Malpighiales Temporal range: | |
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Flower of Calophyllum inophyllum (Calophyllaceae) | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Eudicots |
Clade: | Rosids |
Clade: | Fabids |
Order: | Malpighiales Juss. ex Bercht. & J.Presl [1] |
Type genus | |
Malpighia | |
Families | |
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Synonyms | |
Rhizophorales |
The Malpighiales comprise one of the largest orders of flowering plants, containing about 36 families and more than 16,000 species, about 7.8% of the eudicots. [2] [3] The order is very diverse, containing plants as different as the willow, violet, poinsettia, manchineel, rafflesia and coca plant, and are hard to recognize except with molecular phylogenetic evidence. It is not part of any of the classification systems based only on plant morphology. Molecular clock calculations estimate the origin of stem group Malpighiales at around 100 million years ago (Mya) and the origin of crown group Malpighiales at about 90 Mya. [4]
The Malpighiales are divided into 32 to 42 families, depending upon which clades in the order are given the taxonomic rank of family. [5] In the APG III system, 35 families were recognized. [1] Medusagynaceae, Quiinaceae, Peraceae, Malesherbiaceae, Turneraceae, Samydaceae, and Scyphostegiaceae were consolidated into other families. The largest family, by far, is the Euphorbiaceae, with about 6300 species in about 245 genera. [6]
In a 2009 study of DNA sequences of 13 genes, 42 families were placed into 16 groups, ranging in size from one to 10 families. The relationships among these 16 groups remain poorly resolved. [5] Malpighiales and Lamiales are the two large orders whose phylogeny remains mostly unresolved. [7]
Some examples of notable species include cassava, a tuber that is a major staple food crop in much of the world; the stinking corpse lily, which produces the largest known flower of any plant; the willows; flaxseed, an important food and fiber crop; Saint John's wort, a herb with a long history of medicinal uses; castor bean, the source of the infamous poison ricin; passionfruit, which produces an edible fruit and psychoactive flowers with a history of traditional medicinal uses; poinsettia, a common ornamental plant; the mangosteen; manchineel tree, one of the most toxic trees in the world; poplars, aspens and cottonwoods which are commonly used for timber – and many more.
Malpighiales is a member of a supraordinal group called the COM clade, which consists of the orders Celastrales, Oxalidales, and Malpighiales. [8] Some describe it as containing a fourth order, Huales, separating the family Huaceae into its own order, separate from Oxalidales. [9]
Some recent studies have placed Malpighiales as sister to Oxalidales sensu lato (including Huaceae), [5] [10] while others have found a different topology for the COM clade. [4] [8] [11]
The COM clade is part of an unranked group known as malvids (rosid II), though formally placed in Fabidae (rosid I). [12] [13] These in turn are part of a group that has long been recognized, namely, the rosids. [3]
The French botanist Charles Plumier named the genus Malpighia in honor of Marcello Malpighi's work on plants; Malpighia is the type genus for the Malpighiaceae, a family of tropical and subtropical flowering plants.
The family Malpighiaceae was the type family for one of the orders created by Jussieu in his 1789 work Genera Plantarum. [14] Friedrich von Berchtold and Jan Presl described such an order in 1820. [15] Unlike modern taxonomists, these authors did not use the suffix "ales" in naming their orders. The name "Malpighiales" is attributed by some to Carl von Martius. [3] In the 20th century, it was usually associated with John Hutchinson, who used it in all three editions of his book, The Families of Flowering Plants. [16] The name was not used by those who wrote later, in the 1970s, '80s, and '90s.
The taxon was largely presaged by Hans Hallier in 1912 in an article in the Archiv. Néerl. Sci. Exact. Nat. titled "L'Origine et le système phylétique des angiospermes", in which his Passionales and Polygalinae were derived from Linaceae (in Guttales), with Passionales containing seven (of eight) families that also appear in the current Malpighiales, namely Passifloraceae, Salicaceae, Euphorbiaceae, Achariaceae, Flacourtiaceae, Malesherbiaceae, and Turneraceae, and Polygalinae containing four (of 10) families that also appear in the current Malpighiales, namely Malpighiaceae, Violaceae, Dichapetalaceae, and Trigoniaceae. [17]
The molecular phylogenetic revolution led to a major restructuring of the order. [2] The first semblance of Malpighiales as now known came from a phylogeny of seed plants published in 1993 and based upon DNA sequences of the gene rbcL. [18] This study recovered a group of rosids unlike any group found in any previous system of plant classification. To make a clear break with classification systems being used at that time, the Angiosperm Phylogeny Group resurrected Hutchinson's name, though his concept of Malpighiales included much of what is now in Celastrales and Oxalidales. [19]
Malpighiales is monophyletic and in molecular phylogenetic studies, it receives strong statistical support. [2] Since the APG II system was published in 2003, minor changes to the circumscription of the order have been made. The family Peridiscaceae has been expanded from two genera to three, and then to four, and transferred to Saxifragales. [5] [20]
The genera Cyrillopsis (Ixonanthaceae), Centroplacus (Centroplacaceae), Bhesa (Centroplacaceae), Aneulophus (Erythroxylaceae), Ploiarium (Bonnetiaceae), Trichostephanus (Samydaceae), Sapria (Rafflesiaceae), Rhizanthes (Rafflesiaceae), and Rafflesia (Rafflesiaceae) had been either added or confirmed as members of Malpighiales by the end of 2009. [5]
Some family delimitations within the order have changed, as well, most notably, the segregation of Calophyllaceae from Clusiaceae sensu lato when it was shown that the latter is paraphyletic. [5] Some differences of opinion on family delimitation exist, as well. For example, Samydaceae and Scyphostegiaceae may be recognized as families or included in a large version of Salicaceae. [21]
The group is difficult to characterize phenotypically, due to sheer morphological diversity, ranging from tropical holoparasites with giant flowers, such as Rafflesia , to temperate trees and herbs with tiny, simple flowers, such as Salix. [2] Members often have dentate leaves, with the teeth having a single vein running into a congested and often deciduous apex (i.e., violoid, salicoid, or theoid). [22] Also, zeylanol has recently been discovered in Balanops and Dichapetalum [23] which are in the balanops clade (so-called Chrysobalanaceae s. l.). The so-called parietal suborder (the clusioid clade and Ochnaceae s. l. were also part of Parietales) corresponds with the traditional Violales as 8 (Achariaceae, Violaceae, Flacourtiaceae, Lacistemataceae, Scyphostegiaceae, Turneraceae, Malesherbiaceae, and Passifloraceae) of the order's 10 families along with Salicaceae, which have usually been assigned as a related order or suborder, [24] are in this most derived malpighian suborder, so that eight of the 10 families of this suborder are Violales. The family Flacourtiaceae has proven to be polyphyletic as the cyanogenic members have been placed in Achariaceae and the ones with salicoid teeth were transferred to Salicaceae. [22] Scyphostegiaceae, consisting of the single genus Scyphostegia has been merged into Salicaceae. [25]
The phylogeny of Malpighiales is, at its deepest level, an unresolved polytomy of 16 clades. [2] It has been estimated that complete resolution of the phylogeny will require at least 25000 base pairs of DNA sequence data per taxon. [26] A similar situation exists with Lamiales and it has been analyzed in some detail. [27] The phylogenetic tree shown below is from Wurdack and Davis (2009). The statistical support for each branch is 100% bootstrap percentage and 100% posterior probability, except where labeled, with bootstrap percentage followed by posterior probability.
Malpighiales |
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In 2012, Xi et al. managed to obtain a more resolved phylogenetic tree than previous studies through the use of data from a large number of genes. They included analyses of 82 plastid genes from 58 species (they ignored the problematic Rafflesiaceae), using partitions identified a posteriori by applying a Bayesian mixture model. Xi et al. identified 12 additional clades and three major, basal clades. [28] [29]
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Changes made in the Angiosperm Phylogeny Group (APG) classification of 2016 (APG IV) were the inclusion of Irvingiaceae, Peraceae, Euphorbiaceae and Ixonanthaceae, together with the transfer of the COM clade from the fabids (rosid I) to the malvids (rosid II). [12]
"Litoh family" is a common name for Ctenolophonaceae, and "koteb family" for Lophopyxidaceae. [30]
The Cucurbitales are an order of flowering plants, included in the rosid group of dicotyledons. 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, with thick pointed petals. The pollination is usually performed by insects, but wind pollination is also present.
Rosales is an order of flowering plants. It is sister to a clade consisting of Fagales and Cucurbitales. It contains about 7,700 species, distributed into about 260 genera. Rosales comprise nine families, the type family being the rose family, Rosaceae. The largest of these families are Rosaceae (91/4828) and Urticaceae (53/2625). The order Rosales is divided into three clades that have never been assigned a taxonomic rank. The basal clade consists of the family Rosaceae; another clade consists of four families, including Rhamnaceae; and the third clade consists of the four urticalean families.
Saxifragales is an order of angiosperms, or flowering plants, containing 15 botanical families and around 100 genera, with nearly 2,500 species. Of the 15 families, many are small, with eight of them being monotypic. The largest family is the Crassulaceae (stonecrops), a diverse group of mostly succulent plants, with about 35 genera. Saxifragales are found worldwide, primarily in temperate to subtropical zones, rarely being encountered growing wild in the tropics; however, many species are now cultivated throughout the world as knowledge of plant husbandry has improved. They can be found in a wide variety of environments, from deserts to fully aquatic habitats, with species adapted to alpine, forested or fully-aquatic habitats. Many are epiphytic or lithophytic, growing on exposed cliff faces, on trees or on rocks, and not requiring a highly organic or nutrient-dense substrate to thrive.
