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Scrophulariaceae A. L. Jussieu

玄參科 xuan shen ke+C1130

Authors: Deyuan Hong, Hanbi Yang, Cun-li Jin, Manfred A. Fischer, Noel H. Holmgren & Robert R. Mill

Lancea tibetica

Credit: Harvard University Herbaria

Herbs, sometimes shrubs, rarely trees; mostly autotrophic, less often hemiparasitic or parasitic. Stipules absent. Leaves alternate, opposite, whorled, or basally opposite and apically alternate, simple or sometimes pinnately dissected. Inflorescences racemes, spikes, or thyrsoid panicles, determinate or indeterminate, or flowers solitary. Flowers perfect, usually zygomorphic, rarely actinomorphic. Calyx often persistent, (2-)4- or 5-lobed or -parted, variously connate. Corolla sympetalous; limb (3 or)4- or 5-lobed, often 2-lipped. Stamens mostly 4, didynamous, sometimes 1 or 2 staminodes present, less often 2 or 5 stamens; anther locules 1 or 2, equal or subequal, free or confluent. Nectary often present at base of ovary, ringlike, cupular, or reduced to a gland. Ovary superior, 2-loculed, rarely apically 1-loculed; ovules numerous, rarely 2 per locule, on axile placentas, anatropous or hemitropous. Style simple; stigma capitate, 2-lobed, or 2-lamellate. Fruit a capsule, septicidal, loculicidal, or septifragal, sometimes opening by pores or irregularly dehiscent, rarely a berry. Seeds minute or rarely conspicuous, sometimes winged; testa often reticulate; hilum lateral or ventral; endosperm fleshy or absent; embryo straight or curved.

About 220 genera and 4500 species: cosmopolitan; 61 genera (seven endemic) and 681 species (415 endemic) throughout China but with a greater concentration in SW China.

Tsoong Puchiu & Yang Hanbi, eds. 1979. Scrophulariaceae (1). Fl. Reipubl. Popularis Sin. 67(2): 1-431. Tsoong Puchiu, ed. 1963. Scrophulariaceae (2). Fl. Reipulb. Popularis Sin. 68: 1-449.

1 Trees or shrubs; calyx leathery; stems and leaves often stellate hairy when young.   (2)
+ Herbs, sometimes woody at base; calyx herbaceous or membranous; stems and leaves without stellate hairs.   (4)
2 (1) Corolla tube long, both lips equal in length   3 Paulownia
+ Corolla tube short, upper lip much longer than lower.   (3)
3 (2) Capsule septicidal; calyx irregularly 3- or 4-lobed   4 Wightia
+ Capsule loculicidal; calyx regularly 2- or 5-lobed   2 Brandisia
4 (1) Leaves abaxially glandular punctate.   (5)
+ Leaves not glandular punctate.   (7)
5 (4) Aquatic or marsh herbs   15 Limnophila
+ Terrestrial herbs.   (6)
6 (5) Corolla tube inconspicuous; stamens subequal   8 Scoparia
+ Corolla tubular; stamens didynamous   14 Adenosma
7 (4) Corolla base spurred, lower lip convex and dilated toward upper lip in center, almost sealing throat, making corolla personate   27 Linaria
+ Corolla neither spurred nor personate.   (8)
8 (7) Fruit berrylike.   (9)
+ Fruit capsular or drupaceous (Lagotis).   (10)
9 (8) Leaves uniform; fruit indehiscent   16 Lancea
+ Leaves dimorphic; fruit dehiscent   32 Hemiphragma
10 (8) Plants much branched, diffusely cushionlike; leaves less than 5 mm   25 Microcarpaea
+ Plants erect to creeping, never cushionlike; leaves usually longer than 5 mm.   (11)
11 (10) Plants sometimes floating; stolons creeping; leaves and flowers in a basal tuft   26 Limosella
+ Plants aquatic or terrestrial, never floating; stems creeping and also erect; leaves and flowers never in a basal tuft.   (12)
12 (11) Sepals free, 4 or 5, imbricate, unequal in width.   (13)
+ Sepals connate to almost free, not imbricate at least after flowering, equal in width.   (14)
13 (12) Outermost sepals not cordate; anther locules parallel; stigma dilated, capitate, or 2-lobed; capsule loculicidal or 4-valved   9 Bacopa
+ Outermost sepals cordate; anther locules divaricate; stigma 2-lamellate; capsule septicidal   17 Picria
14 (12) Pedicel coiling after flowering; seeds densely peltate hairy   33 Ellisiophyllum
+ Pedicel not coiling after flowering; seeds glabrous.   (15)
15 (14) Staminode 1, in center of corolla upper lip; anther locules confluent, reniform and transverse; filaments apically dilated.   (16)
+ Staminodes 2 if present, anterior; anthers not transverse (except in some Gratiola and Verbascum); filaments not dilated.   (18)
16 (15) Plants more than 10 cm tall; flowers in cymes, rarely axillary, never fascicled; pedicel conspicuous; corolla tube short and thick   5 Scrophularia
+ Plants less than 10 cm tall; flowers solitary or fascicled in axils; pedicel inconspicuous; corolla tube elongated.   (17)
17 (16) Flowers in fascicles; corolla conspicuously 2-lipped; petiole short   6 Oreosolen
+ Flowers solitary; corolla obscurely 2-lipped, subactinomorphic; petiole ca. 2 X as long as leaf blade   7 Nathaliella
18 (15) Corolla rotate, almost without a tube.   (19)
+ Corolla not rotate, if sometimes subrotate then stamens only 2 and tube conspicuous.   (20)
19 (18) Leaves alternate; flowers 5-merous; filaments woolly   1 Verbascum
+ Leaves opposite or whorled; corolla 4-merous; filaments glabrous   38 Veronica
20 (18) Stamens 2, staminodes absent; corolla (3 or)4-lobed, actinomorphic or slightly zygomorphic, if 2-lipped, then lower lip often 1- or 2-lobed.   (21)
+ Stamens 4 or if 2, then 2 staminodes present abaxially; corolla conspicuously 2-lipped, lower lip 3-lobed, upper 2-lobed or entire, or 5-lobed and subactinomorphic (corolla 4-lobed in Neopicrorhiza but then basal leaves rosulate).   (25)
21 (20) Calyx parted to base on lower side, spathelike or sometimes also 2-lobed adaxially; leaves mostly basal; fruit drupaceous, indehiscent   39 Lagotis
+ Calyx 4- or 5-lobed; leaves on stem; fruit a capsule.   (22)
22 (21) Calyx lobes 5, subequal in length; stamens exserted; stigma small, weakly dilated   35 Veronicastrum
+ Calyx lobes 4 or if 5 then upper one much smaller than other lobes; stamens included in corolla; stigma capitate or clavate.   (23)
23 (22) Corolla limb conspicuously 2-lipped with strongly reflexed lower lip, tube conspicuous; leaves alternate   36 Scrofella
+ Corolla limb weakly 2-lipped, often subrotate, tube usually weakly developed; leaves opposite, whorled, or alternate.   (24)
24 (23) Racemes terminal, long and densely spicate; bracts small and narrow; capsule subglobose, slightly compressed; corolla with conspicuous tube and slightly 2-lipped limb; stems mostly more than 30 cm tall   37 Pseudolysimachion
+ Racemes axillary or terminal, usually short and lax, if raceme terminal then lower bracts similar to leaves; capsule usually strongly compressed; corolla mostly with short tube and subrotate limb; stems mostly less than 25 cm tall   38 Veronica
25 (20) Leaves basal and rosulate; rhizomes stout; inflorescences spicate   34 Neopicrorhiza
+ Leaves on stems and also basal (Triaenophora scapose but leaves very large); rhizomes slender if present; inflorescences mostly racemose.   (26)
26 (25) Upper corolla lip galeate or rostrate; anther locules mucronate or base spurred.   (27)
+ Upper corolla lip straight or revolute, never galeate; anther locules rarely mucronate, base often obtuse.   (41)
27 (26) Leaves alternate; upper corolla lip rostrate, conspicuously longer than lower lip; anther locules unequal in length or free, or anthers 1-loculed.   (28)
+ Leaves opposite; upper corolla lip galeate, as long as lower lip; anther locules equal in length, connivent.   (29)
28 (27) Anther locules 2, unequal in length; calyx unequally 4-lobed, parted to middle medianly, more shallowly laterally; corolla lower lip lobes patent   47 Castilleja
+ Anther essentially 1-loculed; calyx subequally 4-lobed; corolla lower lip shallowly saccate   48 Triphysaria
29 (27) Calyx upper side parted to middle, lower side and laterally shallowly parted, strongly inflated and utriculate in fruit; seeds compressed, often winged; branches and leaves subappressed to main stem   55 Rhinanthus
+ Calyx equally parted or more deeply parted medianly, not inflated in fruit; seeds not compressed; branches and leaves spreading.   (30)
30 (29) Capsule 1-4-seeded; seeds large and smooth; bracts dentate to setose dentate, rarely entire; upper corolla lip densely bearded at margin   49 Melampyrum
+ Capsule with numerous seeds; seeds small and sculptured; bracts often entire; upper corolla lip not bearded at margin.   (31)
31 (30) Bracteoles 2 below calyx.   (32)
+ Bracteoles absent.   (35)
32 (31) Basal leaves well developed.   (33)
+ Basal leaves scalelike.   (34)
33 (32) Calyx ovoid, conspicuously 5-ribbed; leaves entire or dentate; stems often winged, rarely terete   58 Pterygiella
+ Calyx tubular, conspicuously 10-ribbed; leaves pinnatifid; stems terete   59 Siphonostegia
34 (32) Capsule regularly loculicidally dehiscent; sinuses between calyx lobes often 1-3-toothed; corolla yellow, more than 2.5 cm   60 Cymbaria
+ Capsule loculicidally dehiscent on 1 side (only 1 locule dehiscent); sinuses between calyx lobes toothless; corolla pale purple or white, less than 2 cm   61 Monochasma
35 (31) Calyx 4-lobed; capsule apex obtuse.   (36)
+ Calyx 5-lobed; capsule apex tapered or truncate.   (38)
36 (35) Inflorescence a racemose panicle   52 Omphalotrix
+ Inflorescence a spike.   (37)
37 (36) Bracts often larger than leaves; margin of upper corolla lip revolute   51 Euphrasia
+ Bracts smaller than leaves; margin of upper corolla lip straight   54 Odontites
38 (35) Leaves palmatisect; capsule emarginate; upper corolla lip slightly arching forward, obscurely galeate, parted to middle   53 Pseudobartsia
+ Leaves pinnatisect, pinnatifid, or serrate; capsule apex tapered; upper corolla lip conspicuously galeate, entire or shallowly 2-lobed.   (39)
39 (38) Calyx equally parted; leaves pinnatisect   50 Phtheirospermum
+ Calyx more deeply parted on lower side; leaves serrate, pinnatifid, or pinnatisect.   (40)
40 (39) Calyx 5-lobed; corolla upper lip revolute, not extending into a beak; leaves serrate   56 Xizangia
+ Calyx 2-5-lobed; corolla upper lip galeate and often extending into a narrow beak; leaves pinnatifid to pinnatisect   57 Pedicularis
41 (26) Bracteoles 2; corolla subactinomorphic or 2-lipped in Rehmannia and Gratiola.   (42)
+ Bracteoles absent; corolla conspicuously 2-lipped.   (49)
42 (41) Corolla salverform; fertile anthers with 1 locule aborted.   (43)
+ Corolla not salverform; fertile anthers 2-loculed.   (44)
43 (42) Corolla tube straight; inflorescences spicate, flowers crowded; lower leaves broad and dentate, upper narrow and entire; hemiparasitic   45 Buchnera
+ Corolla tube apically curved; inflorescences laxly spicate; all leaves narrow, entire, rarely dentate, sometimes reduced to scales; parasitic   46 Striga
44 (42) Calyx spathelike, deeply parted on lower side, entire or 3-5-toothed   41 Centranthera
+ Calyx campanulate, equally 5-lobed.   (45)
45 (44) Corolla (3.5-)5-7 cm; calyx 1.5-3 cm   30 Rehmannia
+ Corolla less than 1 cm; calyx less than 0.5 cm.   (46)
46 (45) Fertile anthers 2.   (47)
+ Fertile anthers 4.   (48)
47 (46) Plants less than 25 cm tall; anthers 2-loculed; flowers solitary in leaf axils   10 Gratiola
+ Plants more than 25 cm tall; anthers with 1 locule aborted; flowers in racemes   44 Sopubia
48 (46) Calyx covering corolla tube, inflated after flowering; corolla subcampanulate; at least lower leaves opposite, 3-veined   42 Alectra
+ Calyx covering ca. 1/2 of corolla tube, not inflated; corolla tubular; leaves alternate, 1-veined   40 Petitmenginia
49 (41) Calyx 3- or 5-winged or conspicuously 3- or 5-angular, toothed.   (50)
+ Calyx neither winged nor conspicuously angular, parted into 5 conspicuous lobes, lobes sometimes secondarily lobed.   (54)
50 (49) Calyx broadly 3-winged, 3-toothed   18 Legazpia
+ Calyx 5-winged or 5-angled, 5-toothed.   (51)
51 (50) Calyx not inflated in fruit, mouth not truncate; filaments often appendiculate.   (52)
+ Calyx inflated, bladderlike in fruit, mouth truncate or obliquely truncate; filaments not appendiculate.   (53)
52 (51) Calyx 5-angular, actinomorphic; capsule septum persistent   19 Lindernia
+ Calyx 5-winged, rarely conspicuously 5-angular, 2-lipped; capsule septum not persistent   20 Torenia
53 (51) Flowers solitary in leaf axils, ebracteate; slender and weak herbs   21 Mimulus
+ Flowers in cymes, usually bracteate; large and erect herbs, woody at base   22 Cyrtandromoea
54 (49) Fertile stamens 2, staminodes 2; aquatic or marsh herbs.   (55)
+ Fertile stamens 4; terrestrial herbs (some species of Mimulicalyx and Lindernia aquatic or marsh herbs).   (57)
55 (54) Leaves petiolate, dentate; capsule cylindric   19 Lindernia
+ Leaves sessile and entire; capsule ovoid-globose or globose.   (56)
56 (55) Calyx parted to slightly beyond middle; stems succulent; filaments straight at apex; anthers glabrous   11 Dopatrium
+ Calyx parted almost to base; stems not succulent; filaments twisted; anthers hairy   12 Deinostema
57 (54) Corolla more than 3 cm; petiole long; stem leaves alternate or absent.   (58)
+ Corolla less than 3 cm; petiole usually short; at least lower stem leaves opposite.   (60)
58 (57) Leaves woolly; stem leaves absent; calyx 5-lobed, each lobe 3-parted and hastate   31 Triaenophora
+ Leaves glandular hairy; stem leaves present; calyx irregularly toothed or lobes entire.   (59)
59 (58) Calyx tube present; corolla lips nearly equal in length   30 Rehmannia
+ Calyx parted almost to base; upper corolla lip short, middle lobe of lower lip longest   29 Digitalis
60 (57) Stems scaly near base, much branched and broomlike; leaves few, sessile, linear or scalelike, alternate apically on stem; xerophytic   24 Dodartia
+ None with above combination of characters; mesophytic or helophytic.   (61)
61 (60) Stems quadrangular; lower corolla lip outer in bud, enveloping upper one.   (62)
+ Stems terete (quadrangular in Mazus lanceifolius); upper corolla lip outer in bud, enveloping lower one.   (63)
62 (61) Calyx 5-ribbed; seeds numerous   28 Mimulicalyx
+ Calyx 10-ribbed; seeds 2 per locule   43 Leptorhabdos
63 (61) Calyx parted almost to base, sometimes less so; capsule often elongated; filaments often appendiculate   19 Lindernia
+ Calyx parted to about middle; capsule short; filaments not appendiculate.   (64)
64 (63) Stems and leaves glandular hairy; stem leaves opposite; anther locules separated by a stalked connective   13 Lindenbergia
+ Stems and leaves glabrous or eglandular villous; basal leaves usually well developed, sometimes rosulate, upper stem leaves often alternate; anther locules apically connivent   23 Mazus

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