Boraginaceae

Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Cladus: Core eudicots
Cladus: Asterids
Cladus: Lamiids
Ordo: Boraginales

Familia: Boraginaceae
Subfamiliae: Boraginoideae - Cordioideae - Ehretioideae - Heliotropioideae - Hydrophylloideae - Lennooideae – Namoideae – Wellstedioideae

Overview of genera

Actinocarya – Adelinia – Adelocaryum – Aegonychon – Afrotysonia – Alkanna – Amblynotus – Amphibologyne – Amsinckia – Amsinckiopsis – Anchusa – Anchusella – Ancistrocarya – Andersonglossum – Anoplocaryum – Antiotrema – Antiphytum – Arnebia – Asperugo – Austrocynoglossum – Benthamia – Borago – Bothriospermum – Bourreria – Brachybotrys – Brandella – Brunnera – Buglossoides – Caccinia – Cerinthe – Cerinthodes – Chionocharis – Choriantha – Codon – Coldenia – Cordia – Craniospermum – Crucicaryum – Cryptantha – Cynoglossopsis – Cynoglossum – Cynoglottis – Cystostemon – Dasynotus – Draperia – Echinospermum – Echiochilon – Echium – Ehretia – Ellisia – Embadium – Emmenanthe – Eriodictyon – Eritrichium – Eucrypta – Gastrocotyle – Gerascanthus – Glandora – Greeneocharis – Gyrocaryum – Hackelia – Harpagonella – Heliotropium – Hesperochiron – Heterocaryum – Hilsenbergia – Hoplestigma – Hydrophyllum – Ixorhea – Johnstonella – Kuschakewiczia – Lappula – Lasiarrhenum – Lasiocaryum – Lennoa – Lepechiniella – Lepidocordia – Lindelofia – Lithocardium – Lithodora – Lithospermum – Lobostemon – Macromeria – Maharanga – Margarospermum – Mattiastrum – Memoremea – Mertensia – Metaeritrichium – Microcaryum – Microparacaryum – Microula – Miltitzia – Mimophytum – Moltkia – Morelosia – Moritzia – Myosotidium – Myosotis – Myriopus – Nama – Neatostema – Nemophila – Nesocaryum – Nihon – Nogalia – Nomosa – Nonea – Nyctelea – Ogastemma – Omphalodes – Omphalotrigonotis – Oncaglossum – Onosma – Onosmodium – Oreocarya – Paracaryum – Paraskevia – Pardoglossum – Patagonica – Pectocarya – Pentalophus – Phacelia – Pholisma – Pholistoma – Pioctonon – Plagiobothrys – Pontechium – Pseudomertensia – Psilolaemus – Pulmonaria – Rindera – Rochefortia – Rochelia – Romanzoffia – Rotula – Saccellium – Schmidelia – Setulocarya – Simpsonanthus – Sinojohnstonia – Solenanthus – Sonnea – Stenosolenium – Symphytum – Thaumatocaryon – Thyrocarpus – Tiquilia – Tournefortia – Tricardia – Trichodesma – Trigonotis – Turricula – Varronia – Vaupelia – Viticella – Wellstedia – Wigandia
Name

Boraginaceae Juss., Gen. Pl. 128 (1789), nom. cons.

Type genus: Borago L., Sp. Pl. 1: 137. (1753)

Synonyms

Anchusaceae Vest, Anleit. Stud. Bot. 274, 302 (1818).
Cerinthaceae Bercht. & J.Presl, Přir. Rostlin: 244 (1820).
Codonaceae (Retief & A.E. van Wijk) Weigend & Hilger (2010).
Cordiaceae R.Br. ex Dumort., Anal. Fam. Pl.: 20, 25 (1829), nom. cons.
Echiaceae Raf., Fl. Tellur. 2: 61 (1837).
Ehretiaceae Mart., Nov. Gen. Sp. Pl. 2: 136, 138 (1827), nom. cons.
Heliotropiaceae Schrad., Commentat. Soc. Regiae Sci. Gott. Recent. 4: 192 [Asperifol. Linnei Comm.: 22] (1819) (Heliotropieae), nom. cons.
Hoplestigmataceae Gilg in H.G.A. Engler & E.F. Gilg, Syllabus, ed. 9-10: 322 (1924), nom. cons.
Hydrophyllaceae R.Br., Bot. Reg. 3: ad t. 242 (1817) (Hydrophylleae), nom. cons.
Lennoaceae Solms, Abh. Naturf. Ges. Halle 11: 174 (1870), nom. cons.
Onosmaceae Martinov, Tekhno-Bot. Slovar: 437 (1820) (Onosmoides).
Sagoneaceae Martinov, Tekhno-Bot. Slovar: 557 (1820) (Sagoneae).
Wellstediaceae Novák in S. Prát (ed.), Rostlinopis 9: 530 (1943)

Note: This circumscription mainly follows Boraginaceae s.l. APG IV, (2016), but also see Luebert et al., (2016) and Discussion Page for an alternative.
References
Primary references

Jussieu, A.L. de 1789. Genera plantarum, secundum ordines naturales disposita juxta methodum in Horto Regio Parisiensi exaratam. 498 pp. Paris: Herissant et Theophile Barrois. BHL Reference page.

