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Gesneriaceae Dumortier

苦苣苔科 ku ju tai ke

Authors: Wencai Wang, Kai-yu Pan, Zhen-yu Li, Anna L. Weitzman & Laurence E. Skog

Herbs, shrubs, or rarely trees. Leaves opposite or rarely alternate, whorled or basal, rosette forming; exstipulate; usually simple, rarely shallowly to deeply lobed, pinnately or rarely palmately veined. Inflorescences usually cymes, rarely racemes, axillary, often near apex and appearing terminal; usually pedunculate. Flowers perfect, zygomorphic, seldom actinomorphic. Calyx actinomorphic, rarely zygomorphic; usually (4 or)5-divided. Corolla gamopetalous, zygomorphic, rarely actinomorphic; usually 2-lipped. Fertile stamens 2 or 4, then often didynamous, rarely 5, epipetalous; anthers free or coherent, thecae 2, parallel, divergent, or divaricate; staminodes 1-3 or absent. Disc ringlike to cupular, rarely absent. Ovary superior in all Old World taxa [half inferior, or inferior], 1-loculed; gynophore seldom present; placentas (1 or)2, parietal, rarely 2-loculed, placenta 1 per locule and axile; ovules numerous, anatropous. Style 1; stigmas 1 or 2. Fruit usually capsular, loculicidal, septicidal, or circumscissile, rarely a berry, indehiscent. Seeds numerous, fusiform to ellipsoid or ovoid, minute, sometimes with appendages at 1 or both ends, with or without endosperm; embryo straight, cotyledons equal or unequal after germination.

About 133 genera and 3000 species: Africa, Central and South America, E and S Asia, S Europe, Oceania; 56 genera (25 endemic) and 442 species (354 endemic) in China.

A few foreign well-known ornamental species are cultivated in China, including the florist's gloxinia, Sinningia speciosa (Loddiges) Hiern, and African violet, Saintpaulia ionantha Wendland.

The two ovary carpels may each produce a stigma; these stigmas are fused into a single structure. Some students of Gesneriaceae have considered the stigma to be single and either simple (capitate) or 2-lobed, whereas others consider each of the two stigmas as units. We have maintained the latter usage, but a family-wide investigation of stigma development is needed. The distinction can be blurred, however, because the stigmas may be completely fused into one with a capitate apex (as in Didymocarpus) or one of the two carpels or stigmas may be aborted resulting in a single stigma that may or may not be 2-lobed.

Certain characters used in this treatment require some explanation. Disc shape is frequently described as "ringlike," "cupular," or rarely "tubular"; ringlike describes a disc that is shorter than or equalling its diameter, and cupular describes one that is longer than its diameter. Tubular describes a disc which is at least twice as long as wide. The corolla tube diameter measurement is for the widest part, usually the mouth.

Wang Wentsai, Pan Kaiyu, & Li Zhenyu. 1990. Gesneriaceae. In: Wang Wentsai, ed., Fl. Reipubl. Popularis Sin. 69: 125-581.


1 Inflorescences racemose; seeds with scalelike appendages at both ends; flowers sometimes replaced by propagules   56 Titanotrichum
+ Flowers solitary or inflorescences cymose (including umbel-like or paniculate cymes), sometimes racemelike and 1-sided; seeds either not appendiculate or with subulate or hairlike appendages at 1 or both ends (unknown in Calcareoboea. Dayaoshania, Gyrogyne. Lagarosolen. Paraisometrum,and Thamnocharis); flowers never replaced by propagules.   (2)
       
2 (1) Corolla actinomorphic, 4- or 5-lobed; stamens 4 or 5, staminodes absent; seeds unappendaged (unknown in Thamnocharis).   (3)
+ Corolla zygomorphic, usually distinctly 2-lipped or rarely actinomorphic; fertile stamens (1 or)2 or 4, staminodes absent or 1-3; seeds appendaged or unappendaged.   (6)
       
3 (2) Corolla suburceolate, tube longer than limb; stamens included, anther thecae divergent, confluent apically; capsule 4-valved   3 Tengia
+ Corolla rotate or campanulate, tube 1/3-1 X limb; stamens equalling corolla or exserted, anther thecae parallel, not confluent; capsule 2-valved.   (4)
       
