Cucurbitales Genera
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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 (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
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 classificatio ...
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, ...
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; o ...
, 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 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 ...
(begonia family) with around 1500 species and
Cucurbitaceae The Cucurbitaceae (), also called cucurbits or the gourd family, are a plant family (biology), family consisting of about 965 species in 101 genera.
(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 most often refers to: * Squash (sport), the high-speed racquet sport also known as squash racquets * Squash (plant), the fruit of vines of the genus ''Cucurbita'' Squash may also refer to: Sports * Squash (professional wrestling), an extr ...
,
pumpkin A pumpkin is a cultivar, cultivated winter squash in the genus ''Cucurbita''. The term is most commonly applied to round, orange-colored squash varieties, but does not possess a scientific definition. It may be used in reference to many dif ...
(both from ''
Cucurbita 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 know ...
''),
watermelon The watermelon (''Citrullus lanatus'') is a species of flowering plant in the family Cucurbitaceae, that has a large, edible fruit. It is a Glossary of botanical terms#scandent, scrambling and trailing vine-like plant, and is plant breeding ...
(''Citrullus vulgaris''), and
cucumber The cucumber (''Cucumis sativus'') is a widely-cultivated creeping vine plant in the family Cucurbitaceae that bears cylindrical to spherical fruits, which are used as culinary vegetables.melon A melon is any of various plants of the family Cucurbitaceae with sweet, edible, and fleshy fruit. It can also specifically refer to ''Cucumis melo'', commonly known as the "true melon" or simply "melon". The term "melon" can apply to both the p ...
s (''
Cucumis __NOTOC__ ''Cucumis'' is a genus of vine, twining, tendril-bearing plants in the Family (biology), family Cucurbitaceae which includes the cucumber (''Cucumis sativus''), true melons (''Cucumis melo'', including cantaloupe and Honeydew (melon), ...
''). 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, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
. 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 ...
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, sometimes leading to speciation. Divergent evolution is typically exhibited when two populations become separate ...
. 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 comm ...
with other groups not related due to ecological or physical drivers toward a similar solution, including
analogous structures Analogy is a comparison or correspondence between two things (or two groups of things) because of a third element that they are considered to share. In logic, it is an inference or an argument from one particular to another particular, as oppose ...
. 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, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
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 d ...
,
Anisophylleaceae The Anisophylleaceae are a small family with four genera and about 70 species, in the order Cucurbitales, according to the APG II. However, it is more isolated from the other suprafamilial clades in this order, while it shows some similarities 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 ...
, Coriariaceae, Corynocarpaceae,
Cucurbitaceae The Cucurbitaceae (), also called cucurbits or the gourd family, are a plant family (biology), family consisting of about 965 species in 101 genera.
,
Tetramelaceae The Tetramelaceae are a family of plants in the order Cucurbitales, containing two genera of mostly Asian, large trees, ''Octomeles'' and ''Tetrameles'', each with a single species. These genera were formerly placed in the Datiscaceae,van Steeni ...
, 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 in the order Cucurbitales, containing two genera of mostly Asian, large trees, ''Octomeles'' and ''Tetrameles'', each with a single species. These genera were formerly placed in the Datiscaceae,van Steeni ...
,
Anisophylleaceae The Anisophylleaceae are a small family with four genera and about 70 species, in the order Cucurbitales, according to the APG II. However, it is more isolated from the other suprafamilial clades in this order, while it shows some similarities i ...
, and Corynocarpaceae are tall canopy trees in temperate and tropical forests. The genus '' Dendrosicyos'', with the only species being the cucumber tree, is adapted to the arid semidesert island of
Socotra Socotra, locally known as Saqatri, is a Yemeni island in the Indian Ocean. Situated between the Guardafui Channel and the Arabian Sea, it lies near major shipping routes. Socotra is the largest of the six islands in the Socotra archipelago as ...
. Deciduous perennial Cucurbitales lose all of their
leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
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 t ...
in tropical, subtropical and arid regions. In
temperate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
or
polar climate The polar climate regions are characterized by a lack of warm summers but with varying winters. Every month a polar climate has an average temperature of less than . Regions with a 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 d ...
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 k ...
,
Malpighiales The Malpighiales comprise one of the largest Order (biology), orders of flowering plants. The order is very diverse, with well-known members including willows, Viola (plant), violets, aspens and Populus, poplars, Euphorbia pulcherrima, 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 for ...
, 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 Ra ...
. 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 northe ...
'' 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 list of systems of plant taxonomy, taxonomic classification system of angiosperms, flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of ...
, 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 ...
,
Cucurbitaceae The Cucurbitaceae (), also called cucurbits or the gourd family, are a plant family (biology), family consisting of about 965 species in 101 genera.
, 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 for ...
, within the subclass Dilleniidae, with the Tetramelaceae subsumed into the Datiscaceae. Corynocarpaceae was placed in order
Celastrales The Celastrales are an order (biology), order of flowering plants found throughout the tropics and subtropics, with only a few species extending far into the Temperateness, temperate regions. The 1200"Lepidobotryaceae", "Parnassiaceae", and "Celas ...
, and
Anisophylleaceae The Anisophylleaceae are a small family with four genera and about 70 species, in the order Cucurbitales, according to the APG II. However, it is more isolated from the other suprafamilial clades in this order, while it shows some similarities i ...
in order
Rosales Rosales (, ) are 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) Well-known members of Rosales include: ...
, both under subclass Rosidae. Coriariaceae was placed in
Ranunculaceae Ranunculaceae (, buttercup or crowfoot family; Latin "little frog", from "frog") is a family (biology), family of over 2,000 known species of flowering plants in 43 genera, distributed worldwide. The largest genera are ''Ranunculus'' (600 spec ...
, 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 d ...
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 e ...
. 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

{{Authority control Extant Albian first appearances Angiosperm orders