Boraginales is an order of flowering plants in the

asterid In the APG IV system (2016) for the classification of flowering plants, the name asterids denotes a clade (a monophyletic group). Asterids is the largest group of flowering plants, with more than 80,000 species, about a third of the total floweri ...

clade. It includes the

Boraginaceae Boraginaceae, the borage or forget-me-not family, includes about 2,000 species of shrubs, trees and herbs in 146, to 156 genera with a worldwide distribution. The APG IV system from 2016 classifies the Boraginaceae as single family of the order ...

and a number of other families, with a total of about 125 genera and 2,700 species. Its herbs, shrubs, trees and

lianas A liana is a long- stemmed, woody vine that is rooted in the soil at ground level and uses trees, as well as other means of vertical support, to climb up to the canopy in search of direct sunlight. The word ''liana'' does not refer to a ta ...

(vines) have a worldwide distribution.

Taxonomy

History

The classification of plants now known as Boraginales dates to the ''Genera plantarum'' (1789) when

Antoine Laurent de Jussieu Antoine Laurent de Jussieu (; 12 April 1748 – 17 September 1836) was a French botanist, notable as the first to publish a natural classification of flowering plants; much of his system remains in use today. His classification was based on an e ...

named a group of plants Boragineae, to include the genus ''

Borago ''Borago'', or borage, is a genus of five species of herbs native to the Mediterranean, with one species, ''Borago officinalis'', cultivated and naturalized throughout the world. Uses ''Borago officinalis'' is used medicinally, in companion pl ...

'', now the

type genus In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearing type of a nominal f ...

. However, since the first valid description was by

Friedrich von Berchtold Count Friedrich Carl Eugen Vsemir von Berchtold, baron von Ungarschitz ( cz, Bedřich Karel Eugen Všemír Berchtold hrabě z Uherčic; 25 October 1781 – 3 April 1876), was a German-speaking Bohemian physician and botanist from Austrian descen ...

and

Jan Svatopluk Presl Jan Svatopluk Presl (4 September 1791 – 6 April 1849) was a Czech natural scientist. He was the brother of botanist Carl Borivoj Presl (1794–1852). The Czech Botanical Society commemorated the two brothers by naming its principal publicatio ...

(1820), the

botanical authority In botanical nomenclature, author citation is the way of citing the person or group of people who validly published a botanical name, i.e. who first published the name while fulfilling the formal requirements as specified by the '' International Co ...

is given as Juss. ex Bercht. & J.Presl, where the ''ex'' refers to the prior authority of Jussieu. Lindley (1853) changed the name to the modern Boraginaceae. Jussieu divided the Boragineae into five groups. Since then

has been treated either as a large family with several subfamilies, or as a smaller family with several closely related families. The family had been included in a number of higher order taxa, but in 1926 Hutchinson erected a new order, Boraginales, to include the Boraginaceae. Although Boraginales was included in a number of taxonomic classifications including Dahlgren (1980),

Takhtajan Armen Leonovich Takhtajan or Takhtajian ( hy, Արմեն Լևոնի Թախտաջյան; russian: Армен Леонович Тахтаджян; surname also transliterated Takhtadjan, Takhtadzhi︠a︡n or Takhtadzhian, pronounced takh-tuh-JA ...

(1997) and Kubitzki (2016) as an order, it was not recognized in either of two major

systems A system is a group of interacting or interrelated elements that act according to a set of rules to form a unified whole. A system, surrounded and influenced by its environment, is described by its boundaries, structure and purpose and express ...

, the

Cronquist system The Cronquist system is a taxonomic classification system of flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of Flowering Plants'' (1968; 2nd edition, 1988) a ...

and the

APG system The APG system (Angiosperm Phylogeny Group system) of plant classification is the first version of a modern, mostly molecular-based, system of plant taxonomy. Published in 1998 by the Angiosperm Phylogeny Group, it was replaced by the improved AP ...

. In the

, Boraginaceae (including

Cordiaceae Cordioideae is a subfamily of the flowering plant family Boraginaceae. Genera *''Coldenia'' L. *''Cordia ''Cordia'' is a genus of flowering plants in the borage family, Boraginaceae. It contains about 300 species of shrubs and trees, that ...

Ehretiaceae Ehretioideae is a subfamily of the flowering plant family Boraginaceae. Genera *''Bourreria'' P.Browne *'' Cortesia'' Cav. *''Ehretia'' P.Browne *''Halgania'' Gaudich. *'' Ixorhea'' Fenzl *'' Lepidocordia'' Ducke *'' Menais'' Loefl. *'' Pa ...

