SCROPHULARIACEAE  

Small, perennial herbs; roots sometimes with red, yellow or orange pigments; stem a short rhizome. Leaves rosulate, petiolate or sessile, obovate, spathulate or broadly ovate to lanceolate, mostly entire and +/- glabrous above, pubescent beneath; venation palmate. Flowers pedicellate, axillary, either in short racemes, usually pedunculate, sometimes branched, sometimes arising from centre of rosette on a +/- sessile inflorescence, or rarely flowers solitary. Bracts usually opposite, lanceolate, oblong or oval. Calyx tubular, 5-toothed, accrescent; tube campanulate or turbinate, sometimes plicate; lobes subequal, pubescent mainly on veins or +/- pilose, shorter than tube. Corolla bilabiate, 4- or 5-lobed; tube funnel-shaped, pubescent within throat; posterior lip hooded, entire or shortly bilobed, exterior in bud; anterior lip 3-lobed, longer than posterior. Stamens 4, didynamous; upper pair included under hood, with filaments short, straight, arising at top of corolla tube; lower pair with filaments longer, arising +/- in throat, geniculate and with yellow appendage at base, bent forwards and inwards above; anthers bithecate, approximating or cohering in pairs; thecae divaricate; staminodes 0. Nectary 0. Ovary bilocular, ellipsoid or elliptical; ovules many; style filiform, +/- exserted, persistent; stigma capitate, of 2 membranous lobes, papillose, touch-sensitive. Fruit an ovoid or ellipsoid, septicidal capsule. Seeds globose; testa thin, brown, reticulate and punctate, epidermal cells thin-walled; endosperm alveolate and bothrospermous (with small hollows and nonfurrowed).

Small annual herbs; stems erect, decumbent or ascending; brachyblasts 0; hairs on stems, leaves, bracts and calyx thin-walled. Leaves opposite, bases often connate, sometimes becoming alternate upwards, variously shaped, sometimes tapering into petiole. Flowers sessile, in a narrow spike, or several forming a panicle. Bract adnate to part of calyx tube. Calyx 5-lobed or 5-toothed, campanulate; anterior lip sometimes only very shallowly divided. Corolla tubular, uni- or bilabiate, 4- or 5-lobed; tube cylindrical or funnel-shaped; posterior lip 4-lobed, sometimes bearded with minute clavate hairs, without yellow/orange patch, exterior in bud; anterior lip 1-lobed, minute, or lobe wanting. Stamens 4, didynamous, sometimes 2 being somewhat abnormal, or 2 only (anterior pair); filaments arising in upper part of corolla tube or in mouth, shortly decurrent on tube; anthers synthecate, dorsifixed, +/- exserted. Nectary annular but more strongly developed on one side. Ovary bilocular, elliptic-oblong, base oblique; ovule 1 per locule, pendulous; style filiform; stigma lingulate with 2 marginal bands of stigmatic papillae, malformed when flowers autogamous. Fruit a pair of hard-walled cocci, convex on outer face with median longitudinal groove, rugose on either side of groove, +/- plane on inner face, indehiscent. Seeds +/- fusiform, smooth or transversely rugose.

Perennial shrublets or herbs, flowering sometimes as seedlings; stems brittle, leafy, very glandular-pubescent, sometimes woody. Leaves mostly alternate, opposite only at extreme base of seedlings, petiolate, broadly ovate, serrate. Flowers pedicellate, sometimes bracteate and 1- or 2-bracteolate, in axils of upper leaves, solitary, paired or in cymules with up to 11 flowers, forming leafy racemes or narrow panicles. Bracts, if present, very small, free from pedicels. Calyx 5-lobed nearly to base, hardly bilabiate; lobes elliptic-spathulate. Corolla slightly irregular, 5-lobed; tube cylindrical, scarcely expanded at apex, glabrous outside, throat bearded inside with broad band of clavate hairs; upper lip 2-lobed with lobes joined for +/- half their length, exterior in bud; lower lip 3-lobed. Stamens 4, didynamous; posterior pair arising +/- halfway up tube, included, with filaments decurrent to tube base, forming channel for style; anterior pair arising in throat, very shortly exserted; all filaments bearded with long clavate hairs or not, minutely glandular at apex; anthers synthecate, dorsifixed, anterior ones often slightly larger than posterior; staminodes 0. Nectary ill-defined at base of ovary, but probably annular, either convoluted or swollen on one side. Ovary bilocular, cuneate, tapering into style, glabrous; ovules many per locule; stigm a very short, held in throat or very shortly exserted; stigmatic papillae in a lateral groove carried over apex, tip sometimes very shortly bifid. Fruit a sharply beaked, septicidal capsule, tip of each valve with a short loculicidal split. Seeds very small, red-brown, oblong, with 6-8 longitudinal ribs (aulacospermous); testa reticulate in furrows between ribs.

Annual or perennial, often hemiparasitic herbs, usually somewhat rigid and scabrid, turning dark on drying; stems erect, ascending or prostrate, simple or branched, glabrous to hispid-scabrid. Leaves usually opposite or quasiverticillate, sometimes subopposite or alternate above, sessile to shortly petiolate, 1-5-veined, lowest ones usually obovate, entire or dentate, upper ones narrow, entire or denticulate, gradually passing into bracts. Flowers often small, sessile or subsessile in axils of upper leaves or bracts, usually bibracteolate, solitary, in terminal, compact to lax spikes or in terminal, or axillary, +/- globose clusters; inflorescences often lengthen in fruit. Calyx tubular, 4- or 5-toothed, tube cylindrical, narrow, 5- or 10(7 or 8)-veined or -ribbed; teeth short, deltoid to lanceolate-acuminate, usually scabrid on margin. Corolla tubular, 4- or 5(6)-lobed, slightly irregular; tube slender cylindrical, longer than calyx, straight or gently curved, glabrous to softly pilose, throat sometimes densely pilose; limb shorter than tube, spreading; lobes subequal, obovate to subcircular or oblong, entire or emarginate, 2 posterior ones often a little smaller, interior in bud. Stamens 4, slightly didynamous, subequal, arising about middle of corolla tube, subsessile or with short filaments, included; anthers unithecate, erect, dorsifixed, acute, blunt at base. Ovary bilocular, elliptical; ovules many per locule; style terete, straight; stigma thickened or clavate, entire or emarginate, almost as long as style, included. Fruit an ovoid to oblong, loculicidal capsule. Seeds many, obovoid or oblong; testa membranous, reticulate, tightly investing endosperm. x = 10 (polyploidy).

Annual, viscid-pubescent herbs, parasitic on grass roots; stems simple or slightly branched in inflorescence, erect, leafy. Leaves opposite, sessile or lowest shortly petiolate, narrow-lanceolate, obtuse, coarsely toothed or shallowly pinnatifid, spreading. Flowers sessile, in axils of leaves or bracts, in dense terminal spikes, usually unbranched, elongated in fruit. Bracts almost as long as flowers. Calyx ovoid, somewhat inflated, tetragonous, shortly 4-lobed, densely tomentose; tube campanulate; lobes deltoid, obtuse or rounded. Corolla tubular, bilabiate, almost personate, 4-lobed; tube subcylindrical, +/- as long as calyx tube, slightly curved; upper lip erect, narrow, hooded, concave, entire, rather shorter than lower one, inside in bud; lower lip dilated, spreading, 3-lobed, middle lobe the smallest; palate bigibbous, prominent. Stamens 4, all fertile, didynamous, slightly exserted; upper pair arising at base of upper corolla lip, lower pair in lower part of corolla tube; all lying in hollow of upper lip; filaments filiform; anthers free, pendulous, bithecate; thecae oblong, shortly mucronulate at base, equal, parallel, +/- bearded or at length glabrous. Ovary bilocular, ovoid, hirsute; ovules many, on thick, bifid placentas; style linear, exceeding stamens, +/- exserted, curved downwards near apex, hairy; stigma short, somewhat thickened, clavate, bifid. Fruit a somewhat flattened, ovoid capsule, coriaceous, pubescent, turgid, loculicidal, bisulcate and enclosed in calyx. Seeds many, small, convex on one side, flattish on other, longitudinally ribbed; testa membranous. x = 6 (7) (aneuploids, high polyploidy).