Theophrastoideae | |
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Jacquinia pungens flowers | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Eudicots |
Clade: | Asterids |
Order: | Ericales |
Family: | Primulaceae |
Subfamily: | Theophrastoideae A.DC. [1] [2] |
Type genus | |
Theophrasta L. | |
Tribes | |
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Synonyms | |
Theophrastoideae is a small subfamily of flowering plants in the family Primulaceae. It was formerly recognized as a separate family Theophrastaceae. As previously circumscribed, the family consisted of eight genera and 95 species of trees or shrubs, native to tropical regions of the Americas.
The two subclades or tribes of Theophrastoideae, Theophrasteae (Theophrastaceae s.s.) and Samoleae, share only the presence of staminodes. The species of Samolus are herbaceous perennials and characterised by perigynous flowers. [4] The remaining genera (Theophrastaceae s.s.) are generally evergreen shrubs or small trees, with hypogynous flowers. [5] [6]
Linnaeus, in formally describing the genera, placed Theophrasta and related genera in a group he named Pentandria Monogynia (i.e 5 stamens, one pistil), his system being based on sexual characteristics. [7] Jussieu arranged Linnaeus' genera in a hierarchical system of ranks based on the relative value of a much wider range of characteristics. In his Genera plantarum (1789) he organised the primuloid genera into two Ordo (families), within a class (VIII) he called Dicotyledones Monopetalae Corolla Hypogyna, based on the cotyledons (two), form of the petals (fused), and position of the corolla with respect to the ovary (below). [8] Jussieu's families were the Lysimachiae, including Primula and Theophrasta [9] and the Sapotae, including Myrsine, these being the three main lineages in modern understanding of the Primulaceae. [10]
Don described a family of Theophrasteaceae in 1836, with four genera, Theophrasta , Clavija , Jacquinia and Leonia , of which the latter was determined unrelated, and placed this family as closely related to Myrsineae and Sapoteae. [3] Later, De Candolle more formally described a family, Theophrastaceae, based on the genus Theophrasta, in 1844, with six genera, Theophrasta, Clavija, Jacquinia, Oncinus , Monotheca and Reptonia . The latter three are no longer considered related. [2]
Theophrastaceae were included in the order Primulales by Cronquist (1988). [11] The APG system (1998) submerged that order in an enlarged order Ericales (Ericales s.l. ), a basal group in the asterids, where the families of Primulales formed a monophyletic primuloid clade. Subsequent molecular phylogenetic analysis showed that the genus Samolus (brook weeds), [4] with about 12–15 additional species and traditionally placed within Primulaceae, as tribe Samoleae, was more closely related to the Theophrastaceae and suggested its transfer. Briefly Samolus was considered a separate family, Samolaceae. The third revision of the APG, APG III (2009) realigned all the primuloid families within a greatly enlarged Primulaceae (Primulaceae s.l.), in which each of the existing families became a subfamily. The newly described Theophrastoideae included Samolus, vastly increasing the area of distribution. [12] [13]
The cladogram below shows the infrafamilial phylogenetic relationships of Primulaceae, together with the subfamilial crown ages. Maesoideae forms the basal group, while Primuloideae and Myrsinoideae are in a sister group relationship. [14] [15] [16] [17]
Primulaceae s.l. |
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The phylogenetic relationships of the 8 accepted genera are shown in the cladogram, in which Samolus forms the basal group and is sister to all other Theophrastoideae (Theophrastaceae s.s.), [6] the remaining genera forming two subclades. Alternatively these two subclasses have been designated as two tribes, Samoleae and Theophrasteae: [13]
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Phylogenetic relationships of Theophrastoideae [18] [19] |
The Theophrasteae consist of seven genera and about 100 species, while Samoleae has only the single genus Samolus, with about 12–15 dozen species. [4] In 1903, Theophrastaceae consisted of four genera, Clavija, Jacquinia, Deherainia , and Theophrasta. [20] In 1904, a species of Deherainia was segregated to form the novel genus Neomezia , to create five genera [21] and in 1993 a species of Jacquinia was segregated to form a sixth genus, Votschia . Molecular phylogenetic analyses revealed that Jacquinia was still paraphyletic consisting of two separate and distinct clades, necessitating splitting off another new genus, Bonellia , to make seven genera in total in this tribe. [6]
Theophrastoideae takes its name from the nominative and type genus, Theophrasta, named by Linnaeus after the Ancient Greek philosopher and biologist Theophrastus.[ citation needed ]
The botanical authority for the previous family, Theophrastaceae, belongs to David Don (D.Don) for his first description of the family in 1835. [3] The subsequently submerged subfamily bears the authority of Alphonse de Candolle (A.DC.) for his formal conspectus of Theophrastaceae in 1844. [2] The authority for the tribe Theophrasteae is that of Bartling who used the term Theophrastea to describe a grouping of genera, including Theophrasta within the family Ardisiaceae in 1830. This represents the earliest creation of a suprageneric taxon for these genera. The Ardisiaceae were later included in the other primuloid family, Myrsinaceae (Myrsinoideae).