, and

Heliotropiaceae Heliotropiaceae is a cosmopolitan family of flowering plants with approximately 450 species worldwide, though it is concentrated especially in the tropics and subtropics. There are the following four genera in the family: * ''Heliotropium (''incl. ...

) and

Lennoaceae Lennooideae is a subfamily of parasitic flowering plants of southwestern North America and northwestern South America. The relationships of this subfamily to other plants remain uncertain. It was traditionally treated at family rank as Lennoacea ...

were placed in the order

Lamiales The order Lamiales (also known as the mint order) are an order in the asterid group of dicotyledonous flowering plants. It includes about 23,810 species, 1,059 genera, and is divided into about 25 families. These families include Acanthaceae, Bi ...

, while the related Hydrophyllaceae was placed in

Solanales The Solanales are an order of flowering plants, included in the asterid group of dicotyledons. Some older sources used the name Polemoniales for this order. Taxonomy Under the older Cronquist system, the latter three families were placed else ...

. The APG system took a broad view of Boraginaceae (Boraginaceae ''

s.l. SL may refer to: Arts and entertainment * SL (rapper), a rapper from London * ''Second Life'', a multi-user 3D virtual world * Sensei's Library, an Internet site dedicated to the game of Go * Subdominant leittonwechselklänge * Leica SL, a mirror ...

''), including within it the traditionally recognized families Hydrophyllaceae and

based on recent

molecular phylogenies Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

that show that Boraginaceae, as traditionally defined, is

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

over these two families.

APG III APG is an abbreviation with several different meanings: * Aberdeen Proving Ground, a United States Army installation in Aberdeen, Maryland, also ** Phillips Army Airfield, the airfield of the above, from its IATA airport code * Aboriginal Provisiona ...

included Boraginaceae in the

Euasterid In the APG IV system (2016) for the classification of flowering plants, the name asterids denotes a clade (a monophyletic group). Asterids is the largest group of flowering plants, with more than 80,000 species, about a third of the total floweri ...

I (

lamiid In the APG IV system (2016) for the classification of flowering plants, the name asterids denotes a clade (a monophyletic group). Asterids is the largest group of flowering plants, with more than 80,000 species, about a third of the total flowe ...

)

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

but this family was otherwise unplaced; its precise relationship to other families in the Euasterid I group remained unclear. In a

phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

study Study or studies may refer to: General * Education **Higher education * Clinical trial * Experiment * Observational study * Research * Study skills, abilities and approaches applied to learning Other * Study (art), a drawing or series of drawi ...

DNA sequence DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...

s of selected

gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...

s, Boraginales was resolved as

sister A sister is a woman or a girl who shares one or more parents with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a familial relationship, it is sometimes used endearingly to refer to ...

to Lamiales sensu APG, but that result had only 65%

maximum likelihood In statistics, maximum likelihood estimation (MLE) is a method of estimation theory, estimating the Statistical parameter, parameters of an assumed probability distribution, given some observed data. This is achieved by Mathematical optimization, ...

bootstrap support. In the 2016

APG IV system The APG IV system of flowering plant classification is the fourth version of a modern, mostly molecular-based, system of plant taxonomy for flowering plants (angiosperms) being developed by the Angiosperm Phylogeny Group (APG). It was publishe ...

Boraginales is an order with only one family Boraginaceae, which includes the former family Codonaceae. At the time of the APG IV consensus there was insufficient support to further divide this

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

group further. (''For a complete discussion of the history of the taxonomy of Boraginales, see '')

Boraginales Working Group

Following the publication of APG IV, a collaborative group along similar lines to the APG, the Boraginales Working Group, was formed to further elucidate the

relationships within the Boraginaceae ''s.l.''. As a result it has been split into eleven families, including: Boraginaceae '' s.s. or s.str.'',

, and Hydrophyllaceae. A number of these were

monogeneric In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...

. Boraginaceae is hard to characterize morphologically if it includes the

genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...

Codon The genetic code is the set of rules used by living cells to translate information encoded within genetic material ( DNA or RNA sequences of nucleotide triplets, or codons) into proteins. Translation is accomplished by the ribosome, which links ...

