Boraginoideae is a

subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...

of the plant family

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 ...

', with about 42 genera. That family is

defined A definition is a statement of the meaning of a term (a word, phrase, or other set of symbols). Definitions can be classified into two large categories: intensional definitions (which try to give the sense of a term), and extensional defini ...

in a much broader sense (Boraginaceae ') in the

Angiosperm Phylogeny Group The Angiosperm Phylogeny Group (APG) is an informal international group of systematic botanists who collaborate to establish a consensus on the taxonomy of flowering plants (angiosperms) that reflects new knowledge about plant relationships disco ...

(APG) system of classification for flowering plants. The APG has not specified any subfamilial structure within Boraginaceae ''s.l.''

Taxonomy

Some

taxonomist In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given ...

s placed the genera ''

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'' in Boraginoideae. Others place one or both of these in separate,

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 ...

subfamilies. ''Codon'' was long regarded as an odd member of

Hydrophylloideae Hydrophylloideae is a subfamily of the plant family Boraginaceae. Their taxonomic position is somewhat uncertain. Traditionally, and under the Cronquist system, they were given family rank under the name Hydrophyllaceae, and treated as part of th ...

, 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 ...

study suggested that it is closer to Boraginoideae. Neither is included n more modern classifications. Some authors proposed a revision of earlier APG systems, in which Boraginaceae had been included as an unplaced family (i.e. not included in a specified order) within the

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 of

eudicots The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants mainly characterized by having two seed leaves upon germination. The term derives from Dicotyledons. Traditionally they were called tricolpates or non-magnoliid dico ...

. In that system. Boraginaceae was defined broadly (Boraginaceae ''sensu lato'' or ''s.l.''). Instead they proposed recognizing five to eight families in a separate order, the

Boraginales 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 distri ...

. In that system Boraginaceae is treated in a narrow sense (''

sensu stricto ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...

'' or ''s.s.''). Subsequently the order Boraginales was added to the 2016 revision (

APG IV 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 ...

) to include Boraginaceae at the ordinal level, in which it was the sole family. The consensus was to continue the broad usage rather than split it into separate families, based on its

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 ...

composition.Peter F. Stevens. (2001 onwards). "Boraginaceae". At:

Angiosperm Phylogeny Website The Angiosperm Phylogeny Website (or APweb) is a website dedicated to research on angiosperm phylogeny and taxonomy. The site is hosted by the Missouri Botanical Garden website and maintained by researchers, Peter F. Stevens and Hilary M. Davis ...

. At:

Missouri Botanical Garden The Missouri Botanical Garden is a botanical garden located at 4344 Shaw Boulevard in St. Louis, Missouri. It is also known informally as Shaw's Garden for founder and philanthropist Henry Shaw. Its herbarium, with more than 6.6 million spe ...

Website. (see ''External links'' below) The subfamilial structure applies solely to Boraginaceae ''s.s.'', which includes around 1,600 to 1,700 species divided into some 90 genera.

Subdivision

Comparisons of

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 by

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 ...

methods suggested the division of Boraginoideae into four

tribes 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 is in the discipline of anthropology. This definition is contested, in part due to conflic ...

: Echiochileae, Boragineae, Lithospermeae, and Cynoglosseae. This was subsequently resolved into only two tribes, Boragineae and Lithospermeae (25 genera). Boragineae was then further subdivided into two subtribes, Boragininae (15 genera) and Moritziinae (2).

Genera

Genera and approximate number of species in the system of Chacon et al (2016): * Tribe Boragineae * Subtribe Boragininae (140 species) ''Anchusa'', ''Anchusella'', ''Borago'', ''Brunnera'', ''Cynoglottis'', ''Gastrocotyle'', ''Hormuzakia'', ''Lycopsis'', ''Melanortocarya'', ''Nonea'', ''Pentaglottis'', ''Phyllocara'', ''Pulmonaria'', ''Symphytum'', ''Trachystemon'' * Subtribe Moritziinae (6 species) ''Moritzia'', ''Thaumatocaryon'' * Tribe Lithospermeae (460 species) ''Aegonychon'', ''Alkanna'', ''Ancistrocarya'', ''Arnebia'', ''Buglossoides'', ''Cerinthe'', ''Cystostemon'', ''Echiostachys'', ''Echium'', ''Glandora'', ''Halacsya'', ''Huynhia'', ''Lithodora'', ''Lithospermum'', ''Lobostemon'', ''Maharanga'', ''Mairetis'', ''Moltkia'', ''Moltkiopsis'', ''Neatostema'', ''Onosma'', ''Paramoltkia'', ''Podonosma'', ''Pontechium'', ''Stenosolenium''. The following list of genera consists of ''Codon'' plus the genera listed for Boraginoideae at the

Germplasm Resources Information Network Germplasm Resources Information Network or GRIN is an online USDA National Genetic Resources Program software project to comprehensively manage the computer database for the holdings of all plant germplasm collected by the National Plant Germplasm ...

References

Bibliography

* * * *
Distribution Map

Missouri Botanical Garden
{{Taxonbar, from=Q530442 Asterid subfamilies