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Malvaceae (), or the mallows, is a family 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 estimated to contain 244 genera with 4225 known species. Well-known members of economic importance include cacao,
cola Cola is a Carbonation, carbonated soft drink flavored with vanilla, cinnamon, citrus essential oil, oils, and other flavorings. Cola became popular worldwide after the American pharmacist John Stith Pemberton invented Coca-Cola, a trademarked br ...
,
cotton Cotton (), first recorded in ancient India, is a soft, fluffy staple fiber that grows in a boll, or protective case, around the seeds of the cotton plants of the genus '' Gossypium'' in the mallow family Malvaceae. The fiber is almost pure ...
,
okra Okra (, ), ''Abelmoschus esculentus'', known in some English-speaking countries as lady's fingers, is a flowering plant in the Malvaceae, mallow family native to East Africa. Cultivated in tropical, subtropical, and warm temperate regions aro ...
, roselle and
durian The durian () is the edible fruit of several tree species belonging to the genus ''Durio''. There are 30 recognized species, at least nine of which produce edible fruit. ''Durio zibethinus'', native to Borneo and Sumatra, is the only species ...
. There are also some genera containing familiar ornamentals, such as '' Alcea'' (hollyhock), '' Malva'' (mallow), and ''
Tilia ''Tilia'' is a genus of about 30 species of trees or bushes, native throughout most of the temperateness, temperate Northern Hemisphere. The tree is known as linden for the European species, and basswood for North American species. In Great Bri ...
'' (lime or linden tree). The genera with the largest numbers of species include ''
Hibiscus ''Hibiscus'' is a genus of flowering plants in the Malva, mallow family, Malvaceae. The genus is quite large, comprising List of Hibiscus species, several hundred species that are Native plant, native to warm temperate, Subtropics, subtropical ...
'' (434 species), '' Pavonia'' (291 species), '' Sida'' (275 species), '' Ayenia'' (216 species), ''
Dombeya ''Dombeya'' is a flowering plant genus. Traditionally included in the family Sterculiaceae, it is included in the expanded Malvaceae in the APG and most subsequent systematics. These plants are known by a number of vernacular names which sometim ...
'' (197 species), and '' Sterculia'' (181 species).


Taxonomy and nomenclature

The circumscription of the Malvaceae is controversial. The traditional Malvaceae ''
sensu stricto ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...
'' comprise a very homogeneous and cladistically
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
group. Another major circumscription, Malvaceae ''
sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...
'', has been more recently defined on the basis that genetics studies have shown the commonly recognised families
Bombacaceae Bombacaceae were long recognised as a family of flowering plants or Angiospermae. The family name was based on the type genus '' Bombax''. As is true for many botanical names, circumscription and status of the taxon has varied with taxonomic poi ...
,
Tiliaceae Tiliaceae () is a family of flowering plants. It is not a part of the APG, APG II and APG III classifications, being sunk in Malvaceae mostly as the subfamilies Tilioideae, Brownlowioideae and Grewioideae, but has an extensive historical re ...
, and
Sterculiaceae Sterculiaceae was a family of flowering plant based on the genus '' Sterculia''. Genera formerly included in Sterculiaceae are now placed in the family Malvaceae, in the subfamilies: Byttnerioideae, Dombeyoideae, Helicteroideae and Sterculioide ...
, which have always been considered closely allied to Malvaceae ''s.s.'', are not monophyletic groups. The Malvaceae can be expanded to include all of these families so as to compose a monophyletic group. Adopting this circumscription, the Malvaceae incorporate a much larger number of genera.


