Huerteales is the
botanical name
A botanical name is a formal scientific name conforming to the '' International Code of Nomenclature for algae, fungi, and plants'' (ICN) and, if it concerns a plant cultigen, the additional cultivar or Group epithets must conform to the ''Intern ...
for 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.
[Peter F. Stevens (2001 onwards). "Huerteales". In: Angiosperm Phylogeny Website. In: Missouri Botanical Garden Website. (see ''External links'' below)] It is one of the 17 orders that make up the large
eudicot
The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants mainly characterized by having two seed leaves upon germination. The term derives from Dicotyledons.
Traditionally they were called tricolpates or non-magnoliid dicot ...
group known as the
rosids
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 ...
in the
APG III system
The APG III system of flowering plant classification is the third version of a modern, mostly molecular-based, system of plant taxonomy being developed by the Angiosperm Phylogeny Group (APG). Published in 2009, it was superseded in 2016 by a fur ...
of
plant classification.
Within the rosids, it is one of the orders in Malvidae
The rosids are members of a large clade (monophyly, 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 Order (biology), orders, depending upon Circu ...
, a group formerly known as eurosids II
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 ...
and now known informally as the malvids
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 ...
. This is true whether Malvidae is circumscribed
In geometry, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this circle is called the circumcenter and its radius is called the circumradius.
Not every polyg ...
broadly
Vice Media Group LLC is an American-Canadian digital media and broadcasting company. , the Vice Media Group included five main business areas: VICE.com (digital content); VICE STUDIOS (film and TV production) VICE TV (also known as VICELAND); ...
to include eight orders as in APG III, or more narrowly to include only four orders.[ Huerteales consists of four small ]families
Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideall ...
, Petenaeaceae, Gerrardinaceae
''Gerrardina'' is a genus of two species of trees, shrubs, and scrambling shrubs found in southeastern Africa. Until recently, the genus was placed in the polyphyletic family Flacourtiaceae, but it was abnormal there due to its apical placentatio ...
, Tapisciaceae
Tapisciaceae is a family of flowering plants. Until recently it had been abandoned by taxonomists, and it was not recognised in the APG II system of 2003. In the APG III system, however, it has been reinstated to encompass the two genera '' Tapis ...
, and Dipentodontaceae
''Dipentodontaceae'' is a family of flowering plants containing two genera.
# ''Dipentodon'' Dunn - southern China, Assam, Myanmar
# ''Perrottetia'' Kunth in F.W.H.von Humboldt - southern China, Southeast Asia, Papuasia, Queensland, Hawaii, Lati ...
.[Andreas Worberg, Mac H. Alford, Dietmar Quandt, and Thomas Borsch. 2009. "Huerteales sister to Brassicales plus Malvales, and newly circumscribed to include ''Dipentodon, Gerrardina, Huertea, Perrottetia, and ''Tapiscia''. ''Taxon'' 58(2):468-478.]
Petenaeaceae consists of a single genus and species ''Petenaea
''Petenaea cordata'' (from northern Central America) was first described in Elaeocarpaceae and later placed in Tiliaceae, but most authors have been uncertain about its familial affinities. It was considered a taxon ''incertae sedis'' in the Angi ...
cordata'' from Southern Mexico, Guatemala and Belize.[Christenhusz, M. J. M., Fay, M. F., Clarkson, J. J., Gasson, P., Morales Can, J., Jiménez Barrios, J. B. & Chase, M. W. (2010). Petenaeaceae, a new angiosperm family in Huerteales with a distant relationship to ''Gerrardina'' (Gerrardinaceae). ]Botanical Journal of the Linnean Society
The ''Botanical Journal of the Linnean Society'' is a scientific journal publishing original papers relating to the taxonomy of all plant groups and fungi, including anatomy, biosystematics, cytology, ecology, ethnobotany, electron microscopy, mo ...
164: 16–25.
Gerrardinaceae consists of a single genus, '' Gerrardina''.[Mac H. Alford. 2006. "Gerrardinaceae: a new family of African flowering plants unresolved among Brassicales, Huerteales, Malvales, and Sapindales." ''Taxon'' 55(4):959-964.] Tapisciaceae has two genera, '' Tapiscia'' and ''Huertea
''Huertea'' is a genus of plant
Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all ...
''.[Klaus Kubitzki. 2003. "Tapisciaceae" pages 369-370. In: Klaus Kubitski and Clemens Bayer (editors). ''The Families and Genera of Vascular Plants'' volume V. Springer-Verlag: Berlin;Heidelberg, Germany.][Dezhu Li, Jie Cai, and Wen Jun. 2008. "Tapisciaceae" page 496. In: Zhengyi Wu, Peter H. Raven, and Deyuan Hong (editors). ''Flora of China'' volume 11. Science Press: Beijing, China; Missouri Botanical Garden Press: St. Louis, Missouri, USA.]
