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35. Didymocarpus Wallich, Edinburgh Philos. J. 1: 378. 1819.
[nom. cons.]

长蒴苣苔属 chang shuo ju tai shu

Roettlera Vahl (1804), not Willdenow (1797).

Herbs, rarely shrubs or subshrubs (outside China), perennial, terrestrial or epipetric, rhizomatous, stemless or simple stemmed. Leaves few to many, along stem, then opposite, whorled, or alternate or basal, equal to subequal in a pair; leaf blade sparsely to densely puberulent to pubescent, rarely nearly glabrous, short strigose, or woolly, base cuneate to cordate, rarely auriculate. Inflorescences axillary, sometimes umbel-like, lax or sometimes dense, 1- to many-flowered cymes; bracts 2, opposite. Calyx actinomorphic and 5-fid, or zygomorphic and 2-lipped or 5- or 3-sect from base to shallowly lobed; segments equal to unequal. Corolla blue to wine-red, rarely pink, white, or yellow, zygomorphic, inside glabrous, rarely sparsely puberulent; tube cylindric to funnelform-tubular, not swollen, rarely saccate at base, longer than limb, 2-13 mm in diam.; limb 2-lipped; adaxial lip 2-lobed, 1.5-3 X shorter than abaxial lip; abaxial lip 3-lobed, lobes equal or central lobe longer, apex rounded, rarely obtuse to acute. Stamens 2, adnate to abaxial side of corolla tube near or above middle, included; anthers dorsifixed, coherent by adaxial surfaces, thecae divaricate, confluent at apex, dehiscing longitudinally; connective not projecting; staminodes (absent or)2 or 3, adnate to adaxial or adaxial and abaxial sides of corolla tube. Disc ringlike, rarely cupular. Ovary linear, 1-loculed; placentas 2, parietal, slightly projecting inward, 2-cleft. Stigma 1, terminal, depressed-globose to disclike or truncate, undivided. Capsule straight in relation to pedicel, usually linear, much surpassing calyx, dehiscing loculicidally to base; valves 2 or 4, straight, not twisted. Seeds unappendaged.

About 180 species: SE Asia; 31 species in China.

The delimitation of Chirita and Didymocarpus is somewhat problematic. As defined here, the genera are based entirely on stigma shape. Thus, more than a few Chinese taxa described in Didymocarpus (see especially Wang et al., Edinburgh J. Bot. 49: 54. 1991) and some included here (e.g., D. cortusifolius and D. heucherifolius) were included in Chirita by other recent workers (see D. Wood, Notes Roy. Bot. Gard. Edinburgh 33: 123-205. 1974). Didymocarpus nigrescens H. Léveillé & Vaniot and D. violaceus H. Léveillé (1912, not Ridley 1896) cannot be placed reliably in any genus because they are only known from fruiting specimens. The senior author suggests that both taxa probably belong in Didymocarpus, but Burtt suggested that D. nigrescens belongs in Petrocodon (Notes Roy. Bot. Gard. Edinburgh 38: 471. 1980) or Tengia (Notes Roy. Bot. Gard. Edinburgh 22: 314. 1958).