4 (3) Anthers connate into a tube surrounding style, connectives with long apical projections; disc absent; inflorescences umbel-like cymes; leaves 1 to few   4 Conandron
+ Anthers free, connectives without projections; disc ringlike; inflorescences cymes; leaves few to many.   (5)
       
5 (4) Bracts 2, opposite; corolla purple to blue, deeply 4- or 5-lobed, tube rotate; stamens adnate to corolla near base, anthers basifixed; stigma 1, nearly truncate or punctiform; leaf blade base cuneate   1 Thamnocharis
+ Bracts 6-9 to numerous, crowded, involucrate; corolla white, 4- or 5-lobed, tube campanulate; stamens adnate to corolla tube near or above middle, anthers dorsifixed; stigmas 2, equal, oblate to 2-lipped; leaf blade base truncate to cordate   2 Bournea
       
6 (2) Fruit indehiscent berries, fleshy to leathery; plants woody herbs or shrubs.   (7)
+ Fruit dehiscent capsule; plants herbaceous to shrubby.   (8)
       
7 (6) Calyx divided from base; corolla campanulate-tubular; stamens 4   49 Rhynchotechum
+ Calyx divided only from above middle (in China); corolla funnelform-tubular; stamens 2 (in China)   50 Cyrtandra
       
8 (6) Calyx plicate between lobes; corolla tube campanulate, spurred or saccate at base; stamens 4; ovary depressed globose or subglobose.   (9)
+ Calyx not plicate; corolla tube funnelform to tubular, cylindric, salverform, urceolate, or campanulate, base usually not swollen, rarely saccate, sometimes gibbous abaxially; stamens (1 or)2 or 4; ovary linear to oblong or ovoid, rarely subglobose or globose.   (10)
       
9 (8) Leaves slightly unequal in a pair; adaxial corolla lip ca. 1/2 as long as abaxial lip; anthers free, thecae not confluent   51 Gyrogyne
+ Leaves strongly unequal in a pair, smaller much reduced; adaxial corolla lip as long as abaxial lip; anthers coherent laterally to form a shallow cone, thecae confluent at apex   52 Stauranthera
       
10 (8) Capsule ovoid to depressed globose, shorter than calyx; ovary ovoid to depressed globose; inflorescences cymose, sympodial, or scorpioid, racemose; bract apparently absent or 1; stem leaves 1 to few.   (11)
+ Capsule linear to ovoid, much surpassing calyx, rarely shorter than calyx, then leaves all basal (fruit unknown in Paraisometrum); ovary usually linear to oblong, rarely ellipsoid, linear-lanceolate, conical, or ovoid; flowers solitary or inflorescences cymose including paniculate or umbel-like cymes; bracts 2, rarely apparently absent, 1, or 3; leaves basal or on stem, few to many.   (13)
       
11 (10) Leaves opposite, strongly unequal in a pair, smaller much reduced and sessile; inflorescences apparently without bracts; calyx with many longitudinal secretory canals; staminode 1; ovary 2-loculed; placenta 1, axile   53 Whytockia
+ Leaves alternate, or lowest stem leaf only unpaired, adaxial ones usually opposite and equal or nearly so in a pair; bract 1; calyx without secretory canals; staminodes 2, 3, or absent; ovary 1-loculed; placentas 2, parietal.   (12)
       
12 (11) Leaves few, all alternate; inflorescences lax racemes appearing scorpioid; stamens 2 and abaxial, or 4; capsule loculicidal   54 Rhynchoglossum
+ Leaves 1 to few, lowest leaf unpaired, those near apex usually opposite; inflorescences scorpioid cymes, dense; stamens 2, adaxial; capsule circumscissile   55 Epithema
       
13 (10) Fertile stamens 4.   (14)
+ Fertile stamens 2.   (37)
       
14 (13) Calyx zygomorphic, 2-lipped, adaxial lip undivided, abaxial lip 4-lobed; disc tubular; capsule narrowly ellipsoid   15 Cathayanthe
+ Calyx actinomorphic, 5-lobed to 5-sect from base; disc ringlike, rarely cupular or inconspicuous; capsule oblong to linear, rarely conical to ovoid.   (15)
       