Additional references

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 Reference page.
Chacón, J., Luebert, F., Hilger, H.H., Ovchinnikova, S., Selvi, F., Cecchi, L., Guilliams, C.M., Hasenstab-Lehman, K., Sutorý, K., Simpson, M.G. & Weigend, M. 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 ResearchGate PDF Reference page.
Cohen, J.I. 2013 (Online) 2014 (Print). 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 Reference page.
Guilliams, C.M., Hasenstab-Lehman, K.E. & Baldwin, B.G. 2020. Nomenclatural Changes in Western North American Amsinckiinae (Boraginaceae). Novon 28(1): 51–59. DOI: 10.3417/2019469 Paywall PDF Reference page.
Irimia, R-E. & Gottschling, M. 2016. Taxonomic revision of Rochefortia Sw. (Ehretiaceae, Boraginales). Biodiversity Data Journal 4: e7720. DOI: 10.3897/BDJ.4.e7720. Reference page.
Luebert, F., Cecchi, L., Frohlich, M.W., Gottschling, M., Guilliams, C.M., Hasenstab-Lehman, K.E., Hilger, H.H., Miller, J.S., Mittelbach, M., Nazaire, M. & Nepi, M. 2016. Familial classification of the Boraginales. Taxon 65(3): 502–522. DOI: 10.12705/653.5 Full text PDF from ResearchGate Reference page.
Nazaire, M. & Hufford, L. 2012. A broad phylogenetic analysis of Boraginaceae: Implications for the relationships of Mertensia. Systematic Botany 37(3): 758–783. DOI: 10.1600/036364412X648715 Reference page.
Otero, A., Jiménez-Mejías, P., Valcárcel, V. & Vargas, P. 2014: Molecular phylogenetics and morphology support two new genera (Memoremea and Nihon) of Boraginaceae s.s. Phytotaxa 173(4): 241–277. DOI: 10.11646/phytotaxa.173.4.1 Reference page.
Weigend, M. & Hilger, H.H. 2010. Codonaceae — a newly required family name in Boraginales. Phytotaxa 10: 26–30. DOI: 10.11646/phytotaxa.10.1.3 Paywall Reference page. Rejected by APG IV
Weigend, M., Luebert, F., Gottschling, M., Couvreur, T.L.P., Hilger, H.H. & Miller, J.S. 2014. From capsules to nutlets - phylogenetic relationships in the Boraginales. Cladistics 30(5): 508–518. DOI: 10.1111/cla.12061Open access Reference page. Rejected by APG IV

Links

Govaerts, R. et al. 2020. Boraginaceae in Kew Science Plants of the World online. The Board of Trustees of the Royal Botanic Gardens, Kew. Published on the internet. Accessed: 2020 Aug. 9. Reference page.
Hassler, M. 2020. Boraginaceae. World Plants: Synonymic Checklists of the Vascular Plants of the World In: Roskovh, Y., Abucay, L., Orrell, T., Nicolson, D., Bailly, N., Kirk, P., Bourgoin, T., DeWalt, R.E., Decock, W., De Wever, A., Nieukerken, E. van, Zarucchi, J. & Penev, L., eds. 2020. Species 2000 & ITIS Catalogue of Life. Published on the internet. Accessed: 2020 Aug. 9. Reference page.
International Plant Names Index. 2020. Boraginaceae. Published online. Accessed: Aug. 9 2020.
Stevens, P.F. 2001 onwards. Angiosperm Phylogeny Website. Version 14, July 2017 [and more or less continuously updated since]. Online. Reference page.
Tropicos.org 2016. Boraginaceae. Missouri Botanical Garden. Published on the internet. Accessed: 2016 Apr. 12.

Vernacular names
العربية: حمحمية
azərbaycanca: Sümürgənkimilər
беларуская: Бурачнікавыя
български: Грапаволистни
català: Boraginàcies
čeština: Brutnákovité
чӑвашла: Хăяр курăк йышĕ
dansk: Rublad-familien
Deutsch: Raublattgewächse
Esperanto: Boragacoj
eesti: Karelehelised
فارسی: گاوزبانیان
suomi: Lemmikkikasvit
Nordfriisk: Rüchbleeden
עברית: זיפניים
hrvatski: Boražinovke
hornjoserbsce: Wódrakowe rostliny
magyar: Borágófélék
հայերեն: Գաղտրիկազգիներ
íslenska: Munablómsætt
日本語: ムラサキ科
ქართული: ლაშქარასებრნი
қазақша: Айлауықтар тұқымдасы
한국어: 지치과
кыргызча: Эндиктер тукуму
lietuvių: Agurkliniai
македонски: Воловски јазичиња
മലയാളം: ബൊറാജിനേസീ
Nederlands: Ruwbladigenfamilie
norsk nynorsk: Rubladfamilien
norsk: Rubladfamilien
ирон: Джитърийымойхуызтæ
polski: Ogórecznikowate
پنجابی: والپتے
Runa Simi: Qipuncha yura rikch'aq ayllu
русский: Бурачниковые
slovenčina: Borákovité
svenska: Strävbladiga växter
ไทย: วงศ์หญ้างวงช้าง
Türkçe: Hodangiller
українська: Шорстколисті
Tiếng Việt: Họ Mồ hôi
中文: 紫草科

Boraginaceae, the borage or forget-me-not family, includes about 2,000 species of shrubs, trees and herbs in 146 genera with a worldwide distribution. [3]

The APG IV system from 2016 classifies the Boraginaceae as single family of the order Boraginales within the asterids.[4] Under the older Cronquist system it was included in Lamiales, but it is now clear that it is no more similar to the other families in this order than they are to families in several other asterid orders. A revision of the Boraginales, also from 2016, split the Boraginaceae in eleven distinct families:[5] 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.[6] 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.[7]

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.[8]

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

Actinocarya
Adelocaryum
Afrotysonia
Alkanna
Amblynotus
Amphibologyne
Amsinckia
Anchusa
Ancistrocarya
Anoplocaryum
Antiotrema
Antiphytum
Arnebia
Asperugo
Auxemma
Borago
Bothriospermum
Bourreria
Brachybotrys
Brunnera
Buglossoides
Caccinia
Carmona
Cerinthe
Chionocharis
Choriantha
Cordia
Craniospermum
Cryptantha
Cynoglossopsis
Cynoglossum
Cynoglottis
Cysostemon
Dasynotus
Decalepidanthus
Echiochilon
Echiostachys
Echium
Ehretia
Elizaldia
Embadium
Emmenanthe
Eritrichium
Euploca
Gastrocotyle
Gyrocaryum
Hackelia
Halacsya
Halgania
Harpagonella
Heliocarya
Heliotropium
Heterocaryum
Huynhia
Ivanjohnstonia
Ixorhea
Johnstonella
Lacaitaea
Lappula
Lasiarrhenum
Lasiocaryum
Lennoa
Lepechiniella
Lepidocordia
Lindelophia
Lithodora
Lithospermum
Lobostemon
Macromeria
Maharanga
Mairetis
Mattiastrum
Mertensia
Metaeritrichium
Microcaryum
Microula
Mimophytum
Moltkia
Moltkiopsis
Moritzia
Myosotidium
Myosotis
Neatostema
Nesocaryum
Nogalia
Nomosa
Nonea
Ogastemma
Omphalodes
Omphalolappula
Omphalotrigonotis
Onosma
Oxyosmyles
Paracaryum
Pardoglossum
Patagonula
Pectocarya
Pentaglottis
Perittostema
Phacelia
Plagiobothrys
Pseudomertensia
Psilolaemus
Pulmonaria
Rindera
Rochefortia
Rochelia
Rotula
Saccellium
Scapicephalus
Sericostoma
Sinojohnstonia
Solenanthus
Stenosolenium
Stephanocaryum
Suchtelenia
Symphytum
Thaumatocaryum
Thyrocarpus
Tianschaniella
Tiquilia
Tournefortia
Trachelanthus
Trachystemon
Trichodesma
Trigonocaryum
Trigonotis
Ulugbekia
Valentiniella

References

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.
"Boraginaceae Juss., nom. cons". Germplasm Resources Information Network. United States Department of Agriculture. 2007-04-12. Retrieved 2009-04-02.
Boraginaceae. Archived 2015-09-23 at the Wayback Machine Diversityoflife.com
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. open access
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. ISSN 0040-0262. Retrieved 16 June 2018.
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.
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.

Hess, D. 2005. Systematische Botanik. ISBN 3-8252-2673-5

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).