# Boraginoideae > Mediated Wiki article. Canonical URL: https://mediated.wiki/source/Boraginoideae > Markdown URL: https://mediated.wiki/source/Boraginoideae.md > Source: https://en.wikipedia.org/wiki/Boraginoideae > Source revision: 1216177942 > License: Creative Commons Attribution-ShareAlike 4.0 International (https://creativecommons.org/licenses/by-sa/4.0/) Subfamily of plants within the borage family (Boraginaceae) Boraginoideae Borago officinalis Scientific classification Kingdom: Plantae Clade: Embryophytes Clade: Tracheophytes Clade: Spermatophytes Clade: Angiosperms Clade: Eudicots Clade: Asterids Order: Boraginales Family: Boraginaceae Subfamily: Boraginoideae Arn. Type genus Borago L. Tribes Boragineae Lithospermeae **Boraginoideae** is a [subfamily](/source/Subfamily) of the plant family [Boraginaceae](/source/Boraginaceae) *s.s*, with about 42 genera. That family is [defined](/source/Circumscription_(taxonomy)) in a much broader sense (Boraginaceae *s.l.*) in the [Angiosperm Phylogeny Group](/source/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](/source/Taxonomist) placed the genera *[Codon](/source/Codon_(plant))* and *[Wellstedia](/source/Wellstedia)* in Boraginoideae. Others place one or both of these in separate, [monogeneric](/source/Monogeneric) subfamilies. *Codon* was long regarded as an odd member of [Hydrophylloideae](/source/Hydrophylloideae), but in 1998, a [molecular phylogenetic](/source/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](/source/Order_(biology))) within the [lamiid](/source/Lamiid) clade of [eudicots](/source/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](/source/Boraginales).[3][2] In that system Boraginaceae is treated in a narrow sense (*[sensu stricto](/source/Sensu_stricto)* or *s.s.*). Subsequently the order Boraginales was added to the 2016 revision ([APG IV](/source/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](/source/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](https://en.wikipedia.org/wiki/Wikipedia:Citation_needed)*] ### Subdivision Comparisons of [DNA sequences](/source/DNA_sequence) by [cladistic](/source/Cladistic) methods suggested the division of Boraginoideae into four [tribes](/source/Tribe_(biology)): 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](/source/Germplasm_Resources_Information_Network)[7] - *[Actinocarya](/source/Actinocarya)* Benth. - *[Adelocaryum](/source/Adelocaryum)* Brand - *[Afrotysonia](/source/Afrotysonia)* Rauschert - *[Alkanna](/source/Alkanna)* Tausch - *[Amblynotus](/source/Amblynotus)* (A.DC.) I. M. Johnst. - *[Amphibologyne](/source/Amphibologyne)* Brand - *[Amsinckia](/source/Amsinckia)* Lehm. - *[Anchusa](/source/Anchusa)* L. - *[Ancistrocarya](/source/Ancistrocarya)* Maxim. - *[Anoplocaryum](/source/Anoplocaryum)* Ledeb. - *[Antiotrema](/source/Antiotrema)* Hand.-Mazz. - *[Antiphytum](/source/Antiphytum)* DC. ex [Meisn.](/source/Carl_Meissner) - *[Arnebia](/source/Arnebia)* Forssk. - *[Asperugo](/source/Asperugo)* L. - *[Auxemma](/source/Auxemma)* Miers - *[Borago](/source/Borago)* L. - *[Bothriospermum](/source/Bothriospermum)* Bunge - *[Brachybotrys](/source/Brachybotrys)* Maxim. ex Oliv. - *[Brunnera](/source/Brunnera)* Steven - *[Buglossoides](/source/Buglossoides)* Moench - *[Caccinia](/source/Caccinia)* Savi - *[Carmona](/source/Carmona_(plant))* Cav. - *[Cerinthe](/source/Cerinthe)* L. - *[Chionocharis](/source/Chionocharis)* I. M. Johnst. - *[Choriantha](/source/Choriantha)* Riedl - *[Codon](/source/Codon_(plant))* [L.](/source/Carl_Linnaeus) - *[Craniospermum](/source/Craniospermum)* Lehm. - *[Cryptantha](/source/Cryptantha)* Lehm. ex G.Don - *[Cynoglossopsis](/source/Cynoglossopsis)* Brand - *[Cynoglossum](/source/Cynoglossum)* L. - *[Cynoglottis](/source/Cynoglottis)* (Gusul.) Vural & Kit Tan - *[Cystostemon](/source/Cystostemon)* Balf.f. - *[Dasynotus](/source/Dasynotus)* I.M.Johnst. - *[Decalepidanthus](/source/Decalepidanthus)* Riedl - *[Echiochilon](/source/Echiochilon)* Desf. - *[Echiostachys](/source/Echiostachys)* Levyns - *[Echium](/source/Echium)* L. - *[Elizaldia](https://en.wikipedia.org/w/index.php?title=Elizaldia&action=edit&redlink=1)* Willk. - *[Embadium](/source/Embadium)* J.M.Black - *[Eritrichium](/source/Eritrichium)* Schrad. ex Gaudin - *[Gastrocotyle](/source/Gastrocotyle_(plant))* Bunge - *[Gyrocaryum](/source/Gyrocaryum)* Valdés - *[Hackelia](/source/Hackelia)* Opiz. - *[Halacsya](/source/Halacsya)* Dörfl. - *[Heliocarya](/source/Heliocarya)* Bunge - *[Heterocaryum](/source/Heterocaryum)* A.DC. - *[Huynhia](/source/Huynhia)* Greuter - *[Ivanjohnstonia](/source/Ivanjohnstonia)* Kazmi - *[Lacaitaea](https://en.wikipedia.org/w/index.php?title=Lacaitaea&action=edit&redlink=1)* Brand - *[Lappula](/source/Lappula)* Moench - *[Lasiarrhenum](https://en.wikipedia.org/w/index.php?title=Lasiarrhenum&action=edit&redlink=1)* I.M.Johnst. - *[Lasiocaryum](/source/Lasiocaryum)* I.M.Johnst. - *[Lepechiniella](/source/Lepechiniella)*Popov - *[Lindelofia](/source/Lindelofia)* Lehm. - *[Lithodora](/source/Lithodora)* Griseb. - *[Lithospermum](/source/Lithospermum)* L. - *[Lobostemon](/source/Lobostemon)* Lehm. - *[Macromeria](https://en.wikipedia.org/w/index.php?title=Macromeria&action=edit&redlink=1)* D.Don - *[Maharanga](/source/Maharanga)* A.DC. - *[Mairetis](/source/Mairetis)* I.M.Johnst. - *[Mattiastrum](/source/Mattiastrum)* (Boiss.) Brand - *[Mertensia](/source/Mertensia)* Roth - *[Metaeritrichium](https://en.wikipedia.org/w/index.php?title=Metaeritrichium&action=edit&redlink=1)* W.T.Wang - *[Microcaryum](/source/Microcaryum)* I.M.Johnst. - *[Microula](/source/Microula)* Benth. - *[Mimophytum](/source/Mimophytum)* Greenm. - *[Moltkia](/source/Moltkia_(plant))* Lehm. - *[Moltkiopsis](/source/Moltkiopsis)* I.M.Johnst. - *[Moritzia](/source/Moritzia)* DC. ex Meisn. - *[Myosotidium](/source/Myosotidium)* Hook. - *[Myosotis](/source/Myosotis)* L. - *[Neatostema](/source/Neatostema)* I.M.Johnst. - *[Nesocaryum](/source/Nesocaryum)* I.M.Johnst. - *[Nogalia](/source/Nogalia)* Verdc. - *[Nomosa](https://en.wikipedia.org/w/index.php?title=Nomosa&action=edit&redlink=1)* I.M.Johnst. - *[Nonea](/source/Nonea)* Medik. - *[Ogastemma](/source/Ogastemma)* Brummitt - *[Omphalodes](/source/Omphalodes)* Mill. - *[Omphalolappula](/source/Omphalolappula)* Brand - *[Omphalotrigonotis](/source/Omphalotrigonotis)* W.T.Wang - *[Onosma](/source/Onosma)* L. - *[Oxyosmyles](https://en.wikipedia.org/w/index.php?title=Oxyosmyles&action=edit&redlink=1)* Speg. - *[Paracaryum](/source/Paracaryum)* (A.DC.) Boiss. - *[Pardoglossum](/source/Pardoglossum)* E.Barbier & Mathez - *[Pectocarya](/source/Pectocarya)* DC. ex Meisn. - *[Pentaglottis](/source/Pentaglottis)* Tausch - *[Perittostema](https://en.wikipedia.org/w/index.php?title=Perittostema&action=edit&redlink=1)* I.M. Johnst. - *[Plagiobothrys](/source/Plagiobothrys)* Fisch. & C.A.Mey. - *[Pseudomertensia](/source/Pseudomertensia)* Riedl - *[Psilolaemus](https://en.wikipedia.org/w/index.php?title=Psilolaemus&action=edit&redlink=1)* I.M.Johnst. - *[Pulmonaria](/source/Pulmonaria)* L. - *[Rindera](/source/Rindera)* Pall. - *[Rochelia](/source/Rochelia)* Rchb. - *[Scapicephalus](https://en.wikipedia.org/w/index.php?title=Scapicephalus&action=edit&redlink=1)* Ovcz. & Czukav. - *[Selkirkia](/source/Selkirkia_(plant))* Hemsl. - *[Sericostoma](https://en.wikipedia.org/w/index.php?title=Sericostoma_(plant)&action=edit&redlink=1)* Stocks - *[Sinojohnstonia](/source/Sinojohnstonia)* Hu - *[Solenanthus](/source/Solenanthus)* Ledeb. - *[Stenosolenium](/source/Stenosolenium)* Turcz. - *[Stephanocaryum](/source/Stephanocaryum)* Popov - *[Suchtelenia](/source/Suchtelenia)* Kar. ex Meisn. - *[Symphytum](/source/Symphytum)* L. - *[Thaumatocaryon](/source/Thaumatocaryon)* Baill. - *[Thyrocarpus](/source/Thyrocarpus)* Hance - *[Tianschaniella](/source/Tianschaniella)* B.Fedtsch. ex Popov - *[Trachelanthus](/source/Trachelanthus_(plant))* Kunze - *[Trachystemon](/source/Trachystemon)* D.Don - *[Traxara](https://en.wikipedia.org/w/index.php?title=Traxara&action=edit&redlink=1)* Raf. - *[Trichodesma](/source/Trichodesma)* R.Br. - *[Trigonocaryum](/source/Trigonocaryum)* Trautv. - *[Trigonotis](/source/Trigonotis)* Steven - *[Ulugbekia](https://en.wikipedia.org/w/index.php?title=Ulugbekia&action=edit&redlink=1)* Zakirov - *[Valentiniella](https://en.wikipedia.org/w/index.php?title=Valentiniella&action=edit&redlink=1)* Speg. - *[Wellstedia](/source/Wellstedia)* Balf.f. ## References 1. **[^](#cite_ref-FOOTNOTEFerguson1998_1-0)** [Ferguson 1998](#CITEREFFerguson1998). 1. ^ [***a***](#cite_ref-FOOTNOTEChacon_et_al2016_2-0) [***b***](#cite_ref-FOOTNOTEChacon_et_al2016_2-1) [***c***](#cite_ref-FOOTNOTEChacon_et_al2016_2-2) [***d***](#cite_ref-FOOTNOTEChacon_et_al2016_2-3) [Chacon et al 2016](#CITEREFChacon_et_al2016). 1. **[^](#cite_ref-FOOTNOTEWeigand_et_al2014_3-0)** [Weigand et al 2014](#CITEREFWeigand_et_al2014). 1. **[^](#cite_ref-apweb_4-0)** Peter F. Stevens. (2001 onwards). "Boraginaceae". At: [Angiosperm Phylogeny Website](/source/Angiosperm_Phylogeny_Website). At: [Missouri Botanical Garden](/source/Missouri_Botanical_Garden) Website. (see *External links* below) 1. **[^](#cite_ref-FOOTNOTECohen2014_5-0)** [Cohen 2014](#CITEREFCohen2014). 