Flora of China
  FOC Vol.18 (1998)              | Family List  Scrophulariaceae  PDF     | Link Font:+ Big | - Small
   Scrophulariaceae  A. L. Jussieu
玄參科   xuan shen ke+C1130
Authors:Hong De-yuan, Yang Han-bi, Jin Cun-li; Noel H. Holmgren


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.

1Trees 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 length3  Paulownia    泡桐属
+Corolla tube short, upper lip much longer than lower.(3)
3(2)Capsule septicidal; calyx irregularly 3- or 4-lobed4  Wightia    美丽桐属
+Capsule loculicidal; calyx regularly 2- or 5-lobed2  Brandisia    来江藤属
4(1)Leaves abaxially glandular punctate.(5)
+Leaves not glandular punctate.(7)
5(4)Aquatic or marsh herbs15  Limnophila    石龙尾属
+Terrestrial herbs.(6)
6(5)Corolla tube inconspicuous; stamens subequal8  Scoparia    野甘草属
+Corolla tubular; stamens didynamous14  Adenosma    毛麝香属
7(4)Corolla base spurred, lower lip convex and dilated toward upper lip in center, almost sealing throat, making corolla personate27  Linaria    柳穿鱼属
+Corolla neither spurred nor personate.(8)
8(7)Fruit berrylike.(9)
+Fruit capsular or drupaceous (Lagotis).(10)
9(8)Leaves uniform; fruit indehiscent16  Lancea    肉果草属
+Leaves dimorphic; fruit dehiscent32  Hemiphragma    鞭打绣球属
10(8)Plants much branched, diffusely cushionlike; leaves less than 5 mm25  Microcarpaea    小果草属
+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 tuft26  Limosella    水茫草属
+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-valved9  Bacopa    假马齿苋属
+Outermost sepals cordate; anther locules divaricate; stigma 2-lamellate; capsule septicidal17  Picria    苦玄参属
14(12)Pedicel curling after flowering; seeds densely peltate hairy33  Ellisiophyllum    幌菊属
+Pedicel not curling 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 thick5  Scrophularia    玄参属
+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 short6  Oreosolen    藏玄参属
+Flowers solitary; corolla obscurely 2-lipped, subactinomorphic; petiole ca. 2 X as long as leaf blade7  Nathaliella    石玄参属
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 woolly1  Verbascum    毛蕊花属
+Leaves opposite or whorled; corolla 4-merous; filaments glabrous38  Veronica    婆婆纳属
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, indehiscent39  Lagotis    兔耳草属
+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 dilated35  Veronicastrum    腹水草属
+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 alternate36  Scrofella    细穗玄参属
+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 tall37  Pseudolysimachion    穗花属
+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 tall38  Veronica    婆婆纳属
25(20)Leaves basal and rosulate; rhizomes stout; inflorescences spicate34  Neopicrorhiza    胡黄连属
+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 patent47  Castilleja    火焰草属
+Anther essentially 1-loculed; calyx subequally 4-lobed; corolla lower lip shallowly saccate48  Triphysaria    直果草属
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 stem55  Rhinanthus    鼻花属
+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 margin49  Melampyrum    山罗花属
+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 terete58  Pterygiella    翅茎草属
+Calyx tubular, conspicuously 10-ribbed; leaves pinnatifid; stems terete59  Siphonostegia    阴行草属
34(32)Capsule regularly loculicidally dehiscent; sinuses between calyx lobes often 1-3-toothed; corolla yellow, more than 2.5 cm60  Cymbaria    芯芭属
+Capsule loculicidally dehiscent on 1 side (only 1 locule dehiscent); sinuses between calyx lobes toothless; corolla pale purple or white, less than 2 cm61  Monochasma    鹿茸草属
35(31)Calyx 4-lobed; capsule apex obtuse.(36)
+Calyx 5-lobed; capsule apex tapered or truncate.(38)
36(35)Inflorescence a racemose panicle52  Omphalotrix    脐草属
+Inflorescence a spike.(37)
37(36)Bracts often larger than leaves; margin of upper corolla lip revolute51  Euphrasia    小米草属
+Bracts smaller than leaves; margin of upper corolla lip straight54  Odontites    疗齿草属
38(35)Leaves palmatisect; capsule emarginate; upper corolla lip slightly arching forward, obscurely galeate, parted to middle53  Pseudobartsia    五齿萼属
+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 pinnatisect50  Phtheirospermum    松蒿属
+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 serrate56  Xizangia    马松嵩属
+Calyx 2-5-lobed; corolla upper lip galeate and often extending into a narrow beak; leaves pinnatifid to pinnatisect57  Pedicularis    马先蒿属
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; hemiparasitic45  Buchnera    黑草属
+Corolla tube apically curved; inflorescences laxly spicate; all leaves narrow, entire, rarely dentate, sometimes reduced to scales; parasitic46  Striga    独脚金属
44(42)Calyx spathelike, deeply parted on lower side, entire or 3-5-toothed41  Centranthera    胡麻草属
+Calyx campanulate, equally 5-lobed.(45)
45(44)Corolla (3.5-)5-7 cm; calyx 1.5-3 cm30  Rehmannia    地黄属
+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 axils10  Gratiola    水八角属
+Plants more than 25 cm tall; anthers with 1 locule aborted; flowers in racemes44  Sopubia    短冠草属
48(46)Calyx covering corolla tube, inflated after flowering; corolla subcampanulate; at least lower leaves opposite, 3-veined42  Alectra    黑蒴属
+Calyx covering ca. 1/2 of corolla tube, not inflated; corolla tubular; leaves alternate, 1-veined40  Petitmenginia    钟山草属
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-toothed18  Legazpia    三翅萼属
+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 persistent19  Lindernia    母草属
+Calyx 5-winged, rarely conspicuously 5-angular, 2-lipped; capsule septum not persistent20  Torenia    蝴蝶草属
53(51)Flowers solitary in leaf axils, ebracteate; slender and weak herbs21  Mimulus    沟酸浆属
+Flowers in cymes, usually bracteate; large and erect herbs, woody at base22  Cyrtandromoea    囊萼花属
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 cylindric19  Lindernia    母草属
+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 glabrous11  Dopatrium    虻眼属
+Calyx parted almost to base; stems not succulent; filaments twisted; anthers hairy12  Deinostema    泽番椒属
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 hastate31  Triaenophora    崖白菜属
+Leaves glandular hairy; stem leaves present; calyx irregularly toothed or lobes entire.(59)
59(58)Calyx tube present; corolla lips nearly equal in length30  Rehmannia    地黄属
+Calyx parted almost to base; upper corolla lip short, middle lobe of lower lip longest29  Digitalis    毛地黄属
60(57)Stems scaly near base, much branched and broomlike; leaves few, sessile, linear or scalelike, alternate apically on stem; xerophytic24  Dodartia    野胡麻属
+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 numerous28  Mimulicalyx    虾子草属
+Calyx 10-ribbed; seeds 2 per locule43  Leptorhabdos    方茎草属
63(61)Calyx parted almost to base, sometimes less so; capsule often elongated; filaments often appendiculate19  Lindernia    母草属
+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 connective13  Lindenbergia    钟萼草属
+Stems and leaves glabrous or eglandular villous; basal leaves usually well developed, sometimes rosulate, upper stem leaves often alternate; anther locules apically connivent23  Mazus    通泉草属
   List of lower taxa