The species of Theophrasteae are largely neotropical, confined to Mesoamerica, Central and South America and the Caribbean. [22] [23]
In contrast Samolus species are mainly restricted to different continents in the Southern Hemisphere with about 4–6 species restricted to North America. However the generic type, Samolus valerandi is near cosmopolitan in its distribution. [23]
Theophrastaceae s.s. are mainly found in lowland regions which have a seasonal, dry climate, and prefer coastal thickets, dry shrub vegetation, or dry deciduous or semideciduous forests. However a number of species of Clavija are found in low montane and lowland rain forests. [22]
Unlike much of the traditional Theophrastoideae, Samolus is found in either flooded areas around rivers and lakes, or in salt marshes. [4]
The Ericales are a large and diverse order of dicotyledons. Species in this order have considerable commercial importance including for tea, persimmon, blueberry, kiwifruit, Brazil nuts, argan, and azalea. The order includes trees, bushes, lianas, and herbaceous plants. Together with ordinary autophytic plants, the Ericales include chlorophyll-deficient mycoheterotrophic plants and carnivorous plants.
The Ericaceae are a family of flowering plants, commonly known as the heath or heather family, found most commonly in acidic and infertile growing conditions. The family is large, with c. 4250 known species spread across 124 genera, making it the 14th most species-rich family of flowering plants. The many well known and economically important members of the Ericaceae include the cranberry, blueberry, huckleberry, rhododendron, and various common heaths and heathers.
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. 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.
Monocotyledons, commonly referred to as monocots, are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of the major groups into which the flowering plants have traditionally been divided; the rest of the flowering plants have two cotyledons and are classified as dicotyledons, or dicots.
The Actinidiaceae are a small family of flowering plants. The family has three genera and about 360 species and is a member of the order Ericales.
Geraniales is a small order of flowering plants, included within the rosid subclade of eudicots. The largest family in the order is Geraniaceae with over 800 species. In addition, the order includes the smaller Francoaceae with about 40 species. Most Geraniales are herbaceous, but there are also shrubs and small trees.
The Primulaceae, commonly known as the primrose family, are a family of herbaceous and woody flowering plants including some favourite garden plants and wildflowers. Most are perennial though some species, such as scarlet pimpernel, are annuals.
Lysimachia is a genus consisting of 182 accepted species of flowering plants traditionally classified in the family Primulaceae. Based on a molecular phylogenetic study it was transferred to the family Myrsinaceae, before this family was later merged into the Primulaceae.
Anagallis is a genus of about 20–25 species of flowering plants in the family Primulaceae, commonly called pimpernel. The scarlet pimpernel referred to in literature is part of this genus. The botanical name is from the Greek ana and agállein, and it refers to the opening and closing of the flowers in response to environmental conditions.