15 (14) Plants stemless.   (16)
+ Plants with a distinct stem.   (27)
       
16 (15) Adaxial corolla lip 4-lobed, over 2 X longer than abaxial lip, abaxial lip undivided   8 Paraisometrum
+ Adaxial corolla lip 2-lobed, equalling or much shorter than abaxial lip, abaxial lip 3-lobed or 3-sect.   (17)
       
17 (16) Stigma 1.   (18)
+ Stigmas 2.   (21)
       
18 (17) Abaxial corolla lip densely bearded inside; bracts apparently absent; corolla tube tubular; anthers coherent in pairs, thecae divaricate   13 Corallodiscus
+ Corolla glabrous to puberulent inside; bracts 2, rarely absent or several; corolla tube campanulate to broadly tubular, sometimes oblique, sometimes saccate at base, throat some- times constricted; anthers free or all 4 coherent together, thecae parallel or subparallel.   (19)
       
19 (18) Anthers of all 4 stamens coherent together; capsule oblique in relation to pedicel; adaxial corolla lip distinctly shorter than abaxial   16 Beccarinda
+ Anthers free; capsule straight in relation to pedicel; adaxial corolla lip slightly shorter than to as long as abaxial.   (20)
       
20 (19) Ovary oblong; anther thecae parallel, not confluent, seldom horseshoe-shaped, then thecae divaricate and confluent at apex; corolla tube 1-4 X longer than limb; leaf blade base cuneate to subcordate   5 Oreocharis
+ Ovary ovoid or conical; anther thecae subparallel, confluent at apex; corolla tube slightly shorter than limb; leaf blade base cuneate to attenuate   18 Boeica
       
21 (17) Adaxial corolla lip emarginate to nearly entire.   (22)
+ Adaxial corolla lip 2-lobed to 2-sect from base.   (23)
       
22 (21) Corolla tube subcampanulate to funnelform-tubular, not swollen; stamens adnate to corolla near base, all or only abaxial ones exserted, anthers free, thecae confluent at apex   6 Tremacron
+ Corolla tube narrowly funnelform to cylindric, slightly gibbous abaxially; stamens adnate to corolla tube above or rarely below middle, usually included, rarely exserted, anthers coherent in pairs, thecae not confluent   9 Ancylostemon
       
23 (21) Ovary 2-loculed, only adaxial locule fertile; placenta 1, axile; capsule oblique in relation to pedicel; cymes 1-3-flowered   11 Briggsiopsis
+ Ovary 1-loculed; placentas 2, parietal; capsule straight in relation to pedicel; cymes 1- to many flowered.   (24)
       
24 (23) Corolla tube narrowly campanulate, gibbous abaxially, (0.7-)1-2(-2.6) cm in diam., abaxial lip 3-lobed   10 Briggsia
+ Corolla tube cylindric to funnelform, broadly tubular, campanulate, urceolate, or salverform, not swollen, slightly gibbous abaxially, or slightly ampliate basally, 0.2-1(-1.2) cm in diam., abaxial lip 3-sect, rarely 3-lobed.   (25)
       
25 (24) Anthers dehiscing longitudinally from arcuate slits; corolla zygomorphic, rarely actinomorphic, tube campanulate, cylindric, or rarely urceolate or salverform, not swollen   7 Isometrum
+ Anthers dehiscing longitudinally, rarely transversely; corolla zygomorphic, tube campanulate-cylindric to broadly tubular, sometimes ampliate basally, throat sometimes constricted or narrowly funnelform to cylindric, slightly gibbous abaxially.   (26)
       
26 (25) Corolla purple to blue or white or yellow to red, tube campanulate-cylindric to broadly tubular, sometimes ampliate basally, throat sometimes constricted, adaxial lip 2-lobed; anthers free; calyx 5-sect from base   5 Oreocharis
+ Corolla orange to yellow or white, rarely pink, tube narrowly funnelform to cylindric, slightly gibbous abaxially, adaxial lip 2-lobed, emarginate, or rarely undivided; anthers coherent in pairs; calyx 5-sect from base to 5-lobed from above middle   9 Ancylostemon
       
27 (15) Stigmas 2.   (28)
+ Stigma 1.   (31)
       
