Scrophulariaceae

Life-forms

Classification System: APG IV

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

Familia: Scrophulariaceae
Tribus: Aptosimeae – Buddlejeae – Hemimerideae – Leucophylleae – Limoselleae – Myoporeae – Scrophularieae – Teedieae – incertae sedis
Buddlejeae - Freylinieae - Hemimerideae - Leucophylleae - Manuleae - Myoporeae - Scrophularieae

Genera: Agathelpis – Alonsoa – Ameroglossum – Anamaria – Androya – Antherothamnus – Anticharis – Aptosimum – Barthlottia – Boelckea – Bontia – Buddleja – Camptoloma – Capraria – Chaenostoma – Cheilophyllum – Chenopodiopsis – Colpias – Cromidon – Dermatobotrys – Diascia – Diclis – Dischisma – Dolichostemon – Emorya – Eremogeton – Eremophila – Fonkia – Freylinia – Glekia – Globulariopsis – Glumicalyx – Gomphostigma – Gosela – Hebenstretia – Hemimeris – Jamesbrittenia – Leucophyllum – Limosella – Lyperia – Magdalenaea – Manulea – Manuleopsis – Melanospermum – Microdon – Mimulicalyx – Myoporum – Namation – Nathaliella – Nemesia – Oftia – Oreosolen – Parasopubia – Peliostomum – Philcoxia – Phygelius – Phyllopodium – Polycarena – Pseudoselago – Ranopisoa – Reyemia – Rhabdotosperma – Scrophucephalus – Scrophularia – Selago – Stemodiopsis – Strobilopsis – Sutera – Teedia – Tetraselago – Trieenea – Verbascum – Walafrida – Zaluzianskya

Name

Scrophulariaceae Juss., Gen. Pl.: 117 (1789), nom. cons.

Type genus: Scrophularia L.

Synonyms

Heterotypic
Bontiaceae Horan.
Buddlejaceae K.Wilh., nom. cons.
Caprariaceae Martynov
Hebenstretiaceae Horan.
Hemimeridaceae Doweld
Limosellaceae J.Agardh
Myoporaceae R.Br., nom. cons.
Oftiaceae Takht. & Reveal
Selaginaceae Choisy, nom. cons.
Verbascaceae Bercht. & J.Presl

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

Olmstead, R.G., de Pamphilis, C.W., Wolfe, A.D., Young, N.D., Elisens, W.J. & Reeves, P.A. 2001. Disintegration of the Scrophulariaceae. American Journal of Botany 88(2): 348–361. DOI: 10.2307/2657024 Open access Reference page.
Olmstead, R.G. 2002. Whatever happened to the Scrophulariaceae? Fremontia 30(2): 13–22. PDF. Reference page.
Tank, D.C., Beardsley, P.M., Kelchner, S.A. & Olmstead, R.G. 2006. Review of the systematics of Scrophulariaceae s.l. and their current disposition. Australian Systematic Botany 19(4): 289–307. DOI: 10.1071/SB05009 Paywall PDF. Reference page.
Olmstead, R.G. (ed.) 2021. A Synoptical Classification of the Lamiales. Version 2.8.0 (in prog.). 20 pp. PDF Reference page.

Links

Govaerts, R. et al. 2014. Scrophulariaceae in Kew Science Plants of the World Online. The Board of Trustees of the Royal Botanic Gardens, Kew. Published online. Accessed: 2014 Feb. 4. Reference page.
Hassler, M. 2020. Scrophulariaceae. 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 online. Accessed: 2020 Dec. 11. Reference page.
Tropicos.org 2014. Scrophulariaceae. Missouri Botanical Garden. Published online. Accessed: 4 Feb. 2014. Reference page.
Find all Wikispecies pages which cite this reference.

