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Linaria
	Common toadflax (Linaria vulgaris)
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Clade:	Asterids
Order:	Lamiales
Family:	Plantaginaceae
Tribe:	Antirrhineae
Genus:	Linaria; Mill.
Species
	See text
Synonyms
	Elatine Hill; Peloria Adans.; Saccularia Kellogg; Trimerocalyx (Murb.) Murb.;

Linaria is a genus of almost 200 species of flowering plants, one of several related groups commonly called toadflax. They are annuals and herbaceous perennials, and the largest genus in the Antirrhineae tribe of the plantain family Plantaginaceae.

Taxonomy

Linaria was traditionally placed in the family Scrophulariaceae. Phylogenetic analysis has now placed it in the vastly expanded family Plantaginaceae.

Closely related genera include Nuttallanthus (American toadflaxes, recently split from Linaria), Antirrhinum (snapdragons) and Cymbalaria (ivy-leaved toadflaxes).

Cultivation

Several Linaria species are cultivated as garden plants, and some are regarded as having a weedy habit.

Common toadflax or butter-and-eggs (Linaria vulgaris), a European species which is widely introduced elsewhere and grows as a common weed in some areas.^[1]
Broomleaf toadflax or Dalmatian toadflax (Linaria genistifolia, syn. L. dalmatica), a native of southeast Europe that has become a weed in parts of North America.^[2]
Purple toadflax (Linaria purpurea), a species native to the Mediterranean region grown as a garden plant for its dark purple or pink flowers. The version with purple flowers can be mistaken for lavender. Spreads readily.
Pale toadflax (Linaria repens), a species from western Europe similar to L. purpurea, but with paler flowers.
Alpine toadflax (Linaria alpina), purple flowers with orange (or purple) lobes in the center.
Moroccan toadflax (Linaria maroccana), the flower has five lobes arranged into two lips with a spur at the end, often purple with white.

Species

The following species are recognised in the genus Linaria:^[3]