28 (27) Ovary 2-loculed, only adaxial locule fertile; placenta 1, axile; capsule oblique in relation to pedicel; corolla tube tubular-funnelform; cymes 1-3-flowered   11 Briggsiopsis
+ Ovary 1-loculed; placentas 2, parietal; corolla tube cylindric to broadly tubular or narrowly campanulate, sometimes ampliate above; capsule straight in relation to pedicel; cymes 1- to many flowered.   (29)
       
29 (28) Corolla tube cylindric basally, ampliate above, not swollen   12 Didissandra
+ Corolla tube narrowly campanulate to broadly tubular, gibbous abaxially.   (30)
       
30 (29) Seeds unappendaged   10 Briggsia
+ Seeds with 1 hairlike appendage at each end   46 Loxostigma
       
31 (27) Calyx zygomorphic, 2-lipped, adaxial lip undivided, abaxial lip 4-lobed; disc tubular; capsule splitting along 1 suture   15 Cathayanthe
+ Calyx actinomorphic, 5-sect from base or 5-lobed, segments equal or unequal; disc ringlike, inconspicuous or absent; capsule 2(or 4)-valved.   (32)
       
32 (31) Leaves 1(or 2), apical on stem; leaf blade base cordate to auriculate; anthers dehiscing from fissures or cracks; capsule slightly longer than calyx   14 Platystemma
+ Leaves few to many, opposite, rarely whorled, spread along stem or crowded near stem apex; leaf blade base cuneate to attenuate or rounded, rarely cordate; anthers dehiscing longitudinally, poricidally, or transversely; capsule much longer than calyx.   (33)
       
33 (32) Seeds with 1 subulate or hairlike appendage at each end, or with up to 50 appendages at hilar end; corolla tube narrowly tubular to funnelform-tubular or subtubular, longer or much longer than limb, 4-17 mm in diam.; ovary linear; stamens adnate to corolla tube near or above middle, anthers usually coherent in pairs.   (34)
+ Seeds unappendaged; corolla tube campanulate, shorter than to slightly longer than limb, 3-9 mm in diam.; ovary ovoid, ellipsoid, or conical; stamens adnate to corolla near or slightly above base, anthers free or all 4 coherent together.   (35)
       
34 (33) Corolla white to yellowish, tube subtubular, gibbous abaxially near limb, with 2 arcuate projections abaxially inside; stamens included, thecae confluent at apex; seeds with 1 subulate appendage at each end   45 Anna
+ Corolla red to orange, seldom greenish, yellow, or white, tube narrowly tubular to funnelform-tubular, often curved, not swollen, inside sparsely puberulent, sparsely glandular puberulent, glabrous, or with a hair ring; stamens usually exserted, thecae not confluent; seeds with 1 hairlike appendage at each end or up to 50 appendages at hilar end   47 Aeschynanthus
       
35 (33) All anthers coherent together; capsule oblique in relation to pedicel; leaf blade base cordate to cuneate   16 Beccarinda
+ Anthers free; capsule straight in relation to pedicel; leaf blade base cuneate to attenuate.   (36)
       
36 (35) Corolla yellowish, tube slightly longer than limb; anther thecae not confluent; ovary ellipsoid; cymes 3-5-flowered   17 Leptobaea
+ Corolla white to pink or blue to purple, tube slightly shorter than limb; anther thecae confluent at apex; ovary ovoid or conical; cymes 1- to many flowered   18 Boeica
       
37 (13) Plants with a distinct stem.   (38)
+ Plants stemless.   (50)
       
38 (37) Stigmas 2.   (39)
+ Stigma 1.   (43)
       
39 (38) Flowers solitary; capsule valves spirally twisted; corolla tube campanulate-tubular, adaxial lip slightly shorter than abaxial   44 Rhabdothamnopsis
+ Inflorescences cymes, sometime spikelike or racemelike, 2-10-flowered; capsule valves straight, not twisted; corolla tube slender funnelform, funnelform, or funnelform-tubular, adaxial lip much shorter than abaxial.   (40)
       
40 (39) Bracts forming a globose involucre; corolla tube much longer than limb; leaf blade base attenuate to broadly cuneate; stamens adnate to corolla tube below middle, anthers coherent by adaxial surfaces, thecae divaricate, confluent   26 Hemiboeopsis
+ Bracts not forming an involucre; corolla tube longer than limb; leaf blade base cuneate to rounded or cordate; stamens adnate to corolla tube near middle, anthers coherent at apex, thecae parallel, not confluent at apex.   (41)
       
41 (40) Stigmas equal, oblate or semiorbicular, undivided; calyx 5-sect from base; stamens adnate to corolla tube near mouth, slightly exserted   21 Didymostigma
+ Stigmas unequal, smaller 1 truncate or triangular, undivided, larger 1 narrowly obtrapeziform, emarginate; calyx 5-lobed; stamens adnate to corolla tube near middle, included.   (42)
       
42 (41) Ovary 1-loculed; placentas parietal; corolla abaxial lip lobes unequal   22 Pseudochirita
+ Ovary 2-loculed; placentas axile; corolla abaxial lip lobes subequal   23 Allostigma
       
43 (38) Subshrubs or rarely lianas, usually epiphytic, less frequently epipetric or terrestrial; seeds with 1 appendage at each end   48 Lysionotus
+ Herbs, rarely subshrubs (rarely shrubs outside China), always terrestrial or epipetric; seeds unappendaged.   (44)
       
44 (43) Anthers basifixed, thecae parallel, not confluent at apex; bracts 2, forming a globose to ovoid involucre.   (45)
+ Anthers dorsifixed, thecae divaricate, confluent at apex; bracts 2, not forming an involucre, rarely up to 7 and whorled, 1, or absent.   (46)
       
45 (44) Ovary 1-loculed; placenta parietal; capsule dehiscing on both sides; corolla inside glabrous; tube somewhat gibbous abaxially, adaxial lip inconspicuous, much shorter than abaxial lip   24 Metabriggsia
+ Ovary 2-loculed, only adaxial locule fertile; placenta axile; capsule dehiscing only adaxially; corolla inside usually with a ring of hairs; tube not swollen, adaxial lip shorter than to nearly as long as abaxial   25 Hemiboea
       
46 (44) Stamens adnate to abaxial side of corolla tube near or above middle; capsule valves straight, not twisted.   (47)
+ Stamens adnate to abaxial side of corolla tube near base; capsule valves usually at least slightly twisted.   (48)
       
47 (46) Stigma abaxial, 2-lobed to undivided   32 Chirita
+ Stigma terminal, undivided   35 Didymocarpus
       
48 (46) Anthers dehiscing transversely; disc cupular; stigma ligulate; calyx zygomorphic   43 Trisepalum
+ Anthers dehiscing longitudinally from straight or arcuate slits; disc inconspicuous or ringlike; stigma capitate, seldom subligulate; calyx actinomorphic, rarely slightly zygomorphic.   (49)
       
49 (48) Leaf blade adaxially woolly to glabrous, abaxially woolly, pannose, or velutinous, hairs interwoven, usually branched; corolla inside glabrous to glandular puberulent; tube longer than to occasionally nearly equalling limb   40 Paraboea
+ Leaf blade puberulent to woolly, hairs unbranched; corolla inside bearded, tube shorter than limb   42 Ornithoboea
       
50 (37) Corolla adaxial lip 4-lobed, abaxial lip undivided.   (51)
+ Corolla adaxial lip 2-lobed, rarely undivided, abaxial lip 3-lobed, rarely 4-lobed.   (52)
       
51 (50) Inflorescences dense umbel-like cymes; corolla tube 4-5 X longer than limb, adaxial lip longer than abaxial; stamens adnate to corolla tube above middle   38 Calcareoboea
+ Inflorescences lax cymes; corolla tube shorter than limb, adaxial lip slightly shorter than abaxial; stamens adnate to corolla tube near base   39 Allocheilos
       
52 (50) Anthers dehiscing transversely; stigma 1, ligulate; capsule valves spirally twisted   43 Trisepalum
+ Anthers dehiscing longitudinally; stigmas 1 or 2, if 1 then usually capitate, globose, disclike, truncate, lamelliform, oblate, or obtrapeziform, rarely linear or subligulate; capsule valves straight or twisted.   (53)
       
53 (52) Corolla adaxial lip undivided.   (54)
+ Corolla adaxial lip 2-lobed.   (55)
       
54 (53) Corolla tube tubular to funnelform-tubular, longer than limb; stamens exserted, anthers basifixed, free or coherent at apex, thecae parallel, not confluent; leaf blade base broadly cuneate to subrounded   29 Deinocheilos
+ Corolla tube broadly tubular, nearly equalling limb; stamens included, anthers dorsifixed, coherent by adaxial surfaces, thecae divaricate, confluent at apex; leaf blade base cordate   36 Gyrocheilos
       
55 (53) Anthers basifixed or if dorsifixed then stamens slightly exserted, thecae parallel.   (56)
+ Anthers dorsifixed or nearly so, stamens included, thecae divaricate.   (58)
       
56 (55) Anthers dorsifixed, stamens slightly exserted; placentas undivided; inflorescences monochasial cymes, 1- or 2-flowered   20 Dayaoshania
+ Anthers basifixed, stamens included; placentas projecting inward, 2-cleft; inflorescences cymes, sometimes umbel-like, 1- to many flowered.   (57)
       
57 (56) Corolla tube 1.5-4 X longer than limb, 3.5-17 mm in diam.; stamens adnate to adaxial side of corolla tube near middle, rarely near base; ovary and capsule linear   19 Opithandra
+ Corolla tube shorter than limb, 2-5 mm in diam.; stamens adnate to abaxial side of corolla tube near base; ovary conical to broadly ovoid; capsule narrowly ellipsoid to oblong to ovoid   27 Petrocosmea
       
58 (55) Anthers free, thecae not confluent; capsule nearly globose; leaf blade base rounded, peltate   28 Metapetrocosmea
+ Anthers coherent, thecae confluent at apex; capsule linear to narrowly ovoid, ellipsoid, or oblong; leaf blade base attenuate to cordate, rarely auriculate or auriculate-peltate.   (59)
       
59 (58) Stigmas 2; corolla lobe apex acute   30 Lagarosolen
+ Stigma 1; corolla lobe apex rounded, seldom obtuse or acute.   (60)
       
60 (59) Capsule valves spirally twisted or seldom straight, if straight leaf blade abaxially woolly, pannose, or velutinous, hairs dense, interwoven, usually branched.   (61)
+ Capsule valves straight, not twisted; leaf blade abaxially strigose or puberulent to villous, velutinous, or woolly.   (62)
       
61 (60) Capsule valves 4; leaf blade abaxially woolly, pannose, or velutinous, hairs very dense, interwoven, usually branched   40 Paraboea
+ Capsule valves 2; leaf blade villous to puberulent, hairs dense, not interwoven or branched   41 Boea
       
62 (60) Corolla tube slightly shorter than to nearly equalling limb, lobes acute   37 Dolicholoma
+ Corolla tube longer to much longer than limb, lobes rounded, obtuse, or acute.   (63)
       
63 (62) Ovary ovoid or narrowly ovoid; capsule narrowly ovoid to ellipsoid, equalling to shorter than calyx.   (64)
+ Ovary linear; capsule linear to very narrowly ovoid, much surpassing calyx.   (65)
       
64 (63) Corolla nearly salverform, tube cylindric, adaxial lip slightly shorter than abaxial   31 Primulina
+ Corolla tube nearly tubular, adaxial lip slightly to 2-3 X shorter than abaxial   33 Chiritopsis
       
65 (63) Corolla tube suburceolate-tubular, 1-2.5 mm in diam., limb indistinctly 2-lipped, adaxial lip slightly shorter than abaxial; lobe apex acute; stamens adnate to abaxial side of corolla tube slightly above base   34 Petrocodon
+ Corolla tube funnelform-tubular to campanulate or cylindric, 2-20 mm in diam., limb 2-lipped; adaxial lip slightly to 1.5-3.5 X shorter than abaxial; lobe apex rounded, rarely obtuse to acute; stamens adnate to abaxial side of corolla tube near or above middle.   (66)
       
66 (65) Stigma abaxial, 2-lobed to undivided   32 Chirita
+ Stigma terminal, undivided   35 Didymocarpus

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