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 botanical subfamily names with "-oideae", and zo ...

of the plant family

Boraginaceae Boraginaceae, the Borago, borage or forget-me-not family, includes about 2,000 species of shrubs, trees, and herbs in 146 to 154 genus, genera with a worldwide distribution. The APG IV system from 2016 classifies the Boraginaceae as single famil ...

', with about 42 genera. That family is defined 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 disc ...

(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 science, scientific study of naming, defining (Circumscription (taxonomy), circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxon, taxa (si ...

s placed the genera ''

Codon Genetic code is a 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 prote ...

'' and '' Wellstedia'' in Boraginoideae. Others place one or both of these in separate, monogeneric subfamilies. ''Codon'' was long regarded as an odd member of

Hydrophylloideae Hydrophylloideae is a subfamily of the plant family Boraginaceae. Its taxonomic position is somewhat uncertain. Traditionally, and under the Cronquist system, the taxon was given family rank under the name Hydrophyllaceae, and treated as part of ...

, 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 Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...

) within the lamiid clade of

eudicots The eudicots or eudicotyledons are flowering plants that have two seed leaves (cotyledons) upon germination. The term derives from ''dicotyledon'' (etymologically, ''eu'' = true; ''di'' = two; ''cotyledon'' = seed leaf). Historically, authors h ...

. 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. 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 c ...

'' or ''s.s.''). Subsequently the order Boraginales was added to the 2016 revision ( APG IV) 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 biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

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

Angiosperm Phylogeny Website The Angiosperm Phylogeny Website (or APweb) is a website that presents up-to-date research on the phylogeny and taxonomy of flowering plants (angiosperms) in what is intended to be a user-friendly way. The site is hosted by the Missouri Botanical ...

. 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 philanthropy, philanthropist Henry Shaw (philanthropist), Henry Shaw. I ...

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 A nucleic acid sequence is a succession of bases within the nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. This succession is denoted by a series of a set of five different letters that indicate the order of the nu ...

s by

cladistic Cladistics ( ; from Ancient Greek 'branch') 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 ...

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 use of the term in English is in the discipline of anthropology. The definition is contested, in part due to conflict ...

: 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 Germpla ...

References

Bibliography

* * * *
Distribution Map

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