Lohmueller, F.A. 2005. The Botanical System of the Plants [1][2][3]

Vernacular names
aragonés: Escrofulariacias
العربية: غدبية
asturianu: Escrofulariácees
azərbaycanca: Keçiqulağıkimilər
башҡортса: Сираж һымаҡтар
беларуская: Залознікавыя
български: Живеничеви
català: Escrofulariàcia
čeština: Krtičníkovité
dansk: Maskeblomst-familien
Deutsch: Braunwurzgewächse
English: Scrophs, figwort family
Esperanto: Skrofulariacoj
español: Escrofulariáceas
eesti: Mailaselised
euskara: Eskrofulariazeoak
فارسی: گل‌میمونیان
suomi: Naamakukkaiskasvit
français: Scrofulariacées
עברית: לועניתיים
hornjoserbsce: Trudownikowe rostliny
magyar: Görvélyfűfélék, tátogatófélék, tátikafélék
հայերեն: Խլածաղկազգիներ
日本語: ゴマノハグサ科
ქართული: შავწამალასებრნი
қазақша: Сабынкөктер тұқымдасы
한국어: 현삼과
lietuvių: Bervidiniai
македонски: Струпници
Nederlands: Leeuwebekachtigen
norsk nynorsk: Brunrotfamilien
norsk: Scrophulariaceae
polski: Trędownikowate
Runa Simi: Kiswar yura rikch'aq ayllu
русский: Норичниковые
slovenčina: Krtičníkovité
svenska: Flenörtsväxter
ไทย: วงศ์มณเฑียรทอง
Türkçe: Sıraca otugiller
українська: Ранникові
Tiếng Việt: Họ Huyền sâm
中文: 玄参科

The Scrophulariaceae are a family of flowering plants, commonly known as the figwort family. The plants are annual and perennial herbs, as well as shrubs. Flowers have bilateral (zygomorphic) or rarely radial (actinomorphic) symmetry. The Scrophulariaceae have a cosmopolitan distribution, with the majority found in temperate areas, including tropical mountains. The family name is based on the name of the included genus Scrophularia L.
Taxonomy

This article needs attention from an expert in botany. The specific problem is: This article needs a comparison between the current and obsolete circumscriptions of the group, its current composition, and the present location of excluded taxa. None of these issues is adequately dealt with here. WikiProject Botany may be able to help recruit an expert. (August 2015)

In the past, it was treated as including about 275 genera and over 5,000 species, but its circumscription has been radically altered since numerous molecular phylogenies have shown the traditional broad circumscription to be grossly polyphyletic.[2] Many genera have recently been transferred to other families within the Lamiales, notably Plantaginaceae and Orobanchaceae, but also several new families.[3][4] Several families of the Lamiales have had their circumscriptions enlarged to accommodate genera transferred from the Scrophulariacae sensu lato.

Fischer (2004) considered the family to consist of three subfamilies – Antirrhinoideae, Gratioloideae, and Digitalidoideae. He further divided the Gratioloideae into five tribes – Gratioleae, Angeloniaeae, Stemodieae, Limoselleae, and Lindernieae. He then divided the Gratioleae, with its 16 genera (and about 182 species) into three subtribes – Caprarinae, Dopatrinae, and Gratiolinae. The Gratiolinae had 10 genera (about 121 species) distributed through temperate and tropical America – Bacopa and Mecardonia (formerly Herpestis), Amphianthus, Gratiola, Sophronanthe, Benjaminia, Scoparia, Boelkea, Maeviella, and Braunblequetia. Many of these were transferred to the family Plantaginaceae, in the tribe Gratioleae.
Uses

The family includes some medicinal plants, among them:

Scrophularia, figworts
Verbascum, mulleins

Genera

The family Scrophulariaceae in its APG IV (2016)[5] circumscription includes 62 genera and about 1830 known species.[6] As of May 2024, Plants of the World Online accepts 58 genera.[7]
Leucophyllum frutescens
Verbascum sp. near Shiraz, Iran

Tribe Androyeae[8]

Androya H.Perrier

Tribe Aptosimeae[9]

Anticharis Endl.
Aptosimum Burch. ex Benth.
Peliostomum E.Mey. ex Benth.

Tribe Buddlejeae[10]

Buddleja L. – Butterfly-bush
Gomphostigma Turcz.

