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(Last Modified On 8/14/2013)
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(Last Modified On 8/14/2013)
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Family
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SCROPHULARIACEAE
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Contributor
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W. G. D'ARCY
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Description
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Mostly herbs or shrubs, rarely trees or lianas, sometimes paludal or aquatic, sometimes parasitic or saprophytic, glabrous or pubescent with simple or branched, glandular or eglandular hairs, often gland dotted, often drying black. Leaves opposite or alternate, occasionally verticillate, simple or compound, mostly serrate to denticulate, sometimes deeply incised; exstipulate. Inflorescences cymose, racemose or spicate, often with flowers solitary or in pairs, mostly axillary; pedicels mostly subtended by 1-2 bracts, the bractlets sometimes present, the bracts sometimes colorful. Flowers mostly perfect, zygomorphic, the calyx mostly of 5 similar or dissimilar, united or deeply divided lobes; corolla 5 or fewer lobed, mostly 2-lipped, the upper (posterior) lip of 3 lobes, often elaborated into saccate or other irregular forms; stamens (5)4 or 2, staminodes sometimes present, inserted in the corolla tube, mostly didynamous or 2, the anthers 1-2 thecate, the thecae sometimes separated on variously elaborated connective arms, sometimes con- fluent; disc sometimes present; ovary 2 locular at least basally, sometimes 1 locular apically, each locule with many(-few-i) anatropous or amphitropous ovules on elaborated placentas; styles mostly united, the stigmas 2 or 1. Fruit mostly a septicidal or also loculicidal capsule, sometimes a berry or nutlike, the placenta often persistent as a tuberculate peg, sometimes winged; seeds small, the embryo straight or slightly bent, the endosperm fleshy.
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Habit
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herbs or shrubs, rarely trees or lianas
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Reference
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Pennell, F. W. 1935. The Scrophulariaceae of Eastern Temperate North Amer- ica. Acad. Nat. Sci. Philadelphia Monogr. 1: 1-650. Standley, P. C. & L. 0. Williams. 1973. Scrophulariaceae in Flora of Guate- mala, Fieldiana, Bot. 24 (IX 3-4): 319-418. Thieret, J. W. 1954. The tribes and genera of Central American Scrophularia- ceae. Ceiba 4: 164-184. Thieret, J. W. 1967. Supraspecific classification in Scrophulariaceae: a review. Sida 3: 87-106.
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Key
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1. Leaves deeply lobed, dissected or compound. 2. Plants submersed aquatics; leaves lax, the segments filiform ...... 5. Benjaminia 2'. Plants terrestrial; leaves turgid or stiff, the segments linear to broad. 3. Corollas bright yellow, sometimes drying orange; calyces green, not showy; leaves membranaceous, the segments mostly more than 5 mm wide; corolla exserted, the lower lip inflated, entire ...... 7. Calceolaria 3'. Corollas green, red or yellowish; calyces or bracts red, often showy; leaves stiff, the segments mostly less than 3 mm wide; corolla included, the tube elongate, the lower lip 3-toothed ...... 9. Castilleja 1'. Leaves simple, toothed, crenate or entire, rarely lobed or dissected. 4. Fruit a globose capsule packed with hairs; corollas tubular, red or pink, glabrous outside, the flowers in clusters or pairs ...... 18. Russelia 4'. Fruit a capsule lacking internal hairs, or a berry; corollas various, if red and tubular then pubescent outside, flowers variously disposed. 5. Flowers more than 1.3 cm long. 6. Corollas bright yellow, sometimes drying orange; leaves lanceolate, petiolate ...... 7. Calceolaria 6'. Corollas red, white, greenish, sometimes fading yellowish; leaves various, if lanceolate, then sessile. 7. Flowers narrowly tubular or salverform. 8. Leaves mostly linear, mostly less than 4 mm wide; plants drying dark ...... 6. Buchnera 8'. Leaves more than 4 mm wide, ovate; plants drying green or brown. 9. Corolla salverform, the limb flaring, more than 3 cm wide; pedicels stout and elongate, bibracteate partway up ...... 11. Escobedia 9'. Corolla tubular, the limb mostly erect, less than 2 cm wide; pedicels either slender or short, lacking bracts. 10. Leaves entire; corolla glabrous outside, white or purplish ...... 12. Gibsoniothamnus 10'. Leaves dentate or serrate; corolla pubescent outside, red or orange ...... 13. Lamourouxia 7'. Flowers campanulate or irregular (saccate). 11. Leaves linear, less than 4 mm wide; plants drying dark ...... 1. Agalinis 11'. Leaves linear to ovate, more than 4 mm wide; plants mostly drying green or brown. 12. Leaves more than 2 cm wide; flowers white or pinkish. 13. Leaves opposite. 14. Leaves lanceolate; coarse herb mostly over 40 cm tall; fruit a rotund white berry ......14. Leucocarpus 14'. Leaves ovate; forest herb less than 30 cm tall; fruit a narrow capsule ...... 21. Tetranema 13'. Leaves alternate ......10. Digitalis 12'. Leaves less than 2 cm wide; flowers blue or purplish. 15. Capsule enclosed in the winged calyx cup; leaves ovate ...... 22. Torenia 15'. Capsule exserted from the short calyx cup; leaves linear ...... 3. Angelonia 5'. Flowers less than 1.3 cm long. 16. Leaves lanceolate, mostly over 2 cm wide, 10 cm long; large herbs mostly over 40 cm tall; fruit a white berry ...... 