1. **[^](#cite_ref-6)** Armen Takhtajan *[Flowering Plants (2009)](https://books.google.com/books?id=oumyfO-NHuUC&pg=PA542)*, p. 542, at [Google Books](/source/Google_Books) 1. **[^](#cite_ref-7)** ["Genera of *Boraginaceae* subfam. *Boraginoideae*"](https://web.archive.org/web/20150924052400/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2089). *[Germplasm Resources Information Network](/source/Germplasm_Resources_Information_Network)*. [United States Department of Agriculture](/source/United_States_Department_of_Agriculture). Archived from [the original](https://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2089) 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"](https://www.researchgate.net/publication/304454123). *[Taxon](/source/Taxon_(journal))*. **65** (3): 523–546. [doi](/source/Doi_(identifier)):[10.12705/653.6](https://doi.org/10.12705%2F653.6). [hdl](/source/Hdl_(identifier)):[2158/1062785](https://hdl.handle.net/2158%2F1062785). - Cohen, James I. (April 2014). "A phylogenetic analysis of morphological and molecular characters of Boraginaceae: evolutionary relationships, taxonomy, and patterns of character evolution". *[Cladistics](/source/Cladistics_(journal))*. **30** (2): 139–169. [doi](/source/Doi_(identifier)):[10.1111/cla.12036](https://doi.org/10.1111%2Fcla.12036). [PMID](/source/PMID_(identifier)) [34784688](https://pubmed.ncbi.nlm.nih.gov/34784688). [S2CID](/source/S2CID_(identifier)) [25399963](https://api.semanticscholar.org/CorpusID:25399963). - Ferguson, Diane M. (July 1998). "Phylogenetic Analysis and Relationships in Hydrophyllaceae Based on ndhF Sequence Data". *[Systematic Botany](/source/Systematic_Botany)*. **23** (3): 253–268. [doi](/source/Doi_(identifier)):[10.2307/2419504](https://doi.org/10.2307%2F2419504). [JSTOR](/source/JSTOR_(identifier)) [2419504](https://www.jstor.org/stable/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"](https://doi.org/10.1111%2Fcla.12061). *[Cladistics](/source/Cladistics_(journal))*. **30** (5): 508–518. [doi](/source/Doi_(identifier)):[10.1111/cla.12061](https://doi.org/10.1111%2Fcla.12061). [PMID](/source/PMID_(identifier)) [34794245](https://pubmed.ncbi.nlm.nih.gov/34794245). [S2CID](/source/S2CID_(identifier)) [11954615](https://api.semanticscholar.org/CorpusID:11954615). - [Distribution Map](http://www.mobot.org/MOBOT/Research/APweb/maps/boraginaceae.gif) And [Genus list](http://www.mobot.org/MOBOT/Research/APweb/genera/boraginaceaegen.html) At [Boraginaceae](http://www.mobot.org/MOBOT/Research/APweb/orders/boraginalesweb.htm#Boraginaceae) At [Boraginales](http://www.mobot.org/MOBOT/Research/APweb/orders/boraginalesweb.htm#Boraginales) At: [Trees](http://www.mobot.org/MOBOT/Research/APweb/treeapweb2map.html) [Archived](https://web.archive.org/web/20150521153254/http://www.mobot.org/mobot/research/apweb/treeapweb2map.html) 2015-05-21 at the [Wayback Machine](/source/Wayback_Machine) At: [APweb](http://www.mobot.org/MOBOT/Research/APweb/welcome.html) At: [Missouri Botanical Garden](http://www.missouribotanicalgarden.org/) Taxon identifiers Boraginoideae Wikidata: Q530442 Wikispecies: Boraginoideae CoL: BVBBM GRIN: 2089 iNaturalist: 795681 NCBI: 1874399 Open Tree of Life: 6061933 VASCAN: 267 --- Adapted from the Wikipedia article [Boraginoideae](https://en.wikipedia.org/wiki/Boraginoideae) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Boraginoideae?action=history)). 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