Linaria accitensis L.Sáez, Juan, M.B.Crespo, F.B.Navarro, J.Peñas & Roquet
Linaria acutiloba Fisch.
Linaria aeruginea (Gouan) Cav.
Linaria afghanica Podlech & Iranshahr
Linaria alaica Junussov
Linaria albifrons (Sm.) Spreng.
Linaria algarviana Chav.
Linaria alpina (L.) Mill.
Linaria altaica Fisch.
Linaria amethystea (Vent.) Hoffmanns. & Link
Linaria amoi Campo ex Amo
Linaria angustissima (Loisel.) Borbás
Linaria antilibanotica Rech.f.
Linaria arcusangeli Atzei & Camarda
Linaria arenaria DC.
Linaria arenicola Pau & Font Quer
Linaria argillicola Juan, Blanca, Cueto, J.Fuentes & L.Sáez
Linaria armeniaca Chav.
Linaria arvensis (L.) Desf.
Linaria atlantica Boiss. & Reut.
Linaria azerbaijanensis Hamdi & Assadi
Linaria badachschanica Junussov
Linaria badalii Willk.
Linaria bamianica Patzak
Linaria becerrae Blanca, Cueto & J.Fuentes
Linaria bessarabica Kotov
Linaria biebersteinii Besser
Linaria bipartita (Vent.) Willd.
Linaria bipunctata (L.) Dum.Cours.
Linaria bordiana Santa & Simonn.
Linaria boushehrensis Hamdi & Assadi
Linaria brachyphylla Delip.
Linaria bubanii Font Quer
Linaria bungei Kuprian.
Linaria buriatica Turcz. ex Ledeb.
Linaria caesia (Lag. ex Pers.) F.Dietr.
Linaria capraria Moris & De Not.
Linaria cavanillesii Chav.
Linaria chalepensis (L.) Mill.
Linaria clementei Haens. ex Boiss.
Linaria confertiflora Benth.
Linaria corifolia Desf.
Linaria × cornubiensis Druce
Linaria cossoniana Braun-Blanq. & Maire
Linaria cossonii Bonnet & Barratte
Linaria cretacea Fisch. ex Spreng.
Linaria dalmatica (L.) Mill.
Linaria damascena Boiss. & Gaill.
Linaria decipiens Batt.
Linaria depauperata Leresche ex Lange
Linaria diffusa Hoffmanns. & Link
Linaria dissita Pomel
Linaria × dominii Druce
Linaria dumanii A.Duran & Menemen
Linaria elegans Cav.
Linaria elymaitica (Boiss.) Kuprian.
Linaria fallax Coss. ex Batt.
Linaria farsensis Hamdi & Assadi
Linaria fastigiata Chav.
Linaria faucicola Leresche & Levier
Linaria fedorovii Kamelin
Linaria ficalhoana Rouy
Linaria flava (Poir.) Desf.
Linaria genistifolia (L.) Mill.
Linaria gharbensis Batt. & Pit.
Linaria glacialis Boiss.
Linaria glauca (L.) Cav.
Linaria golestanensis Hamdi & Assadi
Linaria grandiflora Desf.
Linaria griffithii Benth.
Linaria grjunerae Knjaz.
Linaria guilanensis Hamdi & Assadi
Linaria haelava (Forssk.) Delile
Linaria hepatica Bunge
Linaria heratensis Podlech & Iranshahr
Linaria hirta (L.) Moench
Linaria hohenackeri Tzvelev
Linaria huteri Lange
Linaria × hybrida Schur
Linaria iconia Boiss. & Heldr.
Linaria ikonnikovii Stasiak
Linaria imzica Gómiz
Linaria incarnata (Vent.) Spreng.
Linaria incompleta Kuprian.
Linaria intricata Coincy
Linaria iranica Hamdi & Assadi
Linaria × jalancina Gómez Nav., R.Roselló, A.Guillén, P.P.Ferrer, E.Laguna & Per
Linaria japonica Miq.
Linaria jaxartica Levichev
Linaria joppensis Bornm.
Linaria kavirensis Hamdi & Assadi
Linaria khalkhalensis Hamdi & Assadi
Linaria khorasanensis Hamdi & Assadi
Linaria × kocianovichii Asch.
Linaria kokanica Regel
Linaria kulabensis B.Fedtsch.
Linaria kurdica Boiss. & Hohen.
Linaria latifolia Desf.
Linaria laxiflora Desf.
Linaria leptoceras Kuprian.
Linaria lineolata Boiss.
Linaria loeselii Schweigg.
Linaria longicalcarata D.Y.Hong
Linaria macrophylla Kuprian.
Linaria macroura (M.Bieb.) M.Bieb.
Linaria maroccana Hook.f.
Linaria maymanica Wendelbo
Linaria mazandaranensis Hamdi & Assadi
Linaria melampyroides Kuprian.
Linaria melanogramma Rech.f., Aellen & Esfand.
Linaria meyeri Kuprian.
Linaria michauxii Chav.
Linaria micrantha (Cav.) Hoffmanns. & Link
Linaria microsepala A.Kern.
Linaria multicaulis (L.) Mill.
Linaria munbyana Boiss. & Reut.
Linaria musilii Velen.
Linaria nachitschevanica Tzvelev
Linaria nigricans Lange
Linaria nivea Boiss. & Reut.
Linaria nurensis Boiss. & Hausskn.
Linaria nuristanica Patzak
Linaria oblongifolia (Boiss.) Boiss. & Reut.
Linaria odora (M.Bieb.) Fisch.
Linaria oligantha Lange
Linaria × oligotricha Borbás
Linaria onubensis Pau
Linaria orbensis Carretero & Boira
Linaria ordubadica Tzvelev
Linaria pamirica (Junussov) Stasiak
Linaria paradoxa Murb.
Linaria parviracemosa D.A.Sutton
Linaria pedicellata Kuprian.
Linaria pedunculata (L.) Chaz.
Linaria pelisseriana (L.) Mill.
Linaria peloponnesiaca Boiss. & Heldr.
Linaria peltieri Batt.
Linaria pinifolia (Poir.) Thell.
Linaria platycalyx Boiss.
Linaria polygalifolia Hoffmanns. & Link
Linaria popovii Kuprian.
Linaria propinqua Boiss. & Reut.
Linaria pseudolaxiflora Lojac.
Linaria pseudoviscosa Murb.
Linaria purpurea (L.) Mill.
Linaria pyramidalis (Vent.) F.Dietr.
Linaria qartobensis Blanca, Cueto, J.Fuentes, L.Sáez & Tarifa
Linaria quasisessilis Levichev
Linaria reflexa (L.) Desf.
Linaria remotiflora Patzak
Linaria repens (L.) Mill.
Linaria ricardoi Cout.
Linaria riffea Pau
Linaria × rocheri P.Fourn.
Linaria rubioides Vis. & Pancic
Linaria sabulosa Czern. ex Klokov
Linaria salangensis Podlech & Iranshahr
Linaria salzmannii Boiss.
Linaria saposhnikovii Nikitina
Linaria saturejoides Boiss.
Linaria saxatilis (L.) Chaz.
Linaria schelkownikowii Schischk.
Linaria schirvanica Fomin
Linaria semialata D.López, Sánchez-Gómez, J.F.Jiménez, J.B.Vera & Güemes
Linaria × sepium G.J.Allman
Linaria sessilis Kuprian.
Linaria simplex Desf.
Linaria spartea (L.) Chaz.
Linaria striatella Kuprian.
Linaria supina (L.) Chaz.
Linaria tarhunensis Pamp.
Linaria tenuis (Viv.) Spreng.
Linaria thibetica Franch.
Linaria thymifolia (Vahl) DC.
Linaria tingitana Boiss. & Reut.
Linaria tonzigii Lona
Linaria triornithophora (L.) Cav.
Linaria triphylla (L.) Mill.
Linaria tristis (L.) Mill.
Linaria tursica Valdés & Cabezudo
Linaria unaiensis Patzak
Linaria × valdesiana Socorro & Aroza
Linaria venosa Lindl.
Linaria ventricosa Coss. & Balansa
Linaria veratrifolia Patzak
Linaria verticillata Boiss.
Linaria virgata (Poir.) Desf.
Linaria viscosa (L.) Dum.Cours.
Linaria volgensis Rakov & Tzvelev
Linaria vulgaris Mill.
Linaria warionis Pomel
Linaria weilleri Emb. & Maire
Linaria yunnanensis W.W.Sm.
Linaria yusufeliensis A.Galán, Makbul & Hamzaoglu
Linaria × zaborskiana Emb.
Linaria zaissanica Semiotr.