14. Leucocarpus 16'. Leaves various, if lanceolate then less than 2 cm wide, 10 cm long; herbs mostly less than 40 cm tall, or shrubs; fruit a capsule. 17. Capsule apically divided into 2 rounded lobes (Fig. 23.) ...... 23. Veronica 17'. Capsule unlobed, obtuse or globose. 18. Leaves linear, less than 5 mm wide and several times longer than wide, mostly entire, sometimes minutely and sparingly toothed. 19. Flowering calyx tubular; plants drying dark. 20. Corolla pink, purplish or white, the limb of spreading, subequal lobes; calyx green or dark, not showy; anthers 1-thecate ...... 6. Buchnera 20'. Corolla green or yellowish, the limb of unlike erect and recurved lobes; calyx showy red or orange; anthers 2-thecate ...... 9. Castilleja 19'. Flowering calyx either cup shaped, tubular, or leaflike; plants mostly not drying dark. 21. Calyx lobes of different size and shape, mostly ovate ...... 4. Bacopa 21'. Calyx lobes all alike, mostly narrow. 22. Flowers sessile or nearly so ...... 20. Stemodia 22'. Flowers distinctly pedicellate. 23. Fruiting pedicels less than 10 mm long; flowers white ......19. Scoparia 23'. Fruiting pedicels more than 10 mm long; flowers pink ...... 1. Agalinis 18'. Leaves deltoid, ovate, rotund or elliptic, mostly less than twice as long as broad (sometimes less than 5 mm wide), entire or toothed. 24. Leaves alternate; shrubs; corolla white, campanulate ...... 8. Capraria 24'. Leaves opposite; herbs; corolla white, yellow or blue, shapes various. 25. Leaves scabrous; flowers yellow, inconspicuous in spike-like terminal racemes ...... 2. Alectra 25'. Leaves smooth; flowers variously colored, pedunculate or sessile, solitary or in clusters. 26. Plants erect or ascending, mostly over 6 cm tall. 27. Placenta conspicuously winged by the septum after seeds have fallen; flowers blue or purplish. 28. Capsule enclosed in the calyx cup ...... 22. Torenia 28'. Capsule exserted from the calyx cup ...... 15. Lindernia 27'. Placenta peglike, globose or rudimentary, unwinged; flowers white or bluish. 29. Calyx lobes of different size and shape, mostly ovate or cordate. 30. Flowers yellow; anther cells separated on arms ...... 16. Mecardonia 30'. Flowers white; anther cells contiguous, at least in part ...... 4. Bacopa 29'. Calyx lobes all alike, mostly narrow. 31. Capsules globose; flowers white ...... 19. Scoparia 31'. Capsules ovoid or ellipsoid; flowers blue ...... 20. Stemodia 26'. Plants prostrate, creeping, or ascending, less than 6 cm tall. 32. Fruits globose ...... 17. Micranthemum 32'. Fruits ovoid or ellipsoid. 33. Calyx lobes of different size and shape, mostly ovate or cordate. 34. Flowers yellow; anther cells separated on arms ...... 16. Mecardonia 34'. Flowers white; anther cells contiguous, at least in part ...... 4. Bacopa 33'. Calyx lobes all alike, mostly narrow ...... 20. Stemodia
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Distribution
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The Scrophulariaceae includes over 250 genera with over 5,000 species dis- tributed worldwide, but with the greatest number of species in temperate regions.
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Note
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Many of the species are wide ranging. The family can usually be recognized by the 2-lipped flowers with paired stamens and the conical or globose capsules which contain many small seeds and a peglike placenta. There are many exceptions to these features, but the Scrophulariaceae has every appearance of being a natural group. However, some genera usually placed in the family may belong to other families. Thus Niezgoda and Tomb (1975) report that the pollen of Capraria is more like that in the Myoporaceae than in the Srrophulariaceae, and Gibsoniothamnus has characters strongly suggesting placement in the Bignoniaceae or Gesneriaceae rather than in the Scrophulariaceae. Tetranema is quite unlike other members of the Scrophu- lariaceae and is strongly suggestive of the Gesneriaceae. At the same time, Nautilocalyx, which is usually considered to belong to the Gesneriaceae, may actually belong to the Scrophulariaceae. At one time the Scrophulariaceae was considered to be closely related to the Solanaceae and some elements, e.g., Verbascum (not in Panama) were thought to be transitional between the two families. Several seemingly fundamental features, e.g., absence of internal phloem, presence of iridioid compounds, and regular rather than asymmetrical ovary placement argue against close affinity between these two large but somewhat similar families. The same features just noted support affinity of the Scrophularia- ceae with the Bignoniaceae and Gesneriaceae. Classification of the family tends to recognize two major groups, subfamily Scrophularioideae (Antirrhinoideae) in which the posterior (upper) lip of the corolla is external in bud, and subfamily Rhinanthoideae in which it is internal. The Rhinanthoideae are frequently parasitic and they have a greater tendency to dry dark. The greatest Panamanian development in the family is in tribe Gratioleae subfamily Scrophularioideae, which includes many paludal or aquatic herbs of both New and Old Worlds-in Panama: Bacopa, Benjaminia, Capraria, Leucocarpus, Lindernia, Mecardonia, Micranthemum, and Stemodia. Many species of Scrophulariaceae are valuable horticultural items, especially in temperate gardens, and a few, e.g. Digitalis, yield drugs, but the family is of little economic importance.
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