Subfamilies

This article is based on the second circumscription, as presented by the
Angiosperm Phylogeny Website The Angiosperm Phylogeny Website (or APweb) is a website that presents up-to-date research on the phylogeny and taxonomy of flowering plants (angiosperms) in what is intended to be a user-friendly way. The site is hosted by the Missouri Botanical ...
. The Malvaceae ''s.l.'' (hereafter simply "Malvaceae") comprise nine subfamilies. A tentative
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
of the family is shown below. The diamond denotes a poorly supported branching (<80%). Until recently, relationships between these subfamilies were either poorly supported or almost completely obscure. Continuing disagreements focused primarily on the correct circumscription of these subfamilies, including the preservation of the family Bombacaceae. A study published in 2021 presented a fully resolved phylogenetic framework for Malvaceae ''s.l.'' using genomic data for all nine subfamilies. Regarding the traditional Malvaceae ''s.s.'', the subfamily
Malvoideae Malvoideae is a botanical name at the rank of subfamily, which includes in the minimum the genus ''Malva''. It was first used by Burnett in 1835, but was not much used until recently, where, within the framework of the APG System, which unites th ...
approximately corresponds to that group.


Genera

245
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
are currently accepted.


Synapomorphies

The relationships between the "core Malvales" families used to be defined on the basis of shared "malvean affinities". These included the presence of malvoid teeth, stems with
mucilage Mucilage is a thick gluey substance produced by nearly all plants and some microorganisms. These microorganisms include protists which use it for their locomotion, with the direction of their movement always opposite to that of the secretion of ...
canals, and stratified wedge-shaped phloem. These affinities were problematic because they were not always shared within the core families. Later studies revealed more unambiguous synapomorphies within Malvaceae ''s.l..'' Synapomorphies identified within Malvaceae ''s.l.'' include the presence of tile cells, trichomatous nectaries, and an
inflorescence In botany, an inflorescence is a group or cluster of flowers arranged on a plant's Plant stem, stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a mai ...
structure called a bicolor unit. Tile cells consist of vertically positioned cells interspersed between and dimensionally similar to procumbent ray cells. Evidence of Malvean wood fossils has confirmed their evolutionary link in Malvaceae ''s.l.'', as well as explained their diverse structures. Flowers of Malvaceae ''s.l''. exhibit nectaries consisting of densely arranged multicellular hairs resembling trichomes. In most of Malvaceae ''s.l.'', these trichomatous nectaries are located on the inner surface of the sepals, but flowers of the subfamily Tiliodeae also have present nectaries on the petals. Malvean flowers also share a unifying structure known as a bicolor unit, named for its initial discovery in the flowers of ''Theobroma bicolor''. The bicolor unit consists of an ordered inflorescence with determinate cymose structures. The inflorescence can branch off the main axis, creating separate orders of the flowers, with the main axis developing first.
Bract In botany, a bract is a modified or specialized leaf, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also lo ...
s on the peduncle subtend axillary buds that become these lateral stalks. One bract within this whorl is a sterile bract. The bicolor unit is a variable structure in complexity, but the presence of fertile and sterile bracts is a salient characteristic.


Names

The English common name 'mallow' (also applied to other members of Malvaceae) comes from Latin '' malva'' (also the source for the English word "
mauve Mauve ( ; ) is a pale purple color named after the mallow flower (French: ). The first use of the word ''mauve'' as a color was in 1796–1798 according to the ''Oxford English Dictionary'', but its use seems to have been rare before 1859. ...
"). ''Malva'' itself was ultimately derived from the word for the plant in ancient Mediterranean languages.
Cognate In historical linguistics, cognates or lexical cognates are sets of words that have been inherited in direct descent from an etymological ancestor in a common parent language. Because language change can have radical effects on both the s ...
s of the word include
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 into the following periods: Mycenaean Greek (), Greek ...
() or (),
Modern Greek Modern Greek (, or , ), generally referred to by speakers simply as Greek (, ), refers collectively to the dialects of the Greek language spoken in the modern era, including the official standardized form of the language sometimes referred to ...
(), modern () and modern ().


Description

Most species are
herbaceous plant Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition o ...
s or
shrub A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...
s, but some are
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, e.g., including only woody plants with secondary growth, only ...
s or
liana A liana is a long-Plant stem, stemmed Woody plant, woody vine that is rooted in the soil at ground level and uses trees, as well as other means of vertical support, to climb up to the Canopy (biology), canopy in search of direct sunlight. T ...
s.


Leaves and stems

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, ...
are generally
alternate Alternative or alternate may refer to: Arts, entertainment and media * Alternative (''Kamen Rider''), a character in the Japanese TV series ''Kamen Rider Ryuki'' * Alternative comics, or independent comics are an alternative to mainstream superh ...
, often palmately lobed or compound and palmately veined. The margin may be entire, but when dentate, a vein ends at the tip of each tooth (malvoid teeth). Stipules are present. The
stem Stem or STEM most commonly refers to: * Plant stem, a structural axis of a vascular plant * Stem group * Science, technology, engineering, and mathematics Stem or STEM can also refer to: Language and writing * Word stem, part of a word respon ...
s contain mucous canals and often also mucous cavities. Hairs are common, and are most typically stellate. Stems of Bombacoideae are often covered in thick prickles.


Flowers

The flowers are commonly borne in definite or indefinite axillary
inflorescence In botany, an inflorescence is a group or cluster of flowers arranged on a plant's Plant stem, stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a mai ...
s, which are often reduced to a single flower, but may also be cauliflorous, oppositifolious, or terminal. They often bear supernumerary
bract In botany, a bract is a modified or specialized leaf, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also lo ...
s in the structure of a bicolor unit. They can be unisexual or bisexual, and are generally
actinomorphic Floral symmetry describes whether, and how, a flower, in particular its perianth, can be divided into two or more identical or mirror-image parts. Uncommonly, flowers may have no axis of symmetry at all, typically because their parts are spirall ...
, often associated with conspicuous bracts, forming an epicalyx. They generally have five valvate
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 Etymology The term ''sepalum'' ...
s, most frequently basally connate, with five imbricate
petal Petals are modified leaves that form an inner whorl surrounding the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corol ...
s. The
stamen The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...
s are five to numerous, and connate at least at their bases, but often forming a tube around the
pistil Gynoecium (; ; : gynoecia) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl (botany), whorl of a flower; it consists ...
s. The pistils are composed of two to many connate
carpel Gynoecium (; ; : gynoecia) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more ...
s. The
ovary The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/ oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are end ...
is superior, with axial placentation, with capitate or lobed stigma. The flowers have
nectaries Nectar is a viscous, sugar-rich liquid produced by plants in glands called nectaries, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists, ...
made of many tightly packed glandular
hairs Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and fin ...
, usually positioned on the sepals.


Fruits

The fruits are most often loculicidal capsules,
schizocarp A schizocarp is a dry fruit that, when mature, splits up into mericarps. There are different definitions: * Any Dry fruits, dry fruit composed of multiple carpels that separate. : Under this definition the mericarps can contain one or more ...
s or nuts.


Pollination

Self-pollination is often avoided by means of
protandry Sequential hermaphroditism (called dichogamy in botany) is one of the two types of hermaphroditism, the other type being simultaneous hermaphroditism. It occurs when the organism's sex changes at some point in its life. A sequential hermaphrodit ...
. Most species are entomophilous (pollinated by insects). Bees from the tribe
Emphorini The Emphorini are a tribe of apid bees. Genera *'' Alepidosceles'' *'' Diadasia'' *'' Diadasina'' **'' Diadasina'' (''Diadasina'') **'' Diadasina'' ('' Leptometriella'') *'' Meliphilopsis'' *'' Melitoma'' *'' Melitomella'' *'' Ptilothrix ...
of the
Apidae Apidae is the largest family within the superfamily Apoidea, containing at least 5700 species of bees. The family includes some of the most commonly seen bees, including bumblebees and honey bees, but also includes stingless bees (also used for ...
(including '' Ptilothrix'', '' Diadasia'', and '' Melitoma'') are known to specialize on the plants.


Importance

A number of species are pests in
agriculture Agriculture encompasses crop and livestock production, aquaculture, and forestry for food and non-food products. Agriculture was a key factor in the rise of sedentary human civilization, whereby farming of domesticated species created ...
, including ''
Abutilon theophrasti ''Abutilon theophrasti,'' also known as velvetleaf, velvet plant, velvetweed and the Chinese jute is an annual plant in the Family (biology), family Malvaceae that is native to South Asia, southern Asia and it serves as a type species of the genu ...
'' and '' Modiola caroliniana'', and others that are garden escapees.
Cotton Cotton (), first recorded in ancient India, is a soft, fluffy staple fiber that grows in a boll, or protective case, around the seeds of the cotton plants of the genus '' Gossypium'' in the mallow family Malvaceae. The fiber is almost pure ...
(four species of ''
Gossypium ''Gossypium'' () is a genus of flowering plants in the tribe Gossypieae of the Malva, mallow family, Malvaceae, from which cotton is harvested. It is native to tropical and subtropical regions of the Old World, Old and New Worlds. There are abo ...
''),
kenaf Kenaf tymology: Persian ''Hibiscus cannabinus'', is a plant in the family Malvaceae also called Deccan hemp and Java jute. ''Hibiscus cannabinus'' is in the genus ''Hibiscus'' and is native to Africa, though its exact origin is unknown. The n ...
(''Hibiscus cannabinus''), cacao (''Theobroma cacao''),
kola nut The kola nut ( Yoruba: ''obi'', Dagbani: ''guli'', Hausa: goro, Igbo: ''ọjị'', Sängö: ''gôro,'' Swahili: ''mukezu'') is the seed of certain species of plant of the genus ''Cola'', placed formerly in the cocoa family Sterculiaceae and ...
(''Cola spp.''), and
okra Okra (, ), ''Abelmoschus esculentus'', known in some English-speaking countries as lady's fingers, is a flowering plant in the Malvaceae, mallow family native to East Africa. Cultivated in tropical, subtropical, and warm temperate regions aro ...
(''Abelmoschus esculentus'') are important agricultural crops. The fruit and leaves of
baobab ''Adansonia'' is a genus of medium-to-large deciduous trees known as baobabs ( or ). The eight species of ''Adansonia'' are native to Africa, Australia, and Madagascar but have also been introduced to other regions of the world, including Barb ...
s are edible, as is the fruit of the
durian The durian () is the edible fruit of several tree species belonging to the genus ''Durio''. There are 30 recognized species, at least nine of which produce edible fruit. ''Durio zibethinus'', native to Borneo and Sumatra, is the only species ...
. A number of species, including '' Hibiscus syriacus'', ''
Hibiscus rosa-sinensis ''Hibiscus'' is a genus of flowering plants in the Malva, mallow family, Malvaceae. The genus is quite large, comprising List of Hibiscus species, several hundred species that are Native plant, native to warm temperate, Subtropics, subtropical ...
'' and '' Alcea rosea'' are garden plants.


See also

* '' Florissantia'', an extinct
Cenozoic The Cenozoic Era ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterized by the dominance of mammals, insects, birds and angiosperms (flowering plants). It is the latest of three g ...
genus in the subfamily Sterculioideae


References

* * * * Bayer, C. and K. Kubitzki 2003. Malvaceae, pp. 225–311. In K. Kubitzki (ed.), ''The Families and Genera of Vascular Plants'', vol. 5, Malvales, Capparales and non-betalain Caryophyllales. * * * Maas, P. J. M. and L. Y. Th. Westra. 2005. ''Neotropical Plant Families'' (3rd edition). * * (abstract onlin
here
). *


External links






Tree of Life.org: Core Malvales


— ''image gallery''.
GREIF Flora: Malvaceae of Mongolia
{{Authority control Malvales families