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Theophrastoideae

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Theophrastoideae
Jacquinia pungens flowers
Scientific classification Edit this 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
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.

Description

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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

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History

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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

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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

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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]

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

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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

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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

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Distribution

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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

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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]

References

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  1. ^ Reveal 2012.
  2. ^ a b c de Candolle 1844.
  3. ^ a b c Don 1836.
  4. ^ a b c d Wanntorp & Anderberg 2011.
  5. ^ Stahl 2004.
  6. ^ a b c 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. ^ a b 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. ^ a b Ståhl 2010.
  23. ^ a b Jones et al 2012.

Bibliography

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Books

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Articles

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Samolus

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APG

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Websites

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Media related to Theophrastoideae at Wikimedia Commons Data related to Theophrastoideae at Wikispecies

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