Under the International Code of Nomenclature for algae, fungi, and plants (ICN), Rosidae is a botanical name at the rank of subclass. Circumscription of the subclass will vary with the taxonomic system being used; the only requirement being that it includes the family Rosaceae.
The Salicaceae are the willow family of flowering plants. The traditional family included the willows, poplars. Genetic studies summarized by the Angiosperm Phylogeny Group (APG) have greatly expanded the circumscription of the family to contain 56 genera and about 1220 species, including the tropical Scyphostegiaceae and many of the former Flacourtiaceae.
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 to 1350 species are in about 100 genera. All but seven of these genera are in the large family Celastraceae. Until recently, the composition of the order and its division into families varied greatly from one author to another.
The Flacourtiaceae is a defunct family of flowering plants whose former members have been scattered to various families, mostly to the Achariaceae and Salicaceae. It was so vaguely defined that hardly anything seemed out of place there and it became a dumping ground for odd and anomalous genera, gradually making the family even more heterogeneous. In 1975, Hermann Sleumer noted that "Flacourtiaceae as a family is a fiction; only the tribes are homogeneous."
The Angiosperm Phylogeny Group (APG) is an informal international group of systematic botanists who collaborate to establish a consensus on the taxonomy of flowering plants (angiosperms) that reflects new knowledge about plant relationships discovered through phylogenetic studies.
The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants (angiosperms) which are mainly characterized by having two seed leaves (cotyledons) upon germination. The term derives from dicotyledon. Historically, authors have used the terms tricolpates or non-magnoliid dicots. The current botanical terms were introduced in 1991, by evolutionary botanist James A. Doyle and paleobotanist Carol L. Hotton, to emphasize the later evolutionary divergence of tricolpate dicots from earlier, less specialized, dicots.
The rosids are members of a large clade of flowering plants, containing about 70,000 species, more than a quarter of all angiosperms.
Peridiscaceae is a family of flowering plants in the order Saxifragales. Four genera comprise this family: Medusandra, Soyauxia, Peridiscus, and Whittonia., with a total of 12 known species. It has a disjunct distribution, with Peridiscus occurring in Venezuela and northern Brazil, Whittonia in Guyana, Medusandra in Cameroon, and Soyauxia in tropical West Africa. Whittonia is possibly extinct, being known from only one specimen collected below Kaieteur Falls in Guyana. In 2006, archeologists attempted to rediscover it, however, it proved unsuccessful.
The APG II system of plant classification is the second, now obsolete, version of a modern, mostly molecular-based, system of plant taxonomy that was published in April 2003 by the Angiosperm Phylogeny Group. It was a revision of the first APG system, published in 1998, and was superseded in 2009 by a further revision, the APG III system.
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 endoparasites of vines in the genus Tetrastigma (Vitaceae) and lack stems, leaves, roots, and any photosynthetic tissue. They rely entirely on their host plants for both water and nutrients, and only then emerge as flowers from the roots or lower stems of the host plants.
Lepidobotryaceae is a family of plants in the order Celastrales. It contains only two species: Lepidobotrys staudtii and Ruptiliocarpon caracolito.
When the APG II system of plant classification was published in April 2003, fifteen genera and three families were placed incertae sedis in the angiosperms, and were listed in a section of the appendix entitled "Taxa of uncertain position".
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 further revision, the APG IV system.
Huaceae is a family of plant in the rosids group, which has been classed in the orders Malpighiales, Malvales, and Violales or in its own order Huales. The APG II system placed it in the clade eurosids I, whereas the APG III system of 2009 and APG IV (2016) place it within the Oxalidales. The family is endemic to central Africa. It contains four species in the following two genera:
The superrosids are members of a large clade of flowering plants, containing more than 88,000 species, and thus more than a quarter of all angiosperms.
In phylogenetic nomenclature, the Pentapetalae are a large group of eudicots that were informally referred to as the "core eudicots" in some papers on angiosperm phylogenetics. They comprise an extremely large and diverse group accounting for about 65% of the species richness of the angiosperms, with wide variability in habit, morphology, chemistry, geographic distribution, and other attributes. Classical systematics, based solely on morphological information, was not able to recognize this group. In fact, the circumscription of the Pentapetalae as a clade is based on strong evidence obtained from DNA molecular analysis data.
The superasterids are members of a large clade of flowering plants, containing more than 122,000 species.