Until 2006, Dipentodontaceae was treated as consisting of a single genus, ''Dipentodon
Dipentodon is a genus of flowering plants in the Family (biology), family Dipentodontaceae. Its only species, ''Dipentodon sinicus'', is a small, deciduous tree native to southern China, northern Myanmar, and northern India.Jinshuang Ma and Bruce ...
''.[Vernon H. Heywood, Richard K. Brummitt, Ole Seberg, and Alastair Culham. ''Flowering Plant Families of the World''. Firefly Books: Ontario, Canada. (2007).] Since that time, some authors have included ''Perrottetia'' in Dipentodontaceae, even though no formal revision of the family has been published as of 2008.[Jinshuang Ma and Bruce Bartholomew. 2008. "Dipentodontaceae" pages 494-495. In: Zhengyi Wu, Peter H. Raven, and Deyuan Hong (editors). ''Flora of China'' volume 11. Science Press: Beijing, China; Missouri Botanical Garden Press: St. Louis, Missouri, USA.] Thus the order Huerteales consists of six genera. The largest genus, ''Perrottetia'', contains about 15 of the approximate total of 25 species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
in the order.[Mark P. Simmons. 2004. "Celastraceae" page 50. In: Klaus Kubitzki (editor). ''The Families and Genera of Vascular Plants'' volume VI. Springer-Verlag: Berlin;Heidelberg, Germany.]
The Huerteales are 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 or small 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, including only woody plants with secondary growth, plants that are ...
s found in most tropical
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 ...
or warm 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 ...
regions. The flowers of ''Perrottetia'' have been studied in detail,[Merran L. Matthews and Peter K. Endress (2005). "Comparative floral structure and systematics in Celastrales". ''Botanical Journal of the Linnean Society'' 149(2):129-194] but otherwise, all five of the genera are poorly known. The order is based on molecular phylogenetic
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 ...
analysis of DNA sequence
DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...
s.
Description
All of the Huerteales are woody plant
A woody plant is a plant that produces wood as its structural tissue and thus has a hard stem. In cold climates, woody plants further survive winter or dry season above ground, as opposite to herbaceous plants that die back to the ground until sp ...
s. The leaves are alternate
Alternative or alternate may refer to:
Arts, entertainment and media
* Alternative (''Kamen Rider''), a character in the Japanese TV series ''Kamen Rider Ryuki''
* ''The Alternative'' (film), a 1978 Australian television film
* ''The Alternative ...
with toothed margins. The inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...
is cymose
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...
, but sometimes nearly racemose
A raceme ( or ) or racemoid is an unbranched, indeterminate type of inflorescence bearing flowers having short floral stalks along the shoots that bear the flowers. The oldest flowers grow close to the base and new flowers are produced as the s ...
or umbel
In botany, an umbel is an inflorescence that consists of a number of short flower stalks (called pedicels) that spread from a common point, somewhat like umbrella ribs. The word was coined in botanical usage in the 1590s, from Latin ''umbella'' "p ...
liform. The bases of the calyx, corolla and 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 fused to form a 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. It ...
which is in some cases very short. 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 unilocular
A locule (plural locules) or loculus (plural loculi) (meaning "little place" in Latin) is a small cavity or compartment within an organ or part of an organism (animal, plant, or fungus).
In angiosperms (flowering plants), the term ''locule'' usu ...
, at least at the top, with one or two ovule
In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the fe ...
s per carpel
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'' ...
. The number of carpels is variable.
Other characters are generally found in Huerteales, but with the exceptions noted below. ''Gerrardina'' differs from the rest of Huerteales in that the stamens are opposite the 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, instead of being opposite the sepals
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 The term ''sepalum'' was coined b ...
. ''Dipentodon'' and ''Perrottetia'' are distinctive in that the calyx and corolla are not well differentiated, but resemble each other. ''Tapiscia'' and ''Huertea'' have a calyx tube and compound
Compound may refer to:
Architecture and built environments
* Compound (enclosure), a cluster of buildings having a shared purpose, usually inside a fence or wall
** Compound (fortification), a version of the above fortified with defensive struct ...
, rather than simple leaves. ''Tapiscia'' has a uniloculate ovary with a single ovule. ''Huertea'' has one locule containing two ovules, or two locules, each containing one ovule. ''Gerrardina'', ''Dipentodon'', and ''Perrottetia'' have two ovules in each locule. ''Tapiscia'' lacks the nectar
Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
y disk that is characteristic of the order. ''Huertea'' lacks stipules.
History
Until the first decade of the twenty-first century, the five genera of Huerteales had usually been placed into three unrelated families. ''Tapiscia'' and ''Huertea'' had long been known to be related. Most authors had placed them in Staphyleaceae
Staphyleaceae is a small family of flowering plants in the order Crossosomatales, native to Europe, temperate and tropical Asia and the Americas. The largest genus ''Staphylea'', which gives the family its name, contains the "bladdernut" trees. ...
and had placed that family in the order Sapindales
Sapindales is an order of flowering plants. Well-known members of Sapindales include citrus; maples, horse-chestnuts, lychees and rambutans; mangos and cashews; frankincense and myrrh; mahogany and neem.