1 Herbs stemless.   (2)
+ Herbs with stems (0.7-)3-62 cm.   (11)
2 (1) Leaf blade elliptic to oblong, ovate, or oblanceolate, not lobed, base cuneate to rounded, pinnately veined, herbaceous or papery; calyx 5-sect from base.   (3)
+ Leaf blade ovate to orbicular, triangular, oblate, or reniform, often lobed, base cordate to truncate, palmately veined (then leaf blade papery) or rarely pinnately veined (then leaf blade leathery); calyx shallowly 5-lobed (rarely 5-sect from or near base).   (5)
3 (2) Leaf blade shorter than 5 cm, densely white woolly; calyx segments 2-2.5 mm   24 Didymocarpus niveolanosus
+ Leaf blade 2.2-20 cm (mostly over 5 cm), densely appressed puberulent or short strigose; calyx segments 2.5-7 mm.   (4)
4 (3) Leaf blade to 20 X 10.5 cm, densely appressed puberulent; filaments sparsely puberulent and glandular; pistil puberulent; corolla yellow   22 Didymocarpus mollifolius
+ Leaf blade to 10 X 3.6 cm, sparsely short strigose; filaments glabrous; pistil sparsely glandular; corolla pale purple   23 Didymocarpus hancei
5 (2) Calyx divided from or near base; tube to 2 mm.   (6)
+ Calyx divided from or near middle, tube at least 4 mm.   (8)
6 (5) Leaf blade not lobed, margin obtusely dentate or double dentate, apex obtuse; corolla yellow, tube funnelform-cylindric, mouth 3-4 mm in diam.   25 Didymocarpus subpalmatinervis
+ Leaf blade distinctly or indistinctly multilobed, margin denticulate to serrate or entire, apex rounded; corolla pink, tube funnelform-tubular to campanulate, mouth 1-1.3 cm in diam.   (7)
7 (6) Calyx lobes broadly lanceolate, oblanceolate-linear, or triangular, 3-4 mm, margin sparsely denticulate; corolla 2.5-3.2 cm, puberulent outside; anthers puberulent; staminodes 3; leaf blade 3-9 X 3.5-11 cm, eglandular   26 Didymocarpus heucherifolius
+ Calyx lobes spatulate-linear or narrowly obovate, 6-8 mm, margin entire; corolla 1.5-1.8 cm, glabrous outside; anthers white woolly; staminodes 2; leaf blade 1.8-4.6 X 1.6-4 cm, sparsely yellow glandular   27 Didymocarpus sinoprimulinus
8 (5) Leaf blade leathery, pinnately veined; calyx slightly zygomorphic, ca. 13 mm, lobes unequal, abaxial lobe ca. 7 mm, others 2.5-3.5 mm; anthers glabrous; staminodes 2   28 Didymocarpus yuenlingensis
+ Leaf blade papery, palmately veined; calyx actinomorphic, 7-9 mm, lobes equal, 1-3 mm; anthers white woolly or puberulent; staminodes 3.   (9)
9 (8) Bracts semiorbicular, adaxially pubescent, abaxially villous; calyx lobes depressed oblong, overlapping; filaments ca. 15 mm; anthers ca. 5 mm; staminodes 2-3 mm; style glabrous   31 Didymocarpus salviiflorus
+ Bracts ovate to elliptic or narrowly oblong, abaxially puberulent to sparsely so; calyx lobes ovate-triangular to triangular, not overlapping; filaments 6-10 mm; anthers 2.5-3 mm; staminodes 0.5-0.8 mm; style puberulent or sparsely so.   (10)
10 (9) Leaf blade orbicular-ovate to orbicular or triangular, 4.6-10 X 3.2-9 cm; bract margin entire; pedicel 4-10 mm; calyx lobe margin denticulate; corolla white, puberulent outside, adaxial lip 5-6 mm, abaxial lip 7-9 mm; anthers white woolly   29 Didymocarpus cortusifolius
+ Leaf blade reniform or orbicular-reniform, 2-4.5 X 3.5-7.5 cm; bract margin few crenate; pedicel 11-18 mm; calyx lobe margin entire; corolla pink, glabrous outside, adaxial lip ca. 1 cm, abaxial lip ca. 16 mm; anthers sparsely puberulent   30 Didymocarpus reniformis
