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The Cucurbitales are an order of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s, included in the
rosid The rosids are members of a large clade ( monophyletic group) of flowering plants, containing about 70,000 species, more than a quarter of all angiosperms. The clade is divided into 16 to 20 orders, depending upon circumscription and classific ...
group of
dicotyledon The dicotyledons, also known as dicots (or, more rarely, dicotyls), are one of the two groups into which all the flowering plants (angiosperms) were formerly divided. The name refers to one of the typical characteristics of the group: namely, t ...
s. This order mostly belongs to tropical areas, with limited presence in subtropical and temperate regions. The order includes shrubs and trees, together with many herbs and climbers. One major characteristic of the Cucurbitales is the presence of unisexual flowers, mostly
pentacyclic A cyclic flower is a flower type formed out of a series of whorls; sets of identical organs attached around the axis at the same point. Most flowers consist of a single whorl of sepals termed a calyx; a single whorl of petals termed a corolla; one ...
, with thick pointed petals (whenever present). The pollination is usually performed by insects, but
wind pollination Anemophily or wind pollination is a form of pollination whereby pollen is distributed by wind. Almost all gymnosperms are anemophilous, as are many plants in the order Poales, including grasses, sedges, and rushes. Other common anemophilous plan ...
is also present (in
Coriariaceae ''Coriaria'' is the sole genus in the family Coriariaceae, which was described by Linnaeus in 1753. It includes 14 species of small trees, shrubs and subshrubs, with a widespread but disjunct distribution across warm temperate regions of the worl ...
and
Datiscaceae The Datiscaceae are a family of dicotyledonous plants, containing two species of the genus ''Datisca''. Two other genera, ''Octomeles'' and ''Tetrameles'', are now classified in the family Tetramelaceae. Datiscaceae are large herbaceous plants, ...
). The order consists of roughly 2600 species in eight families. The largest families are
Begoniaceae Begoniaceae is a family of flowering plants with two genera and about 2040 species occurring in the subtropics and tropics of both the New World and Old World. All but one of the species are in the genus ''Begonia''. There have been many recent d ...
(begonia family) with around 1500 species and
Cucurbitaceae The Cucurbitaceae, also called cucurbits or the gourd family, are a plant family consisting of about 965 species in around 95 genera, of which the most important to humans are: *''Cucurbita'' – squash, pumpkin, zucchini, some gourds *''Lagena ...
(gourd family) with around 900 species. These two families include the only economically important plants. Specifically, the Cucurbitaceae (gourd family) include some food species, such as
squash Squash may refer to: Sports * Squash (sport), the high-speed racquet sport also known as squash racquets * Squash (professional wrestling), an extremely one-sided match in professional wrestling * Squash tennis, a game similar to squash but pla ...
,
pumpkin A pumpkin is a vernacular term for mature winter squash of species and varieties in the genus ''Cucurbita'' that has culinary and cultural significance but no agreed upon botanical or scientific meaning. The term ''pumpkin'' is sometimes use ...
(both from ''
Cucurbita ''Cucurbita'' (Latin for gourd) 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 ...
''),
watermelon Watermelon (''Citrullus lanatus'') is a flowering plant species of the Cucurbitaceae family and the name of its edible fruit. A scrambling and trailing vine-like plant, it is a highly cultivated fruit worldwide, with more than 1,000 varieti ...
(''Citrullus vulgaris''), and
cucumber Cucumber (''Cucumis sativus'') is a widely-cultivated Vine#Horticultural climbing plants, creeping vine plant in the Cucurbitaceae family that bears usually cylindrical Fruit, fruits, which are used as culinary vegetables.
and
melon A melon is any of various plants of the family Cucurbitaceae with sweet, edible, and fleshy fruit. The word "melon" can refer to either the plant or specifically to the fruit. Botanically, a melon is a kind of berry, specifically a "pepo". The ...
s (''
Cucumis __NOTOC__ ''Cucumis'' is a genus of twining, tendril-bearing plants in the family Cucurbitaceae which includes the cucumber (''Cucumis sativus''), muskmelons (''Cucumis melo'', including cantaloupe and honeydew), the horned melon (''Cucumis m ...
''). The Begoniaceae are known for their horticultural species, of which there are over 130 with many more
varieties Variety may refer to: Arts and entertainment Entertainment formats * Variety (radio) * Variety show, in theater and television Films * ''Variety'' (1925 film), a German silent film directed by Ewald Andre Dupont * ''Variety'' (1935 film), ...
.