The APG III system of 2009 includes it ...
. Armen Takhtajan
Armen Leonovich Takhtajan or Takhtajian ( hy, Արմեն Լևոնի Թախտաջյան; russian: Армен Леонович Тахтаджян; surname also transliterated Takhtadjan, Takhtadzhi︠a︡n or Takhtadzhian, pronounced takh-tuh-JA ...
established the family Tapisciaceae in 1987 and placed it in Sapindales, but this treatment was not followed by many others and it did not stand up to phylogenetic
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analysis. Since that time, Staphyleaceae has been recircumscribed. It no longer includes ''Tapiscia'' and ''Huertea'' [Sarah L. Simmons. 2007. "Staphyleaceae" pages 440-445. In: Klaus Kubitzki (editor). ''The Families and Genera of Vascular Plants'' volume IX. Springer-Verlag: Berlin;Heidelberg, Germany.] and it is in the order Crossosomatales
The Crossosomatales are an order, first recognized as such by APG II. They are flowering plants included within the Rosid eudicots.
Description
Species assigned to the Crossosomatales have in common flowers that are positioned solitarily, with ...
.[Sang-Hun Oh and Daniel Potter. 2006. "Description and Phylogenetic Position of a New Angiosperm Family, Guamatelaceae, Inferred from Chloroplast ''rbcL, atpB,'' and ''matK'' Sequences." ''Systematic Botany'' 31(4):730-738.]
For most of the twentieth century, ''Gerrardina'' and ''Dipentodon'' had usually been placed in Flacourtiaceae, a family that is now recognized by only a few taxonomists
In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given ...
, and then only as a segregate of Salicaceae
The Salicaceae is the willow family of flowering plants. The traditional family (Salicaceae ''sensu stricto'') included the willows, poplar, aspen, and cottonwoods. Genetic studies summarized by the Angiosperm Phylogeny Group (APG) have greatly ...
.[Mark W. Chase, Sue Zmarzty, M. Dolores Lledó, Kenneth J. Wurdack, Susan M. Swensen, and Michael F. Fay. 2002. "When in doubt, put it in Flacourtiaceae: a molecular phylogenetic analysis based on plastid ''rbcL'' DNA sequences." ''Kew Bulletin'' 57(1):141-181.][Sue Zmarzty et alii. (in press). "Salicaceae" In: ''The Families and Genera of Vascular Plants''. Springer-Verlag: Berlin;Heidelberg, Germany.] ''Perrottetia'', meanwhile, had usually been placed, with considerable doubt, in Celastraceae
The Celastraceae (staff-vine or bittersweet) are a family of 97 genera and 1,350 species of herbs, vines, shrubs and small trees, belonging to the order Celastrales. The great majority of the genera are tropical, with only ''Celastrus'' (the staf ...
.[Li-Bing Zhang and Mark P. Simmons (2006). "Phylogeny and Delimitation of the Celastrales Inferred from Nuclear and Plastid Genes". ''Systematic Botany'' 31(1):122-137.]
Ever since ''Dipentodon'' was named in 1911, there had been occasional suggestions that it might be related to ''Tapiscia'' and ''Huertea''. In 2001, Alexander Doweld established the order Huerteales,[James L. Reveal. 2008 onward. "A Checklist of Family and Suprafamilial Names for Extant Vascular Plants." At: Home page of James L. Reveal and C. Rose Broome. (see ''External links'' below).] defining it to consist of ''Tapiscia'', ''Huertea'', and ''Dipentodon''.[Alexander B. Doweld. 2001. ''Tentamen Systematis Plantarum Vascularium (Tracheophyta).'': xxxv. 23 Dec 2001.] This grouping was later supported by molecular phylogenetic
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 ...
studies. In 2006, a study of DNA sequence
DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...
s showed that ''Perrottetia'' was misplaced in 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 that it is sister
A sister is a woman or a girl who shares one or more parents with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a familial relationship, it is sometimes used endearingly to refer to ...
to ''Dipentodon'' in Huerteales. Also in 2006, it was found that ''Gerrardina'' is a malvid, but its placement within this group remained uncertain.
In 2009, Andreas Worberg and co-authors produced the first phylogenetic study that included all of the genera of Huerteales. From one of their data matrices, they derived a well supported phylogeny for the order, as well as strongly supported relationships among the four orders of malvids.
Phylogeny
The phylogeny shown below is the one found by Worberg and co-authors. Monospecific
In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
genera are represented by species names.
References
External links
Huerteales
At
At
Missouri Botanical Garden Website
At
James L. Reveal
{{Taxonbar, from=Q745931
Angiosperm orders