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Boraginaceae
Borage (Borago officinalis)
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Clade:	Asterids
Order:	Boraginales
Family:	Boraginaceae Juss.
Subfamilies
Boraginoideae Cordioideae Ehretioideae Hydrophylloideae Lennooideae

Boraginaceae, the borage or forget-me-not family, includes about 2,000 species of shrubs, trees, and herbs in 146 to 154 genera with a worldwide distribution.

The APG IV system from 2016 classifies the Boraginaceae as single family of the order Boraginales within the asterids. Under the older Cronquist system, it was included in the Lamiales, but clearly is no more similar to the other families in this order than it is to families in several other asterid orders. A revision of the Boraginales, also from 2016, split the Boraginaceae into 11 distinct families: Boraginaceae sensu stricto, Codonaceae, Coldeniaceae, Cordiaceae, Ehretiaceae, Heliotropiaceae, Hoplestigmataceae, Hydrophyllaceae, Lennoaceae, Namaceae, and Wellstediaceae.

These plants have alternately arranged leaves, or a combination of alternate and opposite leaves. The leaf blades usually have a narrow shape; many are linear or lance-shaped. They are smooth-edged or toothed, and some have petioles. Most species have bisexual flowers, but some taxa are dioecious. Most pollination is by hymenopterans, such as bees. Most species have inflorescences that have a coiling shape, at least when new, called scorpioid cymes. The flower has a usually five-lobed calyx. The corolla varies in shape from rotate to bell-shaped to tubular, but it generally has five lobes. It can be green, white, yellow, orange, pink, purple, or blue. There are five stamens and one style with one or two stigmas. The fruit is a drupe, sometimes fleshy.

Most members of this family have hairy leaves. The coarse character of the hairs is due to cystoliths of silicon dioxide and calcium carbonate. These hairs can induce an adverse skin reaction, including itching and rash in some individuals, particularly among people who handle the plants regularly, such as gardeners. In some species, anthocyanins cause the flowers to change color from red to blue with age. This may be a signal to pollinators that a flower is old and depleted of pollen and nectar.

Well-known members of the family include:

• alkanet (Alkanna tinctoria)
• borage (Borago officinalis)
• comfrey (Symphytum spp.)
• fiddleneck (Amsinckia spp.)
• forget-me-not (Myosotis spp.)
• geigertree (Cordia sebestena)
• green alkanet (Pentaglottis sempervirens)
• heliotrope (Heliotropium spp.)
• hound's tongue (Cynoglossum spp.)
• lungwort (Pulmonaria spp.)
• oysterplant (Mertensia maritima)
• purple viper's bugloss/Salvation Jane (Echium plantagineum)
• Siberian bugloss (Brunnera macrophylla)
• viper's bugloss (Echium vulgare)

Genera

According to Kew;

• Actinocarya Benth.
• Adelinia J.I.Cohen
• Adelocaryum Brand
• Aegonychon Gray
• Afrotysonia Rauschert
• Ailuroglossum Sutory
• Alkanna Tausch
• Amblynotus (A.DC.) I.M.Johnst.
• Amphibologyne Brand
• Amsinckia Lehm.
• Amsinckiopsis (I.M.Johnst.) Guilliams, Hasenstab & B.G.Baldwin
• Anchusa L.
• Ancistrocarya Maxim.
• Andersonglossum J.I.Cohen
• Anoplocaryum Ledeb.
• Antiotrema Hand.-Mazz.
• Antiphytum DC. ex Meisn.
• Arnebia Forssk.
• Asperugo L.
• Borago L.
• Bothriospermum Bunge
• Bourreria P.Browne
• Brachybotrys Maxim. ex Oliv.
• Brandella R.R.Mill
• Brunnera Steven
• Buglossoides Moench
• Caccinia Savi
• Cerinthe L.
• Chionocharis I.M.Johnst.
• Codon L.
• Coldenia L.
• Cordia L.
• Craniospermum Lehm.
• Crucicaryum Brand
• Cryptantha Lehm. ex G.Don
• Cynoglossopsis Brand
• Cynoglossum L.
• Cynoglottis (Guşul.) Vural & Kit Tan
• Cystostemon L.
• Dasynotus I.M.Johnst.
• Decalepidanthus Reidl
• Draperia Torr.
• Echiochilon Desf.
• Echium Tourn. ex L.
• Ehretia P.Browne
• Ellisia L.
• Embadium J.M.Black
• Emmenanthe Benth.
• Eremocarya Greene
• Eriodictyon Benth.
• Eritrichium Schrad. ex Gaudin
• Eucrypta Nutt.
• Euploca Nutt.
• Gastrocotyle Bunge
• Glandora D.C.Thomas, Weigend & Hilger
• Greeneocharis Gürke & Harms
• Gyrocaryum Valdés
• Hackelia Opiz
• Halacsya Dörfl.
• Halgania Gaudich.
• Harpagonella A.Gray
• Heliocarya Bunge
• Heliotropium Tourn. ex L.
• Hesperochiron S.Watson
• Hoplestigma Pierre
• Hormuzakia Guşul.
• Huynhia Greuter
• Hydrophyllum L.
• Iberodes M.Serrano, R.Carbajal & S.Ortiz
• Ivanjohnstonia Kazmi
• Ixorhea Fenzl
• Johnstonella Brand
• Keraunea Cheek & Sim.-Bianch.
• Lappula Moench
• Lasiocaryum I.M.Johnst.
• Lennoa Lex.
• Lepechiniella Popov
• Lepidocordia Ducke
• Lindelofia Lehm.
• Lithodora Griseb.
• Lithospermum L.
• Lobostemon Lehm.
• Maharanga DC.
• Mairetis I.M.Johnst.
• Mattiastrum (Boiss.) Brand
• Megacaryon Boiss.
• Melanortocarya Selvi, Bigazzi, Hilger & Papini
• Memoremea A.Otero, Jim.Mejías, Valcárcel & P.Vargas
• Mertensia Roth
• Microcaryum I.M.Johnst.
• Microparacaryum (Popov ex Riedl) Hilger & Podlech
• Microula Benth.
• Mimophytum Greenm.
• Moltkia Lehm.
• Moltkiopsis I.M.Johnst.
• Moritzia DC. ex Meisn.
• Myosotidium Hook.
• Myosotis L.
• Myriopus Small
• Nama L.
• Neatostema I.M.Johnst.
• Nemophila Nutt. ex W.P.C.Barton
• Nesocaryum I.M.Johnst.
• Nihon A.Otero, Jim.Mejías, Valcárcel & P.Vargas
• Nogalia Verdc.
• Nonea Medik.
• Ogastemma Brummitt
• Omphalodes Mill.
• Omphalolappula Brand
• Omphalotrigonotis W.T.Wang
• Oncaglossum Sutory
• Onosma L.
• Oreocarya Greene
• Paracaryum Boiss.
• Paramoltkia Greuter
• Pectocarya DC. ex Meisn.
• Pentaglottis Tausch
• Phacelia Juss.
• Pholisma Nutt. ex Hook.
• Pholistoma Lilja
• Phyllocara Guşul.
• Plagiobothrys Fisch. & C.A.Mey.
• Podonosma Boiss.
• Pontechium Böhle & Hilger
• Pseudolappula Khoshsokhan & Kaz.Osaloo
• Pulmonaria L.
• Rindera Pall.
• Rochefortia Sw.
• Rochelia Rchb.
• Romanzoffia Cham.
• Rotula Lour.
• Sauria Bajtenov
• Selkirkia Hemsl.
• Simpsonanthus Guilliams, Hasenstab & B.G.Baldwin
• Sinojohnstonia Hu
• Solenanthus Ledeb.
• Stenosolenium Turcz.
• Suchtelenia Kar. ex Meisn.
• Symphytum Tourn. ex L.
• Thaumatocaryon Baill.
• Thyrocarpus Hance
• Tianschaniella B.Fedtsch. ex Popov
• Tiquilia Pers.
• Tournefortia L.
• Trachelanthus Kunze
• Trachystemon D.Don
• Tricardia Torr. ex S.Watson
• Trichodesma R.Br.
• Trigonocaryum Trautv.
• Trigonotis Steven
• Turricula J.F.Macbr.
• Varronia P.Browne
• Wellstedia Balf.f
• Wigandia Kunth

References

1. Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x. hdl:10654/18083.
2. "Boraginaceae Juss., nom. cons". Germplasm Resources Information Network. United States Department of Agriculture. 2007-04-12. Retrieved 2009-04-02.
3. Boraginaceae. Archived 2015-09-23 at the Wayback Machine Diversityoflife.com
4. ^ "Boraginaceae Juss. | Plants of the World Online | Kew Science". Plants of the World Online. Retrieved 8 July 2024.
5. Angiosperm Phylogeny Group (2016). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV". Botanical Journal of the Linnean Society. 181 (1): 1–20. doi:10.1111/boj.12385.
6. Luebert, F.; Cecchi, L.; Frohlich, M.W.; et al. (2016). "Familial classification of the Boraginales". Taxon. 65 (3): 502–522. doi:10.12705/653.5. hdl:2158/1062790. ISSN 0040-0262. Retrieved 16 June 2018.
7. Buys, Matt H.; Hilger, Hartmut H. (2003). "Boraginaceae Cymes Are Exclusively Scorpioid and Not Helicoid". Taxon. 52 (4): 719–724. doi:10.2307/3647346. ISSN 0040-0262. JSTOR 3647346.
8. Watson, L. and M. J. Dallwitz. 1992 onwards. Boraginaceae Juss. Archived July 1, 2005, at the Wayback Machine The Families of Flowering Plants. Version: 19 August 2013.
9. Hess, D. 2005. Systematische Botanik. ISBN 3-8252-2673-5

Further reading

• Diane, N., et al. 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(2), 287-95.
• Gottschling, M., et al. (2001). Secondary structure of the ITS1 transcript and its application in a reconstruction of the phylogeny of Boraginales. Plant Biology 3, 629-36.

External links

• Distribution Map And Genus list At Boraginaceae At Boraginales At: Trees At: APweb At: Missouri Botanical Garden
• Boraginaceae In:Mabberley's Plant-Book
• Boraginaceae At: Plant Names At: IPNI
• Home page of James L. Reveal and C. Rose Broome
• Boraginaceae (Search Exact) At Name Search At: Tropicos At: Missouri Botanical Garden
• Boraginaceae. Topwalks.net: Walking Routes in Spain.
• Boraginaceae. Integrated Taxonomic Information System (ITIS).

• Boraginaceae
• Asterid families