially produced pumpkin commonly used in
pumpkin pie Pumpkin pie is a dessert Dessert () is a course (food), course that concludes a meal. The course consists of sweet foods, such as confections, and possibly a beverage such as dessert wine and liqueur. In some parts of the world, such as much o ...

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. Other foods that can be made using members of this genus include biscuits, Pumpkin bread, bread, cheesecake, desserts, donuts, granola, ice cream, lasagna dishes, pancakes, pudding, pumpkin butter, salads, soups, and stuffing. Squash soup is a dish in African cuisine. The Xerophyte, xerophytic species are proving useful in the search for nutritious foods that grow well in arid regions. ''C. ficifolia'' is used to make soft and mildly alcoholic drinks. In India, squashes (''ghia'') are cooked with seafood such as prawns. 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, 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. 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.


Art, music, and literature

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. For example, cucurbits are often represented in Moche (culture), Moche ceramics. 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. 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. 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. 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, time. It became a popular hit in Australia in 1973, and was revived by the Wurzels in Britain on their 2003 album ''Cutler of the West''. John Greenleaf Whittier wrote a poem entitled ''The Pumpkin'' in 1850. "The Great Pumpkin" is a fictional holiday figure in the comic strip ''Peanuts'' by Charles M. Schulz.


Cleansing and personal care 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. Pumpkin is also used in cosmetics.


Folk remedies

''Cucurbita'' have been used in various cultures as folk remedies. Pumpkins have been used by Indigenous peoples of the Americas, 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. In southeastern Europe, seeds of ''C. pepo'' were used to treat irritable bladder and benign prostatic hyperplasia. 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. 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. In China, Cucurbita moschata, ''C. moschata'' seeds were also used in traditional Chinese medicine for the treatment of the parasitic disease schistosomiasis and for the expulsion of tape worms. In Mexico, herbalists use ''C. ficifolia'' in the belief that it reduces blood sugar levels.


Festivals

''Cucurbita'' fruits including pumpkins and marrows are celebrated in festivals in countries such as Argentina, Austria, Bolivia, Britain, Canada, Croatia, France, Germany, India, Italy, Japan, Peru, Portugal, Spain, Switzerland, and the United States. Argentina holds an annual nationwide pumpkin festival ''Fiesta Nacional del Zapallo'' ("Squashes and Pumpkins National Festival"), in Ceres, Santa Fe, on the last day of which a ''Reina Nacional del Zapallo'' ("National Queen of the Pumpkin") is chosen. In Portugal the ''Festival da Abóbora de Lourinhã e Atalaia'' ("Squashes and Pumpkins Festival in Lourinhã and Gavião e Atalaia, Atalaia") is held in Lourinhã city, called the ''Capital Nacional da Abóbora'' (the "National Capital of Squashes and Pumpkins"). Ludwigsburg, Ludwigsburg, Germany annually hosts the world's largest pumpkin festival. In Britain a giant marrow (zucchini) weighing was displayed in the Harrogate Autumn Flower Show in 2012. In the US, pumpkin chucking is practiced competitively, with machines such as trebuchets and Seismic source#Air gun, air cannons designed to throw intact pumpkins as far as possible. 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. Halloween is widely celebrated with jack-o-lanterns made of large orange pumpkins carved with ghoulish faces and illuminated from inside with candles. The pumpkins used for jack-o-lanterns are ''C. pepo'', 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.


See also

* List of gourds and squashes in the genus ''Cucurbita'' * List of squash and pumpkin dishes


Notes


References


External links

* * * {{Authority control Cucurbita, Cucurbitaceae genera Squashes and pumpkins Early agriculture in Mesoamerica Crops originating from indigenous Americans Native American cuisine