Myrsinoideae is a subfamily of the family Primulaceae in the order Ericales. It was formerly recognized as the family Myrsinaceae, or the myrsine family, consisting of 35 genera and about 1000 species. It is widespread in temperate to tropical climates extending north to Europe, Siberia, Japan, Mexico, and Florida, and south to New Zealand, South America, and South Africa.
The Clethraceae are a small family of flowering plants in the order Ericales, composed of two genera, Clethra and Purdiaea, with approximately 75 species. They are native to warm temperate to tropical regions of Asia and the Americas, with one species also on Madeira.
Diapensiaceae is a small family of flowering plants, which includes 15 species in 6 genera. The genera include Berneuxia Decne., Diapensia L., Galax Sims, Pyxidanthera Michx., Shortia Torr. & A.Gray, and Schizocodon Siebold & Zucc.. Members of this family have little economic importance; however, some members are cultivated by florists.
Samolus is a widely distributed genus of about a dozen species of water-loving herbs and amphibious flowering plants. According to the APG III system of classification, the genus falls within the primrose family, Primulaceae, under the order containing the rhododendroid eudicots, Ericales. It was considered closely related to a clade comprising the Theophrastaceae, and was likewise treated as part of that family. The APG III system does not recognize these families, instead classifying all former Theophrastaceae genera within the family Primulaceae.
The Cyrillaceae are a small family of flowering plants in the order Ericales, native to warm temperate to tropical regions of the Americas. The family comprises two genera, Cliftonia and Cyrilla, each containing a single species, Cliftonia monophylla and Cyrilla racemiflora. However, additional species of Cyrilla are now often recognized and the genus is in need of taxonomic revision.
Boraginales is an order of flowering plants in the asterid clade, with a total of about 125 genera and 2,700 species. Different taxonomic treatments either include only a single family, the Boraginaceae, or divide it into up to eleven families. Its herbs, shrubs, trees and lianas (vines) have a worldwide distribution.
Sladeniaceae is a family of flowering plants containing tree species found in subtropical to tropical environments in East Africa (Ficalhoa), Burma, Yunnan, and Thailand (Sladenia). The family consists of trees with alternate, simple leaves without stipules, and flowers arranged in cymose inflorescences.
Pentaphylax euryoides is a species of flowering plant in the Pentaphylacaceae family. It is the sole species in genus Pentaphylax. It is a shrub or small tree native to southern China, Vietnam, Laos, Peninsular Malaysia, and northern Sumatra. In China, it is found in Guangdong, Guangxi, Guizhou, Hainan, and Yunnan provinces, and in southern parts of Fujian, Hunan, and Jiangxi.
Anagallis tenella, known in Britain as the bog pimpernel, is a low growing perennial plant found in a variety of damp habitats from calcareous dune slacks to boggy and peaty heaths in Eurasia. In the United Kingdom it is mostly restricted to the western half of the country, although it was more common in the east before land drainage and intensification of farming in that area.
Lysimachia clethroides, the gooseneck loosestrife, is a species of flowering plant, traditionally classified in the family Primulaceae. It was transferred to the family Myrsinaceae based on a molecular phylogenetic study, but this family was later merged into the Primulaceae.
The taxonomy of the plant family Liliaceae has had a complex history since its first description in the mid-eighteenth century. Originally, the Liliaceae were defined as having a "calix" (perianth) of six equal-coloured parts, six stamens, a single style, and a superior, three-chambered (trilocular) ovary turning into a capsule fruit at maturity. The taxonomic circumscription of the family Liliaceae progressively expanded until it became the largest plant family and also extremely diverse, being somewhat arbitrarily defined as all species of plants with six tepals and a superior ovary. It eventually came to encompass about 300 genera and 4,500 species, and was thus a "catch-all" and hence paraphyletic. Only since the more modern taxonomic systems developed by the Angiosperm Phylogeny Group (APG) and based on phylogenetic principles, has it been possible to identify the many separate taxonomic groupings within the original family and redistribute them, leaving a relatively small core as the modern family Liliaceae, with fifteen genera and 600 species.
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