Description:

• Annual or perennial, terrestrial, amphibious or aquatic herbs, sometimes undershrubs, rarely shrubs or small trees, sometimes hemiparasitic or holoparasitic on roots and lacking chlorophyll
• Stems cylindrical or quadrangular, sometimes ribbed or narrowly winged
• Leaves opposite and/or alternate, or rarely whorled, sometimes fascicled or radical, simple, entire to deeply divided, rarely reduced to scales or aborted; stipules 0
• Flowers bisexual, strongly to slightly irregular, rarely subregular or regular, pedicellate or not, solitary axillary or in terminal or at times subterminal and lateral cymes or racemes, in such forms as spikes, heads, panicles or thyrses
• Bracts opposite or alternate, free or adnate to pedicel, rarely to calyx; bracteoles 0 or 2
• Calyx usually (4)5-lobed, or rarely, by modification, 2- or 3-lobed or spathaceous, usually ± united, persistent, tube from almost lacking to long, usually cylindrical to campanulate; lobes equal or not, valvate, variously overlapping or open in bud
• Corolla sympetalous, (3)4- or 5(6-8)-lobed, usually campanulate, cylindrical or ventricose or enlarged above, or bilabiate with upper (posterior) lip entire, 2- or sometimes 4-lobed, erect, concave, ± hooded, flat or spreading; lower (anterior) lip (1)3(4)-lobed, spreading; sometimes corolla ± unilabiate with tube split in front and limb of 4(5) lobes; tube straight, or variously curved to geniculate, at times oblique, sometimes with 1 or 2 spurs, sacs or hollows at base or in throat; lobes subequal or posterior pair shorter, ± spreading
• Stamens usually 4, didynamous or at times ± equal, or 2 with sometimes 2 reduced to staminodes, or 5 with fifth subrudimentary, rarely perfect, alternating with corolla lobes, exserted or included; filaments arising in corolla tube or at throat, filiform or dilated, free, confluent or decurrent down tube, occasionally with lower pair variously appendaged at base, there sometimes anterior filaments twisted around and thus occupying posterior position; anthers usually bithecate, sometimes synthecate (at length unithecate by confluence of 2 thecae), at times unithecate, free or often coherent; thecae equal or with one smaller and at times sterile; staminodes 0, 1 or 2
• Nectary/disc various, often annular or unilateral at base of ovary, sometimes obsolete
• Ovary superior, bilocular (rarely ± unilocular or trilocular), sessile, entire, with carpels median; placentas central, adnate to septum or rarely parietal; ovules usually many, sometimes few or 1 per locule, often seated on pulvinate funicle with central depression; style terminal, simple, usually filiform; stigma entire or 2(3)-lobed
• Fruit usually a capsule, loculicidal and/or septicidal, sometimes dehiscing by pores or by meridional fissures, or indehiscent and then at times separating into 2 1-seeded nutlets (cocci), or rarely 1 nutlet by abortion, seldom a berry, rarely a fleshy drupe
• Seeds small to minute, 1-many per locule, variously shaped, pitted, furrowed, ridged, ribbed or winged, rarely smooth; embryo straight to weakly curved; endosperm oily

Classification Notes:

• Classification and delimitation of the Scrophulariaceae are problematical, and in its present circumscription the family is not considered monophyletic (Olmstead & Reeves 1995). The Selaginaceae, established as a family by Choisy (1823), and accepted as such by Dyer (1975), Dahlgren (1980), Cronquist (1988, under Globulariaceae) and Hartley & Balkwill (1990), is here included in Scrophulariaceae. This is in agreement with, among others, Hilliard (1990, 1994 and 1999), Philcox (1990), Brummitt (1992), Mielcarek (1996) and Takhtajan (1997)
• *Orobanchaceae is also difficult to separate from Scrophulariaceae. Some genera have been moved back and forth between them, such as Buchnera L., Harveya Hook., Hyobanche L. and *Orobanche L. The two families have been treated as distinct by, for example, Cronquist (1988), Kreutz (1995) and Mabberley (1997). Others recognise *Orobanchaceae as part of Scrophulariaceae, such as Brummitt (1992), Thorne (1992), Mielcarek (1996) and Takhtajan (1997), and they are followed here.
• Recent research is leading to publication of other classification schemes. For instance, based on evidence from morphology, DNA base-pair sequences and chemistry, Judd et al. (1999) divide the genera here placed under Scrophulariaceae into three families which they consider monophyletic:
  • 1. Scrophulariaceae, which includes the closely related Scrophularia L. and *Verbascum L. These two genera are seen as closely linked phenetically to Selago L. and relatives (Selagineae) by the intermediate group of Sutera Roth and relatives (Manuleae)
  • 2. Plantaginaceae, which comprises Plantago L., Bacopa Aubl., Limnophila R.Br., *Linaria Mill., Lindernia All., Mimulus L., Nemesia Vent., *Scoparia L. and others
  • 3. *Orobanchaceae, which includes the parasitic genera, such as *Orobanche L. and Harveya Hook.
• Oftia Adans. and Hiernia S.Moore, listed by Dyer (1975) under Scrophulariaceae, are retained under that family, the first with reservation. Oftia has been placed in Verbenaceae, Myoporaceae, Scrophulariaceae or in a separate family of its own. Takhtajan (1997) placed it in Oftiaceae, next to the Scrophulariaceae, but noted it shows similarities with Teedia, in the last family. Oftia is regarded here as an anomalous genus in Scrophulariaceae. Charadrophila Marloth, usually placed under Gesneriaceae (also by Dyer 1975), was transferred by Weber (1989) to Scrophulariaceae, mainly because of its capsular fruits. This was accepted by Brummitt (1992) and Mabberley (1997) and is followed here. In Scrophulariaceae the genus occupies an isolated position, possibly near Alonsoa Ruiz & Pav. (Weber 1989), although it has features aligning it with the Rhinanathoideae.
• Some exotic species of genera such as *Cymbalaria Hill, *Linaria Mill., Mimulus L., *Verbascum L. and Veronica L. are widely cultivated and may occur as garden escapes. Others are found as weeds, for example *Kickxia Dumort., *Scoparia L., *Misopates Raf. and *Orobanche L.
• The southern Africa genera of Scrophulariaceae can be classified based mainly on the classification system of Takhtajan (1997) and groupings of genera into tribes by Hilliard (1994, 1999), Fischer (1992), Barringer (1993) and Steiner (1996 and pers. com. 2000). However, relationships are far from definite and several genera are of very uncertain affinity (marked by ? in the scheme)

Nomenclature:

• Scrophulariaceae
  • Choisy: 71 (1823)
  • Bentham: 186 (1846)
  • Bentham: 913, 1126 (1876)
  • Wettstein: 39 (1891/1893)
  • Hallier: 181 (1903)
  • Hiern: 121 (1904)
  • Hiern & Stapf: 421 (1904)
  • Hemsley & Skan: 261 (1906)
  • Merxmüller & Roessler: 1 (1967a)
  • Merxmüller & Roessler: 1 (1967b)
  • Webb: 202 (1972)
  • Hilliard & Burtt: 155 (1977)
  • Visser: 82 (1981)
  • Sutton: 1 (1988)
  • Hilliard & Burtt: 179 (1989)
  • Hilliard: 315 (1990)
  • Philcox: 1 (1990)
  • Brummitt: 665 (1992)
  • Fischer: 1 (1992)
  • Hilliard: 1 (1994)
  • Mielcarek: 3 (1996)
  • Steiner: 63 (1996)
  • Mabberley: 1 (1997)
  • Retief & Herman: 548, 603, 617 (1997)
  • Takhtajan: 451 (1997)
  • Hilliard: 1 (1999)

Distribution & Notes:

• Global: Genera ± 290, species ± 4500 (including *Orobanchaceae, Selaginaceae & Charadrophila); cosmopolitan but especially temperate areas and tropical mountains
• Southern Africa: Genera ± 79 (80) genera, ± 760 species, widespread

References:

• BARRINGER, K. 1993. Five new tribes in the Scrophulariaceae. Novon 3
• BENTHAM, G. 1846. Scrophulariaceae. In A.P. de Candolle, Prodromus 10. Masson, Paris
• BENTHAM, G. 1876. Scrophularineae. In G. Bentham & J.D. Hooker, Genera plantarum 2. Lovell Reeve & Co., London
• BRUMMITT, R.K. 1992. Vascular plant families and genera. Royal Botanic Gardens, Kew
• CHOISY, J.D. 1823. Mémoire sur la famille des Sélaginées. Mémoires de la Société de Physique et d' Histoire Naturelle de Genève 2(2)
• FISCHER, E. 1992. Systematik der afrikanischen Lindernieae (Scrophulariaceae). Tropische und subtropische Pflanzenwelt 81
• HALLIER H. 1903. Über die Abgrenzung und Verwandtschaft der einzelnen Sippen bei den Scrophulariaceen. Bulletin de l'Herbier Boissier 2, 3
• HEMSLEY, W.B. & SKAN, S.A. 1906. Scrophulariaceae. Flora of tropical Africa 4
• HIERN, W.P. 1904. Scrophulariaceæ. Flora capensis 4,2
• HIERN, W.P. & STAPF, O. 1904. Orobanchaceæ. Flora capensis 4,2
• HILLIARD, O.M. 1990. A brief survey of Scrophulariaceae - Selagineae. Edinburgh Journal of Botany 47
• HILLIARD, O.M. 1994. The Manuleae. A tribe of Scrophulariaceae. Edinburgh University Press, Edinburgh
• HILLIARD, O.M. 1999. The tribe Selagineae (Scrophulariaceae). Royal Botanic Garden, Edinburgh
• HILLIARD, O.M. & BURTT, B.L. 1977. Notes on some plants of southern Africa chiefly from Natal: VI. Notes from the Royal Botanic Garden Edinburgh 35
• HILLIARD, O.M. & BURTT, B.L. 1989 ('1988'). Notes on some plants of southern Africa chiefly from Natal: XV. Notes from the Royal Botanic Garden Edinburgh 45
• MABBERLEY, D.J. 1997. The plant-book, edn 2. Cambridge University Press, Cambridge
• MERXMÜLLER, H. & ROESSLER, H. 1967a. Scrophulariaceae. Prodromus einer Flora von Südwestafrika 126
• MERXMÜLLER, H. & ROESSLER, H. 1967b. Selaginaceae. Prodromus einer Flora von Südwestafrika 127
• MIELCAREK, R. 1996. Les Scrophulariaceae dans la flore d'Afrique centrale (excl. Linderneae). Fragmenta floristica et geobotanica 41
• PHILCOX, D. 1990. Scrophulariaceae. Flora zambesiaca 8,2
• RETIEF, E. & HERMAN, P.P.J. 1997. Plants of the northern provinces of South Africa: keys and diagnostic characters. Strelitzia 6
• STEINER, K.E. 1996. Chromosome numbers and relationships in tribe Hemimerideae (Scrophulariaceae). Systematic Botany 21
• SUTTON, D.A. 1988. A revision of the tribe Antirrhineae. British Museum (Natural History), London
• TAKHTAJAN, A. 1997. Diversity and classification of flowering plants. Columbia University Press, New York
• VISSER, J. 1981. Scrophulariaceae and Orobanchaceae. South African parasitic flowering plants. Juta, Cape Town
• WEBB, D.A. 1972. Scrophulariaceae. Flora europaea 3
• WETTSTEIN, R. VON. 1891/1893. Scrophulariaceae. Die natürlichen Pflanzenfamilien 4, 3b

Resources:

• Classification of Scrophulariaceae
• Key to Scrophulariaceae genera

• Scrophulariaceae genera: