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56. Mussaenda Linnaeus, Sp. Pl. 1: 177. 1753.
玉叶金花属 yu ye jin hua shu
Authors: Tao Chen & Charlotte M. Taylor
Belilla Adanson.
Trees, shrubs, or clambering or twining lianas, rarely dioecious, unarmed. Raphides absent. Leaves opposite or occasionally in whorls of 3, with or usually without domatia; stipules persistent or caducous, interpetiolar, entire or 2-lobed. Inflorescences terminal and sometimes also in axils of uppermost leaves, cymose, paniculate, or thyrsiform, several to many flowered, sessile to pedunculate, bracteate. Flowers sessile to pedicellate, bisexual and usually distylous or rarely unisexual. Calyx limb 5-lobed nearly to base, frequently some or all flowers of an inflorescence with 1(-5) white to colored, petaloid, persistent or deciduous, membranous, stipitate calycophyll(s) with 3-7 longitudinal veins. Corolla yellow, red, orange, white, or rarely blue (Mussaenda multinervis), salverform with tube usually slender then abruptly inflated around anthers, or rarely constricted at throat (M. hirsuta), inside variously pubescent but usually densely yellow clavate villous in throat; lobes 5, valvate-reduplicate in bud, often long acuminate. Stamens 5, inserted in middle to upper part of corolla tube, included; filaments short or reduced; anthers basifixed. Ovary 2-celled, ovules numerous in each cell, inserted on oblong, fleshy, peltate, axile placentas; stigmas 2-lobed, lobes linear, included or exserted. Fruit purple to black, baccate or perhaps rarely capsular (M. decipiens), fleshy, globose to ellipsoid, often conspicuously lenticellate, with calyx limb persistent or caducous often leaving a conspicuous scar; seeds numerous, small, angled to flattened; testa foveolate-striate; endosperm abundant, fleshy.
About 200 species: widespread in tropical Africa, Asia, Madagascar, and Pacific islands; 29 species (18 endemic, one introduced) in China.
The characteristic large, petaloid calycophylls of Mussaenda are frequently but mistakenly considered to be bracts. Mussaenda is frequently confused with Schizomussaenda; Schizomussaenda can be recognized by its large shrub habit with relatively large leaves and calycophylls, its capsular fruit, and its flower buds with the corollas clavate and rather flat-topped with the acuminate ends of the corolla lobes then bent upward to form an erect appressed group of filaments on the top of the bud. Several species of Mussaenda are frequently cultivated as ornamentals in tropical regions. Mussaenda philippica A. Richard is apparently the most commonly cultivated species and has numerous cultivar forms; Puff et al. (Rubiaceae of Thailand, 215. 2005) presented photos of several of the cultivated forms. This species is pilosulous to villous throughout, with white to pink flowers and calycophylls and with calyx lobes variously 1 to all modified into calycophylls. Also frequently cultivated are M. frondosa of our flora and M. erythrophylla Schumacher & Thonning of Africa; the latter has pilose to villous pubescence on all organs, including the corollas, and deep red inflorescence axes and flowers, including the calycophylls and the outside of the corolla but excepting the corolla limb, which is creamy white. In addition to the species treated here, Hooker and Arnott (Bot. Beechey Voy. 265. 1838) reported Mussaenda glabra Vahl from "Loo Choo" in China. Mussaenda glabra was described from the Himalaya and has medium-sized leaves, a climbing habit, petioles 3-15 mm, a corymbiform branched inflorescence, calyx lobes 1-4 mm and deciduous in fruit, corolla tubes 13-18 mm, and corolla lobes 3-3.5 mm; this species is otherwise known from 300-1300 m in India and Bhutan and has not been confidently recorded from China. The name "M. glabra" has been frequently applied in herb. to Chinese specimens of both M. erosa and M. frondosa. Also Henry 8279 from Hainan was treated by C. E. C. Fisher (Bull. Misc. Inform. Kew 1928: 274. 1928) as M. parryorum C. E. C. Fischer, a species otherwise known only from Assam, NE India. This identification seems unlikely, especially given that M. parryorum has not otherwise been reported or treated from China or Indochina, so the Henry specimen will need re-examination. Mussaenda parryorum is included for reference in the key below, based on its protologue, but the species is not otherwise treated here. H. H. Hsue and H. Wu (in FRPS 71(1): 283-306. 1999), Hutchinson (in Sargent, Pl. Wilson. 3: 395-400. 1916), and some other authors have distinguished Mussaenda species in large part based on leaf shape, ovary length, calyx lobe length, and corolla size, but the consistency and distinctiveness of these characters within species of Mussaenda have been questioned by some other authors. In particular, corolla size is apparently sometimes widely variable among living plants in a population (see comments by Wood, Fl. Bhutan 2(2): 781. 