Scrophulariaceae Explained

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

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.^[1] Many genera have recently been transferred to other families within the Lamiales, notably Plantaginaceae and Orobanchaceae, but also several new families.^{[2] [3]} 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) circumscription includes 62 genera and about 1830 known species.^[4] As of May 2024, Plants of the World Online accepts 58 genera.^[5]

Tribe Androyeae^[6] Androya Tribe Aptosimeae^[7] Anticharis Endl. Aptosimum Burch. ex Benth. Peliostomum E.Mey. ex Benth. Tribe Buddlejeae^[8] Buddleja L.  - Butterfly-bush Gomphostigma Turcz. Tribe Camptolomeae Camptoloma Benth. Tribe Hemimerideae^[9] Alonsoa Ruiz & Pav. Colpias E.Mey. ex Benth. Diascia Link & Otto Diclis Benth. Hemimeris L.f. Nemesia Vent. Tribe Leucophylleae^[10] Capraria L.^[11] Eremogeton Standl. & L.O.Williams Leucophyllum Humb. & Bonpl. Tribe Limoselleae^[12] Barthlottia Eb.Fisch. Chaenostoma Benth. (sometimes included in Sutera but separated by Kornhall and Bremer^[13]) Chenopodiopsis Hilliard Cromidon Compton Dischisma Choisy Glekia Hilliard Globulariopsis Compton Glumicalyx Hiern Gosela Choisy Hebenstretia L Jamesbrittenia Kuntze Limosella L.^[14] Lyperia Benth. Manulea 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^[15] Bontia L. Calamphoreus Chinnock (synonym of Eremophila) Diocirea Chinnock (synonym of Eremophila) Eremophila R.Br. Glycocystis Chinnock Myoporum Sol. ex G.Forst. Tribe Scrophularieae^[16] Antherothamnus N.E.Br. Nathaliella B.Fedtsch. Rhabdotosperma Hartl (synonym of Verbascum) × Scrophulari-verbascum Scrophularia L.  - Figwort Verbascum L.  - Mullein Tribe Teedieae^[17] Dermatobotrys Bolus Freylinia Colla Oftia Adans. Phygelius E.Mey. ex Benth.  - Cape fuchsia^[18] Ranopisoa J.F.Leroy Teedia Rudolphi Not placed in a tribe or unknown tribe Ameroglossum Eb.Fisch., S.Vogel & A.V.Lopes Chamaecrypta Dolichostemon

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^[19] or as part of Lindernia^[20] Cymbalaria → Plantaginaceae Dasistoma Raf. → Orobanchaceae^[21] 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^[22] Limnophila R.Br. → Plantaginaceae Linaria Mill. → Plantaginaceae Lindernia All. → Linderniaceae^[23] 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^[24] 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^[25]

Further reading

• Book: Fischer, E. . Kubitzki, K. . Kadereit, J. W. . Flowering Plants – Dicotyledons: Lamiales . fischer2004scrophulariaceae . 333−432 . Scrophulariaceae . The Families and Genera of Vascular Plants . VII . 2004 . Springer . 978-3-540-40593-1.

External links

• Kew list of Scrophulariaceae
• Family Scrophulariaceae Flowers in Israel
• verbascum.org

Notes and References

1. 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.
2. Olmstead, R. G., dePamphilis, C. W., Wolfe, A. D., Young, N. D., Elisons, W. J. & Reeves P. A.. Disintegration of the Scrophulariaceae . American Journal of Botany. 88. 348–361. 2001 . 10.2307/2657024 . 11222255 . 2 . 2657024. free.
3. Olmstead, R. G. . Whatever happened to the Scrophulariaceae? . Fremontia. 30. 13–22. 2003. - on line here
4. Christenhusz, M. J. M., and Byng, J. W. . 2016 . The number of known plants species in the world and its annual increase . Phytotaxa . 261 . 201–217 . 10.11646/phytotaxa.261.3.1 . 3 . free .
5. https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:30466893-2 Scrophulariaceae Juss.
6. 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
7. Web site: Genera of Scrophulariaceae tribe Aptosimeae . Germplasm Resources Information Network . United States Department of Agriculture . 2015-11-28.
8. Web site: Genera of Scrophulariaceae tribe Buddlejeae . Germplasm Resources Information Network . United States Department of Agriculture . 2015-11-28.
9. Web site: Genera of Scrophulariaceae tribe Hemimerideae . Germplasm Resources Information Network . United States Department of Agriculture . 2015-11-28.
10. Web site: Genera of Scrophulariaceae tribe Leucophylleae . Germplasm Resources Information Network . United States Department of Agriculture . 2015-11-28.
11. Etelvina . Gándara . Victoria Sosa . 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 . March 2013 . 10.1111/j.1095-8339.2012.01327.x. free .
12. Web site: Genera of Scrophulariaceae tribe Limoselleae . Germplasm Resources Information Network . United States Department of Agriculture . 2015-11-28.
13. 10.1111/j.1095-8339.2004.00341.x . New circumscription of the tribe Limoselleae (Scrophulariaceae) that includes the taxa of the tribe Manuleeae . Kornhall, Per . Bremer, Birgitta . amp . Botanical Journal of the Linnean Society . 146 . 453–467 . 4 . 2004 . free .
14. [Plants of the World Online]
15. Web site: Genera of Scrophulariaceae tribe Myoporeae . Germplasm Resources Information Network . United States Department of Agriculture . 2015-11-28.
16. Web site: Genera of Scrophulariaceae tribe Scrophularieae . Germplasm Resources Information Network . United States Department of Agriculture . 2015-11-28.
17. Web site: Genera of Scrophulariaceae tribe Scrophularieae . Germplasm Resources Information Network . United States Department of Agriculture . 2015-11-28.
18. Web site: Genera of Scrophulariaceae . Germplasm Resources Information Network . United States Department of Agriculture . 2015-11-28.
19. 10.3732/ajb.92.2.297 . Piecing together the "new" Plantaginaceae . 2005 . Albach, D. C. . American Journal of Botany . 92 . 297–315 . Meudt . H. M. . Oxelman . B. . 2 . 21652407 .
20. Web site: Lindernia All. . A Catalogue of the Vascular Plants of Madagascar . Missouri Botanical Garden and Muséum National d'Histoire Naturelle .
21. https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:30015204-2 Dasistoma Raf.
22. https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:37771-1 Leucosalpa Scott Elliot
23. Haston, E.. Richardson, J. E., Stevens, P. F., Chase, M. W., Harris, D. J. . A linear sequence of Angiosperm Phylogeny Group II families. Taxon. 56. 1. 2007. 7–12. 10.2307/25065731 . 25065731 . free.
24. The Evolution of Parasitism in Scrophulariaceae/Orobanchaceae: Plastid Gene Sequences Refute an Evolutionary Transition Series . Nelson D. Young . Kim E. Steiner . Claude W. dePamphilis . Annals of the Missouri Botanical Garden . 86 . 4 . Autumn 1999 . 876–893 . 10.2307/2666173 . 2666173.
25. Web site: GRIN genera sometimes placed in Scrophulariaceae . Germplasm Resources Information Network . United States Department of Agriculture . 2011-10-17 . dead . https://web.archive.org/web/20150924062306/http://www.ars-grin.gov/cgi-bin/npgs/html/gnothlist.pl?1023 . 2015-09-24 .