11 (1) Calyx divided from base into 3 or 5 segments, without a tube.   (12)
+ Calyx not divided from base, 2-lipped or 5-lobed, tube 1.5-9 mm.   (17)
12 (11) Calyx actinomorphic or slightly zygomorphic, 5-sect, segments of same width.   (13)
+ Calyx zygomorphic, 3-sect, adaxial lip 2-3 X wider than abaxial 2 segments.   (15)
13 (12) Stems to 5.5 cm; petiole 3-12 cm; leaf blade broadly ovate to orbicular, base cordate to truncate, palmately veined; peduncle 6-15 cm; calyx slightly zygomorphic, segments unequal in length   3 Didymocarpus margaritae
+ Stems 10-30 cm; petiole 0.2-5 cm; leaf blade elliptic to ovate or oblong, base rounded to cuneate, pinnately veined; peduncle 2.3-7 cm; calyx actinomorphic, segments equal in length.   (14)
14 (13) Leaf blade 2.5-4.8 cm; lateral veins 6 or 7 on each side of midrib; peduncle glabrous; corolla 2.5-3 cm; filaments ca. 7 mm; pistil ca. 2 cm   1 Didymocarpus medogensis
+ Leaf blade 4-18.5 cm; lateral veins 8-11 on each side of midrib; peduncle sparsely glandular puberulent or pilose; corolla 1.6-2.2 cm; filaments ca. 3.5 mm; pistil ca. 1.1 cm   2 Didymocarpus glandulosus
15 (12) Calyx 2-3 mm, adaxial segment 3-lobed from near middle, outer surface glabrous   4 Didymocarpus zhenkangensis
+ Calyx 4-5 mm, adaxial segment shallowly 3-lobed, outer surface puberulent or sparsely glandular puberulent.   (16)
16 (15) Leaf blade to 9 cm wide; cymes 3-15-flowered; peduncle 4-8 cm; adaxial calyx lip ca. 3 mm wide, lobes triangular, 2 abaxial calyx segments oblong, ca. 1.5 mm wide; corolla adaxial lip ca. 3 mm, abaxial lip ca. 7 mm; anthers puberulent; staminodes 2   5 Didymocarpus pseudomengtze
+ Leaf blade to 4.3 cm wide; cymes 1-3-flowered; peduncle 1.8-3.5 cm; adaxial calyx lip ca. 1.5 mm wide, lobes lanceolate-linear, 2 abaxial calyx segments narrowly lanceolate, 0.8-0.9 mm wide; corolla adaxial lip ca. 1.8 mm, abaxial lip ca. 4 mm; anthers glabrous; staminodes absent   6 Didymocarpus stenocarpus
17 (11) Calyx actinomorphic, 2-3 mm, 5-lobed nearly from middle; corolla mouth ca. 2.5 mm in diam.; staminodes absent; ovary appressed puberulent   8 Didymocarpus silvarum
+ Calyx slightly to strongly zygomorphic, 3-12 mm, indistinctly to distinctly 2-lipped; corolla mouth 3-10 mm in diam.; staminodes 1-3 (unknown in D. nanophyton); ovary glabrous, rarely densely glandular puberulent, minutely glandular, or sparsely puberulent.   (18)
18 (17) Calyx tuberculate outside; stems 2.5-7 cm; leaf blade 2.5-4.5 X 1.5-3 cm   15 Didymocarpus nanophyton
+ Calyx smooth outside; stems 3-30(-62) cm; leaf blade 1-15(-26) X 1-10(-16) cm.   (19)
19 (18) Corolla 2.5-4.5 cm.   (20)
+ Corolla to 2.4 cm.   (24)
20 (19) Calyx 10-12 mm; pistil sparsely puberulent or minutely glandular.   (21)
+ Calyx 3-6 mm; pistil glabrous, eglandular.   (22)
21 (20) Petiole to 2.5(-5) cm; leaf blade to 5.5 cm wide, adaxially densely appressed puberulent; peduncle glandular puberulent; corolla ca. 3 cm, mouth ca. 3.5 mm in diam.; anthers glabrous; ovary minutely glandular   20 Didymocarpus praeteritus
+ Petiole to 11 cm; leaf blade to 10.5(-16) cm wide, adaxially sparsely appressed puberulent to glabrous; peduncle glabrous; corolla 3.2-4.5 cm, mouth 9-10 mm in diam.; anthers puberulent; ovary sparsely puberulent   21 Didymocarpus purpureobracteatus