Overview

The Cucurbitales are an order of plants with a cosmopolitan distribution, particularly diverse in the
tropics The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
. Most are herbs, climber herbs, woody lianas or shrubs but some genera include canopy-forming evergreen
lauroid This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...
trees. Members of the Cucurbitales form an important component of low to montane tropical forest with greater representation in terms of the number of species. Although not known with certainty the total number of species in the order, conservative estimates indicate about 2600 species worldwide, distributed in 109 genera. Compared to other flowering plant orders, the taxonomy is poorly understood due to their great diversity, difficulty in identification, and limited study. The order Cucurbitales in the eurosid I clade comprises almost 2600 species in 109 or 110 genera in eight families, tropical and temperate, of very different sizes, morphology, and ecology. It is a case of
divergent evolution Divergent evolution or divergent selection is the accumulation of differences between closely related populations within a species, leading to speciation. Divergent evolution is typically exhibited when two populations become separated by a geog ...
. In contrast, there is
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
with other groups not related due to ecological or physical drivers toward a similar solution, including
analogous structures Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
. Some species are trees that have similar foliage to the true laurels due to convergent evolution. The patterns of speciation in the Cucurbitales are diversified in a high number of species. They have a pantropical distribution with centers of diversity in Africa, South America, and Southeast Asia. They most likely originated in West
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
67–107 million years ago, so the oldest split could relate to the break-up of Gondwana in the middle Eocene to late Oligocene, 45–24 million years ago. The group reached their current distribution by multiple intercontinental dispersal events. One factor was product of aridification, other groups responded to favorable climatic periods and expanded across the available habitat, occurring as opportunistic species across wide distribution; other groups diverged over long periods within isolated areas. The Cucurbitales comprise the families:
Apodanthaceae The family Apodanthaceae comprises about 10 species of endoparasitic herbs. They live in the branches or stems of their hosts (as filaments similar to a fungal mycelium), emerging only to flower and fruit. The plants produce no green parts and do ...
,
Anisophylleaceae The Anisophylleaceae are a small family with four genera and about 70 species, in the order Cucurbitales, according to the Angiosperm Phylogeny Group, APG II. However, it is more isolated from the other suprafamilial clades in this order, while i ...
,
Begoniaceae Begoniaceae is a family of flowering plants with two genera and about 2040 species occurring in the subtropics and tropics of both the New World and Old World. All but one of the species are in the genus ''Begonia''. There have been many recent d ...
,
Coriariaceae ''Coriaria'' is the sole genus in the family Coriariaceae, which was described by Linnaeus in 1753. It includes 14 species of small trees, shrubs and subshrubs, with a widespread but disjunct distribution across warm temperate regions of the worl ...
, Corynocarpaceae,
Cucurbitaceae The Cucurbitaceae, also called cucurbits or the gourd family, are a plant family consisting of about 965 species in around 95 genera, of which the most important to humans are: *''Cucurbita'' – squash, pumpkin, zucchini, some gourds *''Lagena ...
,
Tetramelaceae The Tetramelaceae are a family of plants formerly classed in the Datiscaceae family . It contains two genera, '' Octomeles'' and ''Tetrameles ''Tetrameles'' is a genus of flowering plants in the family Tetramelaceae with one species, ''Tetram ...
, and
Datiscaceae The Datiscaceae are a family of dicotyledonous plants, containing two species of the genus ''Datisca''. Two other genera, ''Octomeles'' and ''Tetrameles'', are now classified in the family Tetramelaceae. Datiscaceae are large herbaceous plants, ...
. Some of the synapomorphies of the order are: leaves in spiral with secondary veins palmated, calyx or perianth valvate, and the elevated stomatal calyx/perianth bearing separate styles. The two whorls are similar in texture. ''
Tetrameles nudiflora ''Tetrameles'' is a genus of flowering plants in the family Tetramelaceae with one species, ''Tetrameles nudiflora''. It grows as a large deciduous tree and is found across southern Asia from India through southeast Asia, Malesia, and into northe ...
'' is a tree of immense proportions of height and width;
Tetramelaceae The Tetramelaceae are a family of plants formerly classed in the Datiscaceae family . It contains two genera, '' Octomeles'' and ''Tetrameles ''Tetrameles'' is a genus of flowering plants in the family Tetramelaceae with one species, ''Tetram ...
