SCROPHULARIACEAE  

Annual or rarely perennial herbs, holoparasitic on roots, lacking chlorophyll, coloured but mostly drying dark; stems erect or ascending, simple or branched mostly at base, usually angular, often furrowed. Leaves usually scale-like, at least lower ones, opposite or subopposite, sometimes alternate above, often close or crowded at base and upper ones more distant to scattered, sometimes few. Flowers usually large, +/- sessile to pedicellate, axillary, bracteate and bibracteolate, in terminal, lax or dense, few- to many-flowered racemes or spikes, sometimes elongating in fruit. Bracts opposite or uppermost alternate; bracteoles arising above middle to near or at pedicel apex, rarely adnate to calyx. Calyx tubular, 5-lobed, sometimes bilabiate, pilose or glandular-hairy; tube campanulate or oblong, rarely dimidiate, somewhat inflated or lax, angular or ribbed; lobes unequal, usually longer than tube, valvate in bud. Corolla tubular, 5-lobed, somewhat bilabiate, various colours but if yellow, veins not dark; tube gently curved, usually narrow, globose at base and much enlarged, subcylindrical or funnel-shaped above, or ventricosely inflated; limb oblique or nearly regular, spreading or obliquely cup-shaped; lobes rounded or broadly obovate, entire or denticulate; upper 2 lobes interior in bud, sometimes connate higher up than others. Stamens 4, didynamous or +/- equal, included or scarcely exserted; filaments linear, arising about or below middle of corolla tube; anthers approximated in pairs, bithecate; thecae parallel or transverse, unequal, one fertile, acuminate, acute or mucronate at base, the other barren, longer and subulate-acuminate, or rarely subobsolete. Ovary bilocular, obovate, elliptic or ovate; placentation axile; ovules very many; style filiform, somewhat flattened, +/- as long as corolla tube, curved downwards near apex when ready for pollen; stigma thickened and various in shape. Fruit a globose, ovoid or conical, somewhat compressed, 2-grooved, loculicidal capsule. Seeds very many, small, irregularly oblong, truncate; testa reticulate, loose, membranous.

Compact, perennial, semiwoody herbs or undershrubs, many-stemmed from a woody rootstock; stems erect or +/- trailing, usually branched at or near base, leafy. Leaves opposite, alternate, or whorled, ovate-elliptic to lanceolate or linear, incised-dentate to entire. Flowers solitary in axils of upper leaves, subsessile to shortly pedicellate, bibracteolate at or near base of calyx. Calyx tubular, 5-lobed; tube campanulate or funnel-shaped, 10-nerved; lobes +/- equal, triangular-lanceolate, equalling or rather longer than tube, somewhat accrescent. Corolla tubular, somewhat bilabiate, 5-lobed, pilose; tube funnel-shaped, somewhat ventricose, exserted; limb spreading; lobes obovate, entire, subequal, the 2 posterior ones interior, shorter than tube. Stamens 4, didynamous, included; posterior pair short, arising below middle of corolla tube; anterior pair longer, arising near base of tube; filaments filiform, hairy at least towards base; anthers bithecate, free; thecae separate, diverging, mucronulate at base, with one usually narrower than other. Ovary bilocular, ovate to globular, compressed; ovules many per locule; style filiform, glabrous, somewhat exserted, incurved above, persistent; stigma short, bifid, sometimes also slightly thickened or subglobose. Fruit an acute or acuminate, loculicidal capsule, flattened longitudinally, therefore appearing unilaterally winged, included in persistent calyx. Seeds many, obovoid-cylindrical; testa membranous, reticulate.

Perennial herbs or dwarf shrublets; stems leafy throughout, pubescent, hairs retrorse and minutely glandular. Leaves opposite below, alternate above, simple, variously elliptical and oblong to oblanceolate, often narrowed to petiole-like, sometimes half-clasping or connate base, usually toothed, thick or leathery, glandular-punctate, sometimes also with glandular and/or eglandular hairs; radical leaves sometimes present. Flowers in a terminal, cone-shaped, very congested head, nodding in flower, erect and elongated in fruit. Bracts broad, clearly distinct from leaves, adnate to pedicel, free from calyx. Calyx unequally 5-lobed, rarely 2 anterior lobes obsolete, split nearly or quite to base at least on anterior side, thin-textured. Corolla slightly bilabiate, glabrous, thick, leathery when dry; tube funnel-shaped; posterior lobes exterior in bud, lobes spreading, oblong to suborbicular, entire, margins pale, thin-textured. Stamens 4, didynamous, arising near top of corolla tube, 2 or 4 exserted; posterior filaments decurrent +/- to base of tube; anthers synthecate, dorsifixed; staminodes 0. Nectary at base of ovary on posterior side. Ovary bilocular, +/- oblong; ovules 7-many per locule; stigma lingulate, flattened, with 2 marginal bands of stigmatic papillae, exserted. Fruit a septicidal capsule with a short loculicidal split at tip of each valve, glabrous. Seeds angled by pressure, sinuously colliculate, reddish brown. x = 7.

Twiggy shrublets; stems leafy, pubescence confined to two narrow ridges spanning internodes, hairs +/- retrorse, eglandular, but sometimes minute, patent, glandular. Leaves (and bracts) usually opposite and decussate, rarely subopposite, elliptical, gradually narrowed below into very short petiolar part, bases connate and decurrent in two narrow ridges, finely callose-toothed, thick, glandular-punctate and partly puberulous. Flowers solitary in upper leaf axils, forming terminal, racemose, round heads, oblong in fruit, sometimes further arranged in corymbose panicles. Bracts adnate to pedicel and halfway up calyx tube. Calyx bilabiate, 5-lobed, membranous; lobes unequal, ciliate. Corolla bilabiate, thick-textured, not persistent; tube cylindrical below, abruptly campanulate above, glabrous outside, inside heavily bearded round mouth with unicellular clavate hairs; posterior lip 2-lobed with orange patch at base extending down inflated part of tube, exterior in bud; anterior lip 3-lobed; lobes entire, rotund. Stamens 4, didynamous; filaments short, arising in throat, not decurrent down tube; posterior pair lying in throat; anterior pair exceeding posterior pair, shortly exserted; anthers dorsifixed, synthecate; staminodes 0. Nectary at base of ovary, on shorter side. Ovary bilocular, elliptical; ovules +/- 9-16 per locule; style long, filiform, passing gradually into lingulate stigma with two marginal bands of stigmatic papillae, exserted. Fruit a septicidal capsule with a short loculicidal split at tip of each valve. Seeds obscurely 3-angled or flattened, irregularly wrinkled on one face; testa tightly investing endosperm, opaque, creamy or pale amber.

Shrubs or subshrubs; branches usually glabrous with ribs due to decurrent leaf bases. Leaves opposite, verticillate or scattered, usually variously linear or lanceolate to ovate, sometimes obovate or oblanceolate, sessile to shortly petiolate, elliptical or mostly acute, often apiculate or mucronate, entire, or rarely toothed in upper half, flat, undulate or crisped, base sometimes decurrent, midrib usually prominent, often leathery. Flowers solitary or in terminal and axillary cymes, shortly pedicellate, usually bracteate and bracteolate. Bracts usually similar to leaves but smaller. Calyx campanulate, 5-lobed; lobes ovate, elliptic or lanceolate, imbricate. Corolla 5-lobed, regular; tube funnel-shaped or subcylindrical, straight, exceeding calyx; limb patent, regular; lobes much shorter than tube, spreading, obtuse, imbricate, 2 posterior lobes exterior in bud. Stamens 4, didynamous, included or scarcely exserted; filaments terete or linear, arising +/- at or above middle of corolla tube; anthers with 2 +/- parallel, erect thecae; staminodes sometimes 1, rudimentary. Nectary saucer- or cushion-shaped. Ovary bilocular, ellipsoid; ovules several; style thick, linear, scarcely exserted, with capitate or globose stigma. Fruit an ovoid, obtuse, septicidal capsule; valves bifid. Seeds discoid with a membranous wing and testa.