22 (20) Corolla sparsely pubescent outside, mouth 4-6 mm in diam.; adaxial leaf surface eglandular; bracts 2-20 mm, often leaflike, margin crenate to entire   19 Didymocarpus yunnanensis
+ Corolla glabrous outside, mouth 7-8 mm in diam.; adaxial leaf surface with yellow or orange glands (at least sparsely); bracts 3-6 mm, never leaflike, margin entire.   (23)
23 (22) Stem leaves in 2 whorls; petiole to 7 cm; leaf blade adaxially sparsely yellow glandular, base oblique, cuneate to cordate; peduncle to 4 cm; bracts free; calyx 3-4.5 mm; staminodes 3, 1.8-3.6 mm   12 Didymocarpus pulcher
+ Stem leaves opposite; petiole to 2.5 cm; leaf blade adaxially orange glandular, base oblique, broadly cuneate; peduncle 5.5-9 cm; bracts connate at base; calyx 5-6 mm; staminodes 2, ca. 6 mm   17 Didymocarpus leiboensis
24 (19) Leaves crowded near stem apex, nodes indistinct; leaves sparsely yellow glandular on both surfaces; staminode 1.   (25)
+ Leaves spread along stem or sometimes apical nodes dense, opposite, rarely whorled or basal few alternate, nodes 1-5; leaves eglandular, sometimes sparsely yellow, red, or purple glandular; staminodes 2 or 3.   (26)
25 (24) Stems spreading villous; lateral veins 3 or 4 on each side of midrib; bracts ca. 2 mm; calyx tube 2.2-2.5 mm   11 Didymocarpus villosus
+ Stems sparsely appressed puberulent; lateral veins 6-8 on each side of midrib; bracts 4-7 mm; calyx tube 3.5-4 mm   13 Didymocarpus primulifolius
26 (24) Abaxial leaf surface sparsely red glandular; petiole 0.5-10 cm; leaf blade ovate to reniform; pedicel ca. 2 mm   14 Didymocarpus grandidentatus
+ Abaxial leaf surface yellow glandular or eglandular; petiole 0.1-3.5(-4.7) cm; leaf blade elliptic to broadly ovate or obovate, rarely triangular, oblong, or rhombic; pedicel usually over 2 mm.   (27)
27 (26) Adaxial leaf surface sparsely purple glandular; bracts connate basally; anthers glabrous; ovary ca. 6 mm, densely glandular puberulent   18 Didymocarpus adenocalyx
+ Adaxial leaf surface eglandular or yellow glandular; bracts free or sometimes connate basally (in some D. stenanthos); anthers glabrous, rarely puberulent; ovary 10-15 mm, glabrous.   (28)
28 (27) Corolla puberulent outside, tube funnelform, mouth ca. 6 mm in diam.; filaments ca. 6 mm; leaf blade base cordate-auriculate   9 Didymocarpus zhufengensis
+ Corolla glabrous outside, tube cylindric or nearly tubular, mouth 3-4 mm in diam.; filaments 3-4 mm; leaf blade base cuneate to cordate.   (29)
29 (28) Leaves mostly opposite, basal 2 often alternate, leaf blade 2-6.8(-11) X 1.3-4.2(-5.6) cm, sometimes yellow glandular; cymes 2-5-flowered; calyx slightly zygomorphic, 5-lobed, lobes nearly equal, sparsely glandular puberulent outside; staminodes 2.5-4 mm   10 Didymocarpus aromaticus
+ Leaves opposite, leaf blade 2-11.5(-15) X 1.8-6.8 cm, adaxial surface eglandular, abaxial surface eglandular or sometimes very sparsely yellow glandular; cymes 5- to many flowered; calyx zygomorphic, 2-lipped from near middle, adaxial lip 3 lobed, abaxial lip 2-lobed, glabrous outside; staminodes 0.3-1.2 mm.   (30)
30 (29) Leaf blade abaxial surface sparsely pubescent and very sparsely yellow glandular; bracts free; anthers glabrous   7 Didymocarpus mengtze
+ Leaf blade abaxial surface puberulent along veins, eglandular; bracts connate at base or free; anthers nearly glabrous   16 Didymocarpus stenanthos

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