,
Anisophylleaceae The Anisophylleaceae are a small family with four genera and about 70 species, in the order Cucurbitales, according to the Angiosperm Phylogeny Group, APG II. However, it is more isolated from the other suprafamilial clades in this order, while i ...
, and Corynocarpaceae are tall canopy trees in temperate and tropical forests. The genus ''
Dendrosicyos ''Dendrosicyos'' is a monotypic genus in the plant family (biology), family Cucurbitaceae. The only species is ''Dendrosicyos socotranus'', the cucumber tree. The species is endemism, endemic to the island of Socotra in Yemen, and is the only spe ...
'', with the only species being the cucumber tree, is adapted to the arid semidesert island of
Socotra Socotra or Soqotra (; ar, سُقُطْرَىٰ ; so, Suqadara) is an island of the Republic of Yemen in the Indian Ocean, under the ''de facto'' control of the UAE-backed Southern Transitional Council, a secessionist participant in Yemen’s ...
. Deciduous perennial Cucurbitales lose all of their
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
for part of the year depending on variations in rainfall. The leaf loss coincides with the
dry season The dry season is a yearly period of low rainfall, especially in the tropics. The weather in the tropics is dominated by the tropical rain belt, which moves from the northern to the southern tropics and back over the course of the year. The te ...
in tropical, subtropical and arid regions. In
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
or
polar climate The polar climate regions are characterized by a lack of warm summers but with varying winters. Every month in a polar climate has an average temperature of less than . Regions with polar climate cover more than 20% of the Earth's area. Most of ...
s, the dry season is due to the inability of the plant to absorb water available in the form of ice.
Apodanthaceae The family Apodanthaceae comprises about 10 species of endoparasitic herbs. They live in the branches or stems of their hosts (as filaments similar to a fungal mycelium), emerging only to flower and fruit. The plants produce no green parts and do ...
are obligatory endoparasites that only emerge once a year in the form of small flowers that develop into small berries, however taxonomists have not agreed on the exact placement of this family within the Cucurbitales. Over half of the known members of this order belong to the greatly diverse begonia family Begoniaceae, with around 1500 species in two genera. Before modern DNA-molecular classifications, some Cucurbitales species were assigned to orders as diverse as
Ranunculales Ranunculales is an order of flowering plants. Of necessity it contains the family Ranunculaceae, the buttercup family, because the name of the order is based on the name of a genus in that family. Ranunculales belongs to a paraphyletic group kno ...
,
Malpighiales The Malpighiales comprise one of the largest orders of flowering plants, containing about 36 families and more than species, about 7.8% of the eudicots. The order is very diverse, containing plants as different as the willow, violet, poinsett ...
,
Violales Violales is a botanical name of an order of flowering plants and takes its name from the included family Violaceae; it was proposed by Lindley (1853). The name has been used in several systems, although some systems used the name Parietales f ...
, and
Rafflesiales Rafflesiales is a botanical name of an order of flowering plants. The name was first published by Oliver in 1895. The Cronquist system used this name for an order placed in subclass Rosidae with the following circumscription (1981) : * order Raf ...
. Early molecular studies revealed several surprises, such as the nonmonophyly of the traditional
Datiscaceae The Datiscaceae are a family of dicotyledonous plants, containing two species of the genus ''Datisca''. Two other genera, ''Octomeles'' and ''Tetrameles'', are now classified in the family Tetramelaceae. Datiscaceae are large herbaceous plants, ...
, including ''
Tetrameles ''Tetrameles'' is a genus of flowering plants in the family Tetramelaceae with one species, ''Tetrameles nudiflora''. It grows as a large deciduous tree and is found across southern Asia from India through southeast Asia, Malesia, and into nort ...
'' and ''
Octomeles ''Octomeles'' is a monotypic genus of plant in family Tetramelaceae. The sole species is ''Octomeles sumatrana'', sometimes written ''O. sumatranum''. ''Octomeles sumatrana'', commonly called Benuang, or Ilimo, is found in Brunei, Indonesia, Ma ...
'', but the exact relationships among the families remain unclear. The lack of knowledge about the order in general is due to many species being found in countries with limited economic means or unstable political environments, factors unsuitable for plant collection and detailed study. Thus the vast majority of species remain poorly determined, and a future increase in the number of species is expected.


Classification

Under the
Cronquist system The Cronquist system is a taxonomic classification system of flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of Flowering Plants'' (1968; 2nd edition, 1988) a ...
, the families
Begoniaceae Begoniaceae is a family of flowering plants with two genera and about 2040 species occurring in the subtropics and tropics of both the New World and Old World. All but one of the species are in the genus ''Begonia''. There have been many recent d ...
,
Cucurbitaceae The Cucurbitaceae, also called cucurbits or the gourd family, are a plant family consisting of about 965 species in around 95 genera, of which the most important to humans are: *''Cucurbita'' – squash, pumpkin, zucchini, some gourds *''Lagena ...
, and
Datiscaceae The Datiscaceae are a family of dicotyledonous plants, containing two species of the genus ''Datisca''. Two other genera, ''Octomeles'' and ''Tetrameles'', are now classified in the family Tetramelaceae. Datiscaceae are large herbaceous plants, ...
were placed in the order
Violales Violales is a botanical name of an order of flowering plants and takes its name from the included family Violaceae; it was proposed by Lindley (1853). The name has been used in several systems, although some systems used the name Parietales f ...
, within the subclass Dilleniidae, with the Tetramelaceae subsumed into the Datiscaceae. Corynocarpaceae was placed in order
Celastrales The Celastrales are an order of flowering plants found throughout the tropics and subtropics, with only a few species extending far into the temperate regions. The 1200"Lepidobotryaceae", "Parnassiaceae", and "Celastraceae" In: Klaus Kubitzki (ed ...
, and
Anisophylleaceae The Anisophylleaceae are a small family with four genera and about 70 species, in the order Cucurbitales, according to the Angiosperm Phylogeny Group, APG II. However, it is more isolated from the other suprafamilial clades in this order, while i ...
in order
Rosales Rosales () is an order of flowering plants. Peter F. Stevens (2001 onwards). "Rosales". At: Trees At: Angiosperm Phylogeny Website. At: Missouri Botanical Garden Website. (see ''External links'' below) It is sister to a clade consisting of Faga ...
, both under subclass Rosidae.
Coriariaceae ''Coriaria'' is the sole genus in the family Coriariaceae, which was described by Linnaeus in 1753. It includes 14 species of small trees, shrubs and subshrubs, with a widespread but disjunct distribution across warm temperate regions of the worl ...
was placed in
Ranunculaceae Ranunculaceae (buttercup or crowfoot family; Latin "little frog", from "frog") is a family of over 2,000 known species of flowering plants in 43 genera, distributed worldwide. The largest genera are ''Ranunculus'' (600 species), ''Delphinium' ...
, subclass Magnoliidae.
Apodanthaceae The family Apodanthaceae comprises about 10 species of endoparasitic herbs. They live in the branches or stems of their hosts (as filaments similar to a fungal mycelium), emerging only to flower and fruit. The plants produce no green parts and do ...
was not recognised as a family, its genera being assigned to another parasitic plant family, the
Rafflesiaceae The Rafflesiaceae are a family of rare parasitic plants comprising 36 species in 3 genera found in the tropical forests of east and southeast Asia, including ''Rafflesia arnoldii'', which has the largest flowers of all plants. The plants are end ...
. The present classification is due to APG III (2009).


Systematics

Modern
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
suggest the following relationships:


References


Further reading

* *


External links

{{Taxonbar, from=Q21875 Extant Albian first appearances Angiosperm orders