Tribe Camptolomeae[8]

Camptoloma Benth.

Tribe Hemimerideae[11]

Alonsoa Ruiz & Pav.
Colpias E.Mey. ex Benth.
Diascia Link & Otto
Diclis Benth.
Hemimeris L.f.
Nemesia Vent.

Tribe Leucophylleae[12]

Capraria L.[13]
Eremogeton Standl. & L.O.Williams
Leucophyllum Humb. & Bonpl.

Tribe Limoselleae[14]

Barthlottia Eb.Fisch.
Chaenostoma Benth. (sometimes included in Sutera but separated by Kornhall and Bremer[15])
Chenopodiopsis Hilliard
Cromidon Compton
Dischisma Choisy
Glekia Hilliard
Globulariopsis Compton
Glumicalyx Hiern
Gosela Choisy
Hebenstretia L
Jamesbrittenia Kuntze[15]
Limosella L.[16]
Lyperia Benth.
Manulea L.
Manuleopsis Thell. ex Schinz
Melanospermum Hilliard
Microdon Choisy

Phyllopodium Benth.
Polycarena Benth.
Selago L.
Strobilopsis Hilliard & B.L.Burtt
Sutera Roth – Dwarf snapdragon, "bacopa"
Tetraselago Junell
Trieenea Hilliard
Zaluzianskya F.W.Schmidt

Tribe Myoporeae[17]

Bontia L.
Calamphoreus Chinnock (synonym of Eremophila)
Diocirea Chinnock (synonym of Eremophila)
Eremophila R.Br.
Glycocystis Chinnock
Myoporum Sol. ex G.Forst.

Tribe Scrophularieae[18]

Antherothamnus N.E.Br.
Nathaliella B.Fedtsch.
Rhabdotosperma Hartl (synonym of Verbascum)
× Scrophulari-verbascum P.Fourn.
Scrophularia L. – Figwort
Verbascum L. – Mullein

Tribe Teedieae[8][19]

Dermatobotrys Bolus
Freylinia Colla
Oftia Adans.
Phygelius E.Mey. ex Benth. – Cape fuchsia[20]
Ranopisoa J.F.Leroy
Teedia Rudolphi

Not placed in a tribe or unknown tribe

Ameroglossum Eb.Fisch., S.Vogel & A.V.Lopes
Chamaecrypta Schltr. & Diels
Dolichostemon Bonati

Excluded genera

The following genera, traditionally included in the Scrophulariaceae, have been transferred to other families as indicated:

Acanthorrhinum Rothm. → Plantaginaceae
Agalinis Raf. → Orobanchaceae
Angelonia Humb. & Bonpl. → Plantaginaceae
Antirrhinum L. → Plantaginaceae
Asarina Mill. → Plantaginaceae
Bacopa Aubl. → Plantaginaceae
Bartsia L. → Orobanchaceae
Basistemon Turcz. → Plantaginaceae
Besseya Rydb. → Plantaginaceae
Calceolaria L. → Calceolariaceae
Castilleja Mutis ex L.f. → Orobanchaceae
Chaenorhinum (DC.) Rchb. → Plantaginaceae
Chelone L. → Plantaginaceae
Chionophila Benth. → Plantaginaceae
Collinsia Nutt. → Plantaginaceae
Cordylanthus Nutt. ex Benth. → Orobanchaceae
Craterostigma Hochst. → Linderniaceae[21] or as part of Lindernia[22]
Cymbalaria → Plantaginaceae
Dasistoma Raf. → Orobanchaceae[23]
Digitalis L. → Plantaginaceae
Dopatrium Buch.-Ham. ex Benth. → Plantaginaceae
Epixiphium (Engelm. ex A.Gray) Munz → Plantaginaceae
Euphrasia L. → Orobanchaceae
Galvezia Dombey ex Juss. → Plantaginaceae
Gambelia Nutt. → Plantaginaceae
Glossostigma Wight & Arn. → Phrymaceae
Gratiola L. → Plantaginaceae
Hebe Comm. ex Juss. → Plantaginaceae
Holmgrenanthe Elisens → Plantaginaceae
Howelliella Rothm. → Plantaginaceae
Keckiella Straw → Plantaginaceae
Kickxia Dumort. → Plantaginaceae
Lagotis Gaertn. → Plantaginaceae
Leucosalpa Scott Elliot → Orobranchaceae[24]
Limnophila R.Br. → Plantaginaceae
Linaria Mill. → Plantaginaceae
Lindernia All. → Linderniaceae[25]
Lophospermum D.Don → Plantaginaceae

Mabrya Elisens → Plantaginaceae
Maurandella (A.Gray) Rothm. → Plantaginaceae
Maurandya Ortega → Plantaginaceae
Mazus Lour. → Mazaceae
Mecardonia Ruiz & Pav. → Plantaginaceae
Melampyrum L. → Orobanchaceae
Micranthemum Michx. → Plantaginaceae
Mimulus L. → Phrymaceae
Misopates Raf. → Plantaginaceae
Mohavea A.Gray → Plantaginaceae
Neogaerrhinum Rothm. → Plantaginaceae
Nothochelone (A.Gray) Straw → Plantaginaceae
Nuttallanthus D.A.Sutton → Plantaginaceae
Odontites Ludw. → Orobanchaceae
Orobanche L. → Orobanchaceae
Orthocarpus Nutt. → Orobanchaceae
Parentucellia Viv. → Orobanchaceae
Paulownia Siebold & Zucc. → Paulowniaceae
Pedicularis L. → Orobanchaceae
Peltanthera Benth. → Gesneriaceae (or Peltantheraceae)
Penstemon Schmidel → Plantaginaceae
Pseudorontium (A.Gray) Rothm. → Plantaginaceae
Rhinanthus L. → Orobanchaceae
Russelia Jacq. → Plantaginaceae
Sairocarpus D.A.Sutton → Plantaginaceae
Sanango G.S.Bunting & Duke → Gesneriaceae
Schistophragma Benth. ex Endl. → Plantaginaceae
Scoparia L. → Plantaginaceae
Stemodia L. → Plantaginaceae
Striga Lour. → Orobanchaceae[3][26]
Synthyris Benth. → Plantaginaceae
Tonella Nutt. ex A.Gray → Plantaginaceae
Torenia L. → Linderniaceae
Tozzia L. → Orobanchaceae
Triphysaria Fisch. & C.A.Mey. → Orobanchaceae
Veronica L. → Plantaginaceae
Veronicastrum Heist. ex Fabr. → Plantaginaceae[27]

