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Boraginoideae
	Borago officinalis
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Clade:	Asterids
Order:	Boraginales
Family:	Boraginaceae
Subfamily:	Boraginoideae; Arn.
Type genus
	Borago; L.
Tribes
	Boragineae; Lithospermeae;

Boraginoideae is a subfamily of the plant family Boraginaceae s.s, with about 42 genera. That family is defined in a much broader sense (Boraginaceae s.l.) in the Angiosperm Phylogeny Group (APG) system of classification for flowering plants. The APG has not specified any subfamilial structure within Boraginaceae s.l.

Taxonomy

Some taxonomists placed the genera Codon 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, but in 1998, a molecular phylogenetic study suggested that it is closer to Boraginoideae.^[1] Neither is included n more modern classifications.^[2]

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 clade of eudicots. 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.^[3]^[2] In that system Boraginaceae is treated in a narrow sense (sensu stricto 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 composition.^[4] The subfamilial structure applies solely to Boraginaceae s.s., which includes around 1,600 to 1,700 species divided into some 90 genera.^{[citation needed]}

Subdivision

Comparisons of DNA sequences by cladistic methods suggested the division of Boraginoideae into four tribes: Echiochileae, Boragineae, Lithospermeae, and Cynoglosseae.^[5] 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).^[2]

Genera

Genera and approximate number of species in the system of Chacon et al (2016):^[2]

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.^[6]

The following list of genera consists of Codon plus the genera listed for Boraginoideae at the Germplasm Resources Information Network^[7]

Actinocarya Benth.
Adelocaryum Brand
Afrotysonia Rauschert
Alkanna Tausch
Amblynotus (A.DC.) I. M. Johnst.
Amphibologyne Brand
Amsinckia Lehm.
Anchusa L.
Ancistrocarya Maxim.
Anoplocaryum Ledeb.
Antiotrema Hand.-Mazz.
Antiphytum DC. ex Meisn.
Arnebia Forssk.
Asperugo L.
Auxemma Miers
Borago L.
Bothriospermum Bunge
Brachybotrys Maxim. ex Oliv.
Brunnera Steven
Buglossoides Moench
Caccinia Savi
Carmona Cav.
Cerinthe L.
Chionocharis I. M. Johnst.
Choriantha Riedl
Codon L.
Craniospermum Lehm.
Cryptantha Lehm. ex G.Don
Cynoglossopsis Brand
Cynoglossum L.
Cynoglottis (Gusul.) Vural & Kit Tan
Cystostemon Balf.f.
Dasynotus I.M.Johnst.
Decalepidanthus Riedl
Echiochilon Desf.
Echiostachys Levyns
Echium L.
Elizaldia Willk.
Embadium J.M.Black
Eritrichium Schrad. ex Gaudin
Gastrocotyle Bunge
Gyrocaryum Valdés
Hackelia Opiz.
Halacsya Dörfl.
Heliocarya Bunge
Heterocaryum A.DC.
Huynhia Greuter
Ivanjohnstonia Kazmi
Lacaitaea Brand
Lappula Moench
Lasiarrhenum I.M.Johnst.
Lasiocaryum I.M.Johnst.
LepechiniellaPopov
Lindelofia Lehm.
Lithodora Griseb.
Lithospermum L.
Lobostemon Lehm.
Macromeria D.Don
Maharanga A.DC.
Mairetis I.M.Johnst.
Mattiastrum (Boiss.) Brand
Mertensia Roth
Metaeritrichium W.T.Wang
Microcaryum I.M.Johnst.
Microula Benth.
Mimophytum Greenm.
Moltkia Lehm.
Moltkiopsis I.M.Johnst.
Moritzia DC. ex Meisn.
Myosotidium Hook.
Myosotis L.
Neatostema I.M.Johnst.
Nesocaryum I.M.Johnst.
Nogalia Verdc.
Nomosa I.M.Johnst.
Nonea Medik.
Ogastemma Brummitt
Omphalodes Mill.
Omphalolappula Brand
Omphalotrigonotis W.T.Wang
Onosma L.
Oxyosmyles Speg.
Paracaryum (A.DC.) Boiss.
Pardoglossum E.Barbier & Mathez
Pectocarya DC. ex Meisn.
Pentaglottis Tausch
Perittostema I.M. Johnst.
Plagiobothrys Fisch. & C.A.Mey.
Pseudomertensia Riedl
Psilolaemus I.M.Johnst.
Pulmonaria L.
Rindera Pall.
Rochelia Rchb.
Scapicephalus Ovcz. & Czukav.
Selkirkia Hemsl.
Sericostoma Stocks
Sinojohnstonia Hu
Solenanthus Ledeb.
Stenosolenium Turcz.
Stephanocaryum Popov
Suchtelenia Kar. ex Meisn.
Symphytum L.
Thaumatocaryon Baill.
Thyrocarpus Hance
Tianschaniella B.Fedtsch. ex Popov
Trachelanthus Kunze
Trachystemon D.Don
Traxara Raf.
Trichodesma R.Br.
Trigonocaryum Trautv.
Trigonotis Steven
Ulugbekia Zakirov
Valentiniella Speg.
Wellstedia Balf.f.

References

^ Ferguson 1998.
^ ^a ^b ^c ^d Chacon et al 2016.
^ Weigand et al 2014.
^ Peter F. Stevens. (2001 onwards). "Boraginaceae". At: Angiosperm Phylogeny Website. At: Missouri Botanical Garden Website. (see External links below)
^ Cohen 2014.
^ Armen Takhtajan Flowering Plants (2009) , p. 542, at Google Books
^ "Genera of Boraginaceae subfam. Boraginoideae". Germplasm Resources Information Network. United States Department of Agriculture. Archived from the original on 2015-09-24. Retrieved 2009-04-02.

Bibliography

Chacón, Juliana; Luebert, Federico; Hilger, Hartmut H.; Ovchinnikova, Svetlana; Selvi, Federico; Cecchi, Lorenzo; Guilliams, C. Matt; Hasenstab-Lehman, Kristen; Sutorý, Karel; Simpson, Michael G.; Weigend, Maximilian (24 June 2016). "The borage family (Boraginaceae s.str.): A revised infrafamilial classification based on new phylogenetic evidence, with emphasis on the placement of some enigmatic genera". Taxon. 65 (3): 523–546. doi:10.12705/653.6. hdl:2158/1062785.
Cohen, James I. (April 2014). "A phylogenetic analysis of morphological and molecular characters of Boraginaceae: evolutionary relationships, taxonomy, and patterns of character evolution". Cladistics. 30 (2): 139–169. doi:10.1111/cla.12036. PMID 34784688. S2CID 25399963.
Ferguson, Diane M. (July 1998). "Phylogenetic Analysis and Relationships in Hydrophyllaceae Based on ndhF Sequence Data". Systematic Botany. 23 (3): 253–268. doi:10.2307/2419504. JSTOR 2419504.
Weigend, Maximilian; Luebert, Federico; Gottschling, Marc; Couvreur, Thomas L.P.; Hilger, Hartmut H.; Miller, James S. (October 2014). "From capsules to nutlets-phylogenetic relationships in the Boraginales". Cladistics. 30 (5): 508–518. doi:10.1111/cla.12061. PMID 34794245. S2CID 11954615.
Distribution Map And Genus list At Boraginaceae At Boraginales At: Trees Archived 2015-05-21 at the Wayback Machine At: APweb At: Missouri Botanical Garden