| Theophrastoideae | |
|---|---|
 | |
| 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 | |
| TheophrastaL. | |
| Tribes | |
| |
| Synonyms | |
Theophrastoideae is a small subfamily offlowering plants in the familyPrimulaceae. It was formerly recognized as a separate familyTheophrastaceae. As previously circumscribed, the family consisted of eight genera and 95 species oftrees orshrubs, native to tropical regions of theAmericas.
The two subclades ortribes of Theophrastoideae, Theophrasteae (Theophrastaceaes.s.) and Samoleae, share only the presence ofstaminodes. The species ofSamolus areherbaceousperennials and characterised byperigynousflowers.[4] The remaining genera (Theophrastaceaes.s.) are generally evergreen shrubs or small trees, withhypogynous flowers.[5][6]
Linnaeus, in formally describing the genera, placedTheophrasta and related genera in a group he namedPentandria Monogynia (i.e 5stamens, onepistil), his system being based on sexual characteristics.[7]Jussieu arranged Linnaeus' genera in ahierarchical system of ranks based on the relative value of a much wider range of characteristics. In hisGenera plantarum (1789) he organised the primuloid genera into twoOrdo (families), within aclass (VIII) he calledDicotyledones Monopetalae Corolla Hypogyna, based on thecotyledons (two), form of the petals (fused), and position of thecorolla with respect to theovary (below).[8] Jussieu's families were the Lysimachiae, includingPrimula andTheophrasta[9] and theSapotae, includingMyrsine, these being the three main lineages in modern understanding of thePrimulaceae.[10]
Don described a family of Theophrasteaceae in 1836, with four genera,Theophrasta,Clavija,Jacquinia andLeonia, 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 genusTheophrasta, in 1844, with six genera,Theophrasta,Clavija,Jacquinia,Oncinus,Monotheca andReptonia. The latter three are no longer considered related.[2]
Theophrastaceae were included in the orderPrimulales byCronquist (1988).[11] TheAPG system (1998) submerged that order in an enlarged orderEricales (Ericaless.l.), abasal group in theasterids, where the families of Primulales formed amonophyletic primuloidclade. Subsequentmolecular phylogenetic analysis showed that the genusSamolus (brook weeds),[4] with about 12–15 additional species and traditionally placed within Primulaceae, astribe Samoleae, was more closely related to the Theophrastaceae and suggested its transfer. BrieflySamolus 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 (Primulaceaes.l.), in which each of the existing families became a subfamily. The newly described Theophrastoideae includedSamolus, vastly increasing the area of distribution.[12][13]
Thecladogram below shows the infrafamilial phylogenetic relationships of Primulaceae, together with the subfamilialcrown ages. Maesoideae forms the basal group, while Primuloideae and Myrsinoideae are in asister 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 whichSamolus forms the basal group and is sister to all other Theophrastoideae (Theophrastaceaes.s.),[6] the remaining genera forming two subclades. Alternatively these two subclasses have been designated as two tribes, Samoleae and Theophrasteae:[13]
| Phylogenetic relationships of Theophrastoideae[18][19] |
The Theophrasteae consist of seven genera and about 100 species, while Samoleae has only the single genusSamolus, with about 12–15 dozen species.[4] In 1903, Theophrastaceae consisted of four genera,Clavija,Jacquinia,Deherainia, andTheophrasta.[20] In 1904, a species ofDeherainia wassegregated to form the novel genusNeomezia, to create five genera[21] and in 1993 a species ofJacquinia was segregated to form a sixth genus,Votschia. Molecular phylogenetic analyses revealed thatJacquinia was stillparaphyletic 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 biologistTheophrastus.[citation needed]
The botanical authority for the previous family, Theophrastaceae, belongs toDavid Don (D.Don) for his first description of the family in 1835.[3] The subsequently submerged subfamily bears the authority ofAlphonse de Candolle (A.DC.) for his formalconspectus of Theophrastaceae in 1844.[2] The authority for the tribe Theophrasteae is that ofBartling who used the term Theophrastea to describe a grouping of genera, including Theophrasta within the familyArdisiaceae 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 largelyneotropical, confined toMesoamerica, Central and South America and the Caribbean.[22][23]
In contrastSamolus species are mainly restricted to different continents in the Southern Hemisphere with about 4–6 species restricted to North America. However thegeneric type,Samolus valerandi is nearcosmopolitan in its distribution.[23]
Theophrastaceaes.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 ofClavija 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 insalt marshes.[4]
Media related toTheophrastoideae at Wikimedia Commons
Data related toTheophrastoideae at Wikispecies