Etymology

The members of this genus are known in English as toadflax, a name shared with several related genera. The 'toad' in toadflax may relate to the plants having historically been used to treat bubonic plague, a false link having been drawn between the words 'bubo' and 'Bufo'. The scientific name Linaria means "resembling linum" (flax), which the foliage of some species superficially resembles.

Distribution and habitat

The genus is native to temperate regions of Europe, northern Africa and Asia, with the highest species diversity in the Mediterranean region.

Ecology

Some Linaria are regarded as noxious weeds. They are likely toxic to livestock, but ruminants generally avoid them.^[4]

Chemical composition

Linaria species are rich in alkaloids, iridoids, terpenes, phenolic acids and flavonoids.

Vasicine, vasicinone, 7-hyrdoxyvasicine, linarinic acid, choline, linavuline, luteolin, acacetin, apigenin, chrysin, quercetin, myricetin, linarioside, aucubin, linaride, iridolinaroside A-D, and iridolinarin A-C are some compounds found in plants of this genus.^[5]

Uses

Toadflaxes are used as food plants by the larvae of some Lepidoptera species, including the mouse moth (Amphipyra tragopoginis) and the common buckeye (Junonia coenia).

Traditional medicine

Linaria vulgaris has been used as a medicinal herb.^[6]

References

^ Brickell, Christopher, ed. (2008). The Royal Horticultural Society A-Z Encyclopedia of Garden Plants. United Kingdom: Dorling Kindersley. p. 636. ISBN 9781405332965.
^ Dalmatian Toadflax (Linaria dalmatica). National Invasive Species Information Center, United States National Agricultural Library.
^ "Linaria Mill. | Plants of the World Online | Kew Science". Plants of the World Online. Retrieved 2022-07-04.
^ Sing, S. E. and R. K. Peterson. (2011). Assessing environmental risks for established invasive weeds: Dalmatian (Linaria dalmatica) and yellow (L. vulgaris) toadflax in North America. International Journal of Environmental Research and Public Health 8(7) 2828-53.
^ "Chemical constituents and biological activities of the genus Linaria (Scrophulariaceae)".
^ Duke, J. A. Ethnobotanical uses: Linaria vulgaris. Dr. Duke's Phytochemical and Ethnobotanical Databases.

Bibliography

A Phylogeny of Toadflaxes (Linaria Mill.) Based on Nuclear Internal Transcribed Spacer Sequences: Systematic and Evolutionary Consequences. Mario Fernández-Mazuecos, José Luis Blanco-Pastor, and Pablo Vargas. International Journal of Plant Sciences, Vol. 174, No. 2 (February 2013), pp. 234–249 Published by: The University of Chicago Press, Article DOI: 10.1086/668790
Vargas P, JA Rosselló, R Oyama, J Güemes. 2004 Molecular evidence for naturalness of genera in the tribe Antirrhineae (Scrophulariaceae) and three independent evolutionary lineages from the New World and the Old. Plant Systematics and Evolution 249:151–172.
Media related to Linaria at Wikimedia Commons