'' and '' Wellstedia''. ''Codon'' was long regarded as an unusual member of Hydrophyllaceae, but in 1998, a

molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

showed that it is closer to Boraginaceae, and both ''Codon'' and ''Wellstedia'' have been allocated to their own families, Codonaceae and Wellstediaceae. The

achlorophyllous Myco-heterotrophy (from Greek μύκης , "fungus", ἕτερος ', "another", "different" and τροφή ', "nutrition") is a symbiotic relationship between certain kinds of plants and fungi, in which the plant gets all or part of its food fro ...

holoparasite An obligate parasite or holoparasite is a parasitic organism that cannot complete its life-cycle without exploiting a suitable host. If an obligate parasite cannot obtain a host it will fail to reproduce. This is opposed to a facultative parasite, ...

s ''

Lennoa ''Lennoa'' is a monotypic genus of flowering plants belonging to the family Boraginaceae. It only contains one known species, Lennoa madreporoides Lex. It is within the subfamily of Lennoaceae. Its native range is Mexico to Venezuela. It is foun ...

'' and ''

Pholisma The genus ''Pholisma'' (Nutt. ex Hook.) consists of three to five species of desert-dwelling, primarily subterranean plants. Most notable of the genus is '' Pholisma sonorae'', native to the Southwestern United States and Mexico. The species is ...

'' were once regarded as a family,

, but it is now known that they form a

that is nested within Ehretiaceae. Some studies have indicated that Hydrophyllaceae is paraphyletic if the

tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English language, English is in the discipline of anthropology. This definition is contested, in p ...

Nameae is included within it, but further studies will be needed to resolve this issue. The inclusion of the genus '' Hoplestigma'' in Boraginales was occasionally doubted until it was strongly

confirmed In Christian denominations that practice infant baptism, confirmation is seen as the sealing of the covenant created in baptism. Those being confirmed are known as confirmands. For adults, it is an affirmation of belief. It involves laying on ...

in a

cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...

study in 2014. ''Hoplestigma'' is the closest relative of Cordiaceae and it has been recommended that the latter be expanded to include it. ''

Hydrolea ''Hydrolea'' (false fiddleleaf) is the only genus of the family Hydroleaceae of the order Solanales. Synonyms * '' H. capsularis'' synonym for ''Hydrolea spinosa'' L. * '' H. glabra'' synonym for ''Hydrolea spinosa'' L. * '' H. prostrata'' sy ...

'' was thought to belong in Hydrophyllaceae for more than a century after it was placed there by

Asa Gray Asa Gray (November 18, 1810 – January 30, 1888) is considered the most important American botanist of the 19th century. His ''Darwiniana'' was considered an important explanation of how religion and science were not necessarily mutually excl ...

, but it is now known to belong in the order

as sister to ''

Sphenoclea ''Sphenoclea'' is a genus of succulent erect annual herbs. They occur in damp habitats throughout the tropics. There are two species, '' S. zeylanica'' and '' S. pongatium''. The genus is placed alone in family Sphenocleaceae. The posi ...

''. ''

Pteleocarpa ''Pteleocarpa'' is a genus of flowering plants. The only member of the genus is the western Malesian tree ''Pteleocarpa lamponga''.David J. Mabberley. 2008. ''Mabberley's Plant-Book; a portable dictionary of plants, their classification and uses ...

'' was long regarded as an anomaly, and was usually placed in Boraginales, but with considerable doubt. The

molecular A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bioche ...

evidence Evidence for a proposition is what supports this proposition. It is usually understood as an indication that the supported proposition is true. What role evidence plays and how it is conceived varies from field to field. In epistemology, evidenc ...

strongly supports it as sister to

Gelsemiaceae Gelsemiaceae is a family of flowering plants, belonging to the order Gentianales. The family contains only three genera: ''Gelsemium'', ''Mostuea'' and ''Pteleocarpa''. ''Gelsemium'' has three species, one native to Southeast Asia and southern C ...

, and that family has been expanded to include it.

References

Bibliography

* * * * * * * * * Diane, N., H. Förther, and H. H. Hilger. 2002. A systematic analysis of ''Heliotropium'', ''Tournefortia'', and allied taxa of the Heliotropiaceae (Boraginales) based on ITS1 sequences and morphological data. ''American Journal of Botany'' 89: 287-295 (online abstrac
here
. * Gottschling, M., H. H. Hilger 1, M. Wolf 2, N. Diane. 2001. Secondary Structure of the ITS1 Transcript and its Application in a Reconstruction of the Phylogeny of Boraginales. ''Plant Biology (Stuttgart)'' 3: 629-636 (abstract onlin
here

Historical sources

* * *

External links

Distribution MapMissouri Botanical Garden
{{Taxonbar, from=Q2538910 Asterids Angiosperm orders