1999), and the flower buds of Mussaenda apparently often open prematurely when collected and thus are shorter than flowers at anthesis (pers. obs.). The calyx lobes and the ovary often elongate after fertilization of the ovary, and the calyx lobes then fall off; thus, fruiting plants are very difficult to identify. Also Hutchinson misidentified several widely distributed sets of Chinese Mussaenda specimens, generating confusion in this group (e.g., he identified Henry 12157 as M. pubescens in spite of its pedicellate flowers with corolla tubes ca. 32 mm, vs. sessile and 11-20 mm in M. pubescens); and he described only the "typical" leaf size and shape of several species although he saw material with more variability. The weakness in some cases of Hutchinson’s characters and species concepts is shown by his simultaneous description in his Mussaenda treatment of two new species that were separated primarily on vegetative features and actually are both based on different specimens of a single distinctive species, Schizomussaenda dehiscens. As noted below, Mussaenda multinervis and M. decipiens were both described as having capsular fruit and most likely belong in other genera, but which other genus or genera is not yet clear so they are maintained here. The measurements below of inflorescence size do not include the corollas and calycophylls.
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| 1 |
Individual flowers with all 5 calyx lobes enlarged into petaloid calycophylls |
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(2) |
| + |
Individual flowers with only 1 or 2 calyx lobes enlarged into a petaloid calycophyll, or without calycophylls |
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(3) |
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| 2 (1) |
Petiole 0.3-0.8 cm, leaf blade 2-9 × 1-4 cm; calyx lobes linear. |
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25 M. pubescens |
| + |
Petiole 1.5-3.5 cm, leaf blade 6-20 × 3.5-13 cm; calyx lobes lanceolate to ovate, 2-2.5 mm wide. |
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27 M. shikokiana |
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| 3 (1) |
Calyx lobes (i.e., not calycophylls) subleaflike, 1.5-5 mm wide, lanceolate, oblanceolate, oblong-lanceolate, ligulate, or broadly triangular |
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(4) |
| + |
Calyx lobes less than 1.5 mm wide, linear to narrowly triangular, not at all leaflike |
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(7) |
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| 4 (3) |
Corolla tube 20-25 mm, sericeous with appressed indument, sericeous with apices of trichomes spreading, or villous |
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(5) |
| + |
Corolla tube 9-14 mm, sericeous |
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(6) |
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| 5 (4) |
Stems sericeous with appressed indument; stipules 5-8 × 4-5 mm; calyx lobes 4-11 mm. |
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19 M. macrophylla |
| + |
Stems villous or hirsute with widely spreading trichomes; stipules 7-10 × 6-12 mm; calyx lobes 7-18 mm. |
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29 M. treutleri |
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| 6 (4) |
Stipules triangular-ovate to broadly ovate, acute to cuspidate; corolla tube 10-11 mm; berries ellipsoid. |
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9 M. emeiensis |
| + |
Stipules triangular to ovate-lanceolate, acute; corolla tube 9-14 mm; berries ellipsoid to subglobose. |
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27 M. shikokiana |
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| 7 (3) |
Stipules 13-20 mm, lobed for less than half their length; corolla yellow or pale blue |
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(8) |
| + |
Stipules 2-8 mm, deeply 2-parted; corolla white, yellow, or orange |
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(10) |
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| 8 (7) |
Corolla pale blue; leaves 8.5-12 cm wide; calyx lobes ca. 2 mm; fruit baccate. |
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22 M. multinervis |
| + |
Corolla yellow; leaves 2.5-7.5 cm wide; calyx lobes 1-4 mm; fruit capsular |
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(9) |
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| 9 (8) |
Calyx lobes 3-4 mm; leaves 6-7.5 cm wide. |
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5 M. decipiens |
| + |
Calyx lobes 1-2 mm; leaves 2.5-6 cm wide. |
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Schizomussaenda dehiscens |
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| 10 (7) |