References

"Family: Scrophulariaceae Juss., nom. cons". Germplasm Resources Information Network. United States Department of Agriculture. 2003-01-17. Retrieved 2011-10-17.
Fischer says that, if we consider morphology, it has been obvious for the last decades that the Scrophulariaceae do not represent a monophyletic group. See FISCHER (2004), p. 346.
Olmstead, R. G., dePamphilis, C. W., Wolfe, A. D., Young, N. D., Elisons, W. J. & Reeves P. A. (2001). "Disintegration of the Scrophulariaceae". American Journal of Botany. 88 (2): 348–361. doi:10.2307/2657024. JSTOR 2657024. PMID 11222255.
Olmstead, R. G. (2003). "Whatever happened to the Scrophulariaceae?". Fremontia. 30: 13–22. - on line here
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, 161 (2): 105–20, doi:10.1111/boj.12385
Christenhusz, M. J. M., and Byng, J. W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. 261 (3): 201–217. doi:10.11646/phytotaxa.261.3.1.
Scrophulariaceae Juss. Plants of the World Online. Retrieved 13 May 2024.
Villaverde, T., Larridon, I., Shah, T., Fowler, R.M., Chau, J.H., Olmstead, R.G. and Sanmartín, I. (2023), Phylogenomics sheds new light on the drivers behind a long-lasting systematic riddle: the figwort family Scrophulariaceae. New Phytol, 240: 1601-1615. https://doi.org/10.1111/nph.18845
"Genera of Scrophulariaceae tribe Aptosimeae". Germplasm Resources Information Network. United States Department of Agriculture. Retrieved 2015-11-28.
"Genera of Scrophulariaceae tribe Buddlejeae". Germplasm Resources Information Network. United States Department of Agriculture. Retrieved 2015-11-28.
"Genera of Scrophulariaceae tribe Hemimerideae". Germplasm Resources Information Network. United States Department of Agriculture. Retrieved 2015-11-28.
"Genera of Scrophulariaceae tribe Leucophylleae". Germplasm Resources Information Network. United States Department of Agriculture. Retrieved 2015-11-28.
Gándara, Etelvina; Victoria Sosa (March 2013). "Testing the monophyly and position of the North American shrubby desert genus Leucophyllum (Scrophulariaceae: Leucophylleae)". Botanical Journal of the Linnean Society. 171 (3): 508–518. doi:10.1111/j.1095-8339.2012.01327.x.
"Genera of Scrophulariaceae tribe Limoselleae". Germplasm Resources Information Network. United States Department of Agriculture. Retrieved 2015-11-28.
Kornhall, Per & Bremer, Birgitta (2004). "New circumscription of the tribe Limoselleae (Scrophulariaceae) that includes the taxa of the tribe Manuleeae". Botanical Journal of the Linnean Society. 146 (4): 453–467. doi:10.1111/j.1095-8339.2004.00341.x.
Plants of the World Online and World Flora Online place Limosella in family Plantaginaceae – Limosella L. (POWO), Limosella L. (WFO), Retrieved 13 May 2024.
"Genera of Scrophulariaceae tribe Myoporeae". Germplasm Resources Information Network. United States Department of Agriculture. Retrieved 2015-11-28.
"Genera of Scrophulariaceae tribe Scrophularieae". Germplasm Resources Information Network. United States Department of Agriculture. Retrieved 2015-11-28.
"Genera of Scrophulariaceae tribe Scrophularieae". Germplasm Resources Information Network. United States Department of Agriculture. Retrieved 2015-11-28.
"Genera of Scrophulariaceae". Germplasm Resources Information Network. United States Department of Agriculture. Retrieved 2015-11-28.
Albach, D. C.; Meudt, H. M.; Oxelman, B. (2005). "Piecing together the "new" Plantaginaceae". American Journal of Botany. 92 (2): 297–315. doi:10.3732/ajb.92.2.297. PMID 21652407.
"Lindernia All". A Catalogue of the Vascular Plants of Madagascar. Missouri Botanical Garden and Muséum National d'Histoire Naturelle.
Dasistoma Raf. Plants of the World Online. Retrieved 13 May 2024.
Leucosalpa Scott Elliot. Plants of the World Online. Retrieved 13 May 2024.
Haston, E., Richardson, J. E., Stevens, P. F., Chase, M. W., Harris, D. J. (2007). "A linear sequence of Angiosperm Phylogeny Group II families". Taxon. 56 (1): 7–12. doi:10.2307/25065731. JSTOR 25065731.
Nelson D. Young; Kim E. Steiner; Claude W. dePamphilis (Autumn 1999). "The Evolution of Parasitism in Scrophulariaceae/Orobanchaceae: Plastid Gene Sequences Refute an Evolutionary Transition Series". Annals of the Missouri Botanical Garden. 86 (4): 876–893. doi:10.2307/2666173. JSTOR 2666173.

"GRIN genera sometimes placed in Scrophulariaceae". Germplasm Resources Information Network. United States Department of Agriculture. Archived from the original on 2015-09-24. Retrieved 2011-10-17.

Further reading
Fischer, E. (2004). "Scrophulariaceae". In Kubitzki, K.; Kadereit, J. W. (eds.). Flowering Plants – Dicotyledons: Lamiales. The Families and Genera of Vascular Plants. Vol. VII. Springer. pp. 333−432. ISBN 978-3-540-40593-1.