Malvaceae - Lavatera Punctata-5
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Malvaceae, or the mallows, is a family of flowering plants estimated to contain 244 genera with 4225 known species. Well-known members of economic importance include okra, cotton,
cacao Cacao is the seed from which cocoa and chocolate are made, from Spanish cacao, an adaptation of Nahuatl cacaua, the root form of cacahuatl ("bean of the cocoa-tree"). It may also refer to: Plants *''Theobroma cacao'', a tropical evergreen tree ** ...
and
durian The durian (, ) is the edible fruit of several tree species belonging to the genus ''Durio''. There are 30 recognised ''Durio'' species, at least nine of which produce edible fruit. ''Durio zibethinus'', native to Borneo and Sumatra, is the onl ...
. 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 Britain a ...
'' (lime or linden tree). The largest genera in terms of number of species include '' Hibiscus'' (300 species), '' Sterculia'' (250 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 ...
'' (250 species), '' Pavonia'' (200 species) and '' Sida'' (200 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 co ...
'' comprise a very homogeneous and cladistically
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
group. Another major circumscription, Malvaceae '' sensu lato'', 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 point ...
,
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 record o ...
, and Sterculiaceae, which have always been considered closely allied to Malvaceae ''s.s.'', are not monophyletic groups. Thus, 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 Malvaceae ''s.l.'' (hereafter simply "Malvaceae") comprise nine subfamilies. A tentative cladogram 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.


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 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 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. Bracts on the
peduncle Peduncle may refer to: *Peduncle (botany), a stalk supporting an inflorescence, which is the part of the shoot of seed plants where flowers are formed *Peduncle (anatomy), a stem, through which a mass of tissue is attached to a body **Peduncle (art ...
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"). ''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 in different languages that have been inherited in direct descent from an etymology, etymological ancestor in a proto-language, common parent language. Because language c ...
s of the word include Ancient Greek () or (), Modern Greek (), modern ar, ملوخية () and modern he, מלוחיה ().


Description

Most species are herbaceous plants or
shrub A shrub (often also called a 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 ...
s, but some are trees or
liana A liana is a long- stemmed, 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 in search of direct sunlight. The word ''liana'' does not refer to a ta ...
s.


Leaves and stems

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 ...
are generally alternate, 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 may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
s contain mucous canals and often also mucous cavities. Hairs are common, and are most typically
stellate Stellate, meaning star-shaped, may refer to: * Stellate cell * Stellate ganglion * Stellate reticulum * Stellate veins * Stellate trichomes (hairs) on plants * Stellate laceration or incision Wound#Open * Stellate fan-shaped Espalier (one form ...
. Stems of Bombacoideae are often covered in thick prickles.


Flowers

The flowers are commonly borne in definite or indefinite axillary inflorescences, 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, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...
s in the structure of a bicolor unit. They can be unisexual or bisexual, and are generally actinomorphic, often associated with conspicuous bracts, forming an
epicalyx 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 ...
. They generally have five valvate sepals, most frequently basally connate, with five imbricate
petal Petals are modified Leaf, leaves that surround the reproductive parts of flowers. They are often advertising coloration, brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''c ...
s. The
stamen 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 filame ...
s are five to numerous, and connate at least at their bases, but often forming a tube around the
pistil Gynoecium (; ) 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) ''pistils'' ...
s. The pistils are composed of two to many connate carpels. The
ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
is superior, with axial placentation, with capitate or lobed stigma. The flowers have nectaries 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 fine ...
, usually positioned on the sepals.


Fruits

The fruits are most often
loculicidal Dehiscence is the splitting of a mature plant structure along a built-in line of weakness to release its contents. This is common among fruits, anthers and sporangia. Sometimes this involves the complete detachment of a part; structures that o ...
capsules, schizocarps or
nut Nut often refers to: * Nut (fruit), fruit composed of a hard shell and a seed, or a collective noun for dry and edible fruits or seeds * Nut (hardware), fastener used with a bolt Nut or Nuts may also refer to: Arts, entertainment, and media Com ...
s.


Pollination

Self-pollination is often avoided by means of
protandry Sequential hermaphroditism (called dichogamy in botany) is a type of hermaphroditism that occurs in many fish, gastropods, and plants. Sequential hermaphroditism occurs when the individual changes its sex at some point in its life. In particular, ...
. 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 (including ''
Ptilothrix ''Ptilothrix'' is a genus within the tribe Emphorini of the family Apidae (bumblebees, euglossines, honeybees, stingless bees). Bees of this genus can range from . ''Ptilothrix'' species are solitary, ground-nesting bees. These bees have especia ...
'', ''
Diadasia ''Diadasia'' is a genus of bees in family Apidae. Species of ''Diadasia'' are oligolectic, specialized on a relatively small number of plant species. Their host plants include asters, bindweeds, cacti, mallows, and willowherbs, although mall ...
'', and ''
Melitoma ''Melitoma'' is a genus of chimney bees in the family Apidae. There are about 13 described species in ''Melitoma''. Found in the Nearctic and Neotropics. Species These 11 species belong to the genus ''Melitoma'': * '' Melitoma ameghinoi'' (Holmb ...
'') are known to specialize on the plants.


Importance

A number of species are pests in agriculture, including ''
Abutilon theophrasti ''Abutilon theophrasti'' (velvetleaf, velvet plant, velvetweed, Chinese jute, China jute, crown weed, buttonweed, lantern mallow, butterprint, pie-marker, or Indian mallow) is an annual plant in the family Malvaceae, native to southern Asia. It' ...
'' and '' Modiola caroliniana'', and others that are garden escapees. Cotton (four species of '' Gossypium''), kenaf (''Hibiscus cannabinus''),
cacao Cacao is the seed from which cocoa and chocolate are made, from Spanish cacao, an adaptation of Nahuatl cacaua, the root form of cacahuatl ("bean of the cocoa-tree"). It may also refer to: Plants *''Theobroma cacao'', a tropical evergreen tree ** ...
(''Theobroma cacao''),
kola nut The term kola nut usually refers to the seeds of certain species of plant of the genus ''Cola'', placed formerly in the cocoa family Sterculiaceae and now usually subsumed in the mallow family Malvaceae (as subfamily Sterculioideae). These cola ...
(''Cola spp.''), and okra (''Abelmoschus esculentus'') are important agricultural crops. The fruit and leaves of
baobab ''Adansonia'' is a genus made up of eight species of medium-to-large deciduous trees known as baobabs ( or ). They are placed in the Malvaceae family, subfamily Bombacoideae. They are native to Madagascar, mainland Africa, and Australia.Tropic ...
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 recognised ''Durio'' species, at least nine of which produce edible fruit. ''Durio zibethinus'', native to Borneo and Sumatra, is the onl ...
. A number of species, including '' Hibiscus syriacus'', '' Hibiscus rosa-sinensis'' and '' Alcea rosea'' are garden plants.


See also

* '' Florissantia'', an extinct
Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...
genus in the subfamily Sterculioidea


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