Leaves sessile, subsessile, or shortly petiolate with petioles to 3 mm, obtuse, rounded, truncate, or cordulate and then sometimes decurrent at base, if petioles more than 2 mm then leaves truncate or cordulate |
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(11) |
| + |
Leaves distinctly petiolate with petioles 2-15 mm, truncate, obtuse, acute, or attenuate at base, if petioles 3 mm or shorter then leaves acute |
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(14) |
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| 11 (10) |
Leaf blade obtuse to rounded then abruptly acuminate at apex; corolla tube outside densely villous with pubescence drying reddened; stems densely villous. |
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26 M. sessilifolia |
| + |
Leaf blade tapered to acuminate apex; corolla tube outside with sparse to dense villosulous, pilosulous, tomentulose, strigose, or strigillose pubescence drying clear, whitened, or reddened; stems strigillose, villous, villosulous, or tomentulose |
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(12) |
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| 12 (11) |
Corolla tube 20-26 mm; inflorescences shortly branched, flowers with pedicels ca. 1 mm. |
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14 M. hossei |
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Corolla tube 11-20 mm; inflorescences subcapitate to congested, flowers mostly or all sessile to subsessile |
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(13) |
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| 13 (12) |
Calyx lobes ca. 2 mm. |
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2 M. breviloba |
| + |
Calyx lobes 3-6 mm. |
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25 M. pubescens |
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| 14 (10) |
Leaves glabrous throughout or sparsely strigose on principal veins, drying thickly papery and usually brown, with tertiary venation sparsely visible abaxially, widely spaced and subparallel, not areolate; stipules caducous exposing a persistent, reddish brown fringe of trichomes. |
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10 M. erosa |
| + |
Leaves glabrous or pubescent at least sparsely on veins abaxially, drying thinly to thickly papery or membranous and green to brown, with tertiary venation regularly visible abaxially, subparallel or areolate; stipules persistent or deciduous, without persistent fringe of trichomes or this present and clear or whitened |
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(15) |
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| 15 (14) |
Corolla tube constricted at throat. |
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13 M. hirsutula |
| + |
Corolla tube uniformly cylindrical or inflated just below or at throat |
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(16) |
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| 16 (15) |
Calyx lobes 25-30 mm, as long as or longer than corolla tube. |
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15 M. kwangsiensis |
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Calyx lobes 1-15 mm, shorter than corolla tube |
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(17) |
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| 17 (16) |
Corolla tube 5-8 mm; fruit stipitate or pedicellate, pedicels to 12 mm. |
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23 M. parviflora |
| + |
Corolla tube 5-40 mm; fruit sessile or subsessile to pedicellate, pedicels to 8 mm |
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(18) |
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| 18 (17) |
Flowers sessile in 1-5 capitate groups; bracts and calyx lobes linear, 8-15 mm, and so densely hirsute as to obscure inflorescence morphology. |
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6 M. densiflora |
| + |
Flowers sessile to pedicellate in cymes with axes at least shortly developed; bracts and calyx lobes narrowly triangular to lanceolate, 1-15 mm, glabrous to variously pubescent, inflorescence morphology obscured or evident |
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(19) |
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| 19 (18) |
Longest calyx lobes 6-15 mm on flowers at anthesis, longer than hypanthium portion |
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(20) |
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Longest calyx lobes 1-5.9 mm, shorter than to longer than hypanthium portion |
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(30) |
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| 20 (19) |
Corolla lobes 6-8 mm |
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(21) |
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Corolla lobes 2.5-6 mm, at least some shorter than 6 mm |
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(23) |
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| 21 (20) |
Cultivated plants; leaves strigose, strigillose, or glabrescent abaxially. |
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11 M. frondosa |
| + |
Native plants; leaves hirsute, subappressed villous, pilose, or strigose abaxially |
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(22) |
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| 22 (21) |
Calyx densely villous, with pubescence spreading. |
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3 M. caudatiloba |
| + |
Calyx hirsute. |
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6 M. densiflora |
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| 23 (20) |
Corolla tubes 26-30 mm |
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(24) |
| + |
Corolla tubes 11-25 mm |
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(25) |
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| 24 (23) |
Inflorescences densely congested; flowers sessile or subsessile. |
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13 M. hirsutula |
| + |
Inflorescences congested to laxly cymose; flowers with pedicels 1-5 mm. |
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17 M. laxiflora |
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| 25 (23) |
Fruit ellipsoid to ellipsoid-oblong, 18-20 × 11-12 mm, densely lenticellate, somewhat woody; stems and leaves abaxially densely villosulous to tomentulose; Hainan. |
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12 M. hainanensis |
| + |
Fruit subglobose to ellipsoid, 5-10 × 4-10 mm, smooth, fleshy or stiffly papery; stems and leaves abaxially glabrescent to variously pubescent; widespread |
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(26) |
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| 26 (25) |
Corolla with tube ca. 14 mm, lobes ca. 2.5 mm; Yunnan, elevation not noted. |
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24 M. pingbianensis |
| + |
Corolla with tube 11-23 mm, lobes 2.5-5 mm; widespread, below 100-1600 m |
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(27) |
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| 27 (26) |
Leaves 2-11.5 × 1-4.5 cm; flowers sessile to pedicellate, white to yellow; below 100-900 m, throughout China. |
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25 M. pubescens |
| + |
Leaves 6-20 × 3-10 cm; flowers sessile, mixed sessile and pedicellate, or all pedicellate with pedicels to 3 mm, yellow-orange; S China, 300-1400 m |
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(28) |
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| 28 (27) |
Corolla lobes rounded. |
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28 M. simpliciloba |
| + |
Corolla lobes acute or acuminate |
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(29) |
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| 29 (28) |
Climbers; leaves 8-13 × 3-6 cm. |
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6 M. densiflora |
| + |
Erect shrubs; leaves 6-20 × 3-10 cm. |
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8 M. elliptica |
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| 30 (19) |
Corolla tube ca. 40 mm. |
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16 M. kwangtungensis |
| + |
Corolla tube 5-37 mm |
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(31) |
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| 31 (30) |
Stems densely tomentose |
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(32) |
| + |
Stems appressed pubescent to glabrescent |
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(33) |
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| 32 (31) |
Erect shrubs; calyx lobes 3-4 mm; corolla lobes acuminate. |
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21 M. mollissima |
| + |
Climbers; calyx lobes 5-7 mm; corolla lobes rounded. |
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28 M. simpliciloba |
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| 33 (31) |
Most or all calyx lobes 1-2 mm |
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(34) |
| + |
Most or all calyx lobes 2.3-5.9 mm |
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(38) |
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| 34 (33) |
Leaves with 4-6 pairs of lateral veins; Hainan |
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(35) |
| + |
Leaves with 6-8 pairs of lateral veins; mainland |
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(36) |
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| 35 (34) |
Corolla tube 25-31 mm; calycophylls elliptic to ovate. |
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1 M. