Theophrastoideae

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Theophrastoideae
Jacquinia-pungens.jpg
Jacquinia pungens flowers
Scientific classification OOjs UI icon edit-ltr.svg
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
Synonyms

Theophrastaceae D.Don [3]

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.

Contents

Description

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]

Taxonomy

History

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]

Phylogeny

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.

Maesoideae

24.1mya

Theophrastoideae

70mya

Primuloideae

46.4mya

Myrsinoideae

53.2mya

Subdivision

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]

Theophrastoideae
Samoleae

Samolus L.

Theophrasteae

Theophrasta L.

Neomezia Votsch

Clavija Ruiz & Pav.

Jacquinia L.

Deherainia Decne.

Votschia B.Ståhl

Bonellia Bertero ex Colla

(Theophrastaceae s.s.)
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]

Etymology

Theophrastoideae takes its name from the nominative and type genus, Theophrasta, named by Linnaeus after the Ancient Greek philosopher and biologist Theophrastus.[ citation needed ]

Botanical authority

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).

Distribution and habitat

Distribution

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]

Habitat

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]

Related Research Articles

<span class="mw-page-title-main">Ericales</span> Order of eudicot flowering plants

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.

<span class="mw-page-title-main">Ericaceae</span> Heather family of flowering 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.

<span class="mw-page-title-main">Malpighiales</span> Eudicot order of flowering plants

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.

<span class="mw-page-title-main">Monocotyledon</span> Clade of flowering plants

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.

<span class="mw-page-title-main">Actinidiaceae</span> Family of flowering plants

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.

<span class="mw-page-title-main">Geraniales</span> Order of flowering plants in the rosid subclade of eudicots

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.

<span class="mw-page-title-main">Primulaceae</span> Family of flowering plants that includes the primroses

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.

<i>Lysimachia</i> Genus of flowering plants in the family Primulaceae

Lysimachia is a genus consisting of 193 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.

<i>Anagallis</i> Genus of flowering plants

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.

<span class="mw-page-title-main">Myrsinoideae</span> Subfamily of plants, including Cyclamen

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.

<span class="mw-page-title-main">Clethraceae</span> Family of flowering plants

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.

<span class="mw-page-title-main">Diapensiaceae</span> Family of flowering plants

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.

<i>Samolus</i> Genus of flowering plants in the family Primulaceae

Samolus is a widely distributed genus of about a dozen species of water-loving herbs. According to the APG III classification, this genus belongs to the family Primulaceae in the order Ericales. It was considered as closely related to a clade comprising the Theophrastaceae, and was treated as part of that family or in its own monogeneric family, the Samolaceae. The APG III system does not recognize these families and instead includes all genera formerly belonging to Theophrastaceae in the family Primulaceae.

<span class="mw-page-title-main">Peridiscaceae</span> Family of flowering plants in the order Saxifragales

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.

<span class="mw-page-title-main">Cyrillaceae</span> Family of flowering plants

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.

<span class="mw-page-title-main">Boraginales</span> Order of flowering plants within the lammiid clade of eudicots

Boraginales is an order of flowering plants in the asterid clade. It includes the Boraginaceae and a number of other families, with a total of about 125 genera and 2,700 species. 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 is a genus of flowering plants, with one or two species which are shrubs and small trees. The species has simple evergreen leaves that are alternately arranged on the stems. The flower pistil is 5 celled and the anthers dehiscing via pores and basifixed. Flowers have five petals and five sepals that are distinct from each other and five stamens attached oppositely to the sepals. The seeds are winged.

<i>Anagallis tenella</i> Species of flowering plant

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.

<i>Lysimachia clethroides</i> Species of flowering plant

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.

References

  1. Reveal 2012.
  2. 1 2 3 de Candolle 1844.
  3. 1 2 3 Don 1836.
  4. 1 2 3 4 Wanntorp & Anderberg 2011.
  5. Stahl 2004.
  6. 1 2 3 Ståhl & Källersjö 2004.
  7. Linnaeus 1753.
  8. Jussieu 1789, p. 93.
  9. Jussieu 1789, p. 95.
  10. Jussieu 1789, p. 151.
  11. Cronquist 1988.
  12. APG III 2009.
  13. 1 2 Stevens 2021a.
  14. Soltis et al 2018.
  15. Anderberg et al 2002.
  16. Byng 2014.
  17. Rose et al 2018.
  18. Ståhl 2010, p. 20.
  19. Kallersjo et al 2000.
  20. Mez 1903.
  21. Votsch 1904.
  22. 1 2 Ståhl 2010.
  23. 1 2 Jones et al 2012.

Bibliography

Books

Articles

Samolus

APG

Websites

Commons-logo.svg Media related to Theophrastaceae at Wikimedia Commons Wikispecies-logo.svg Data related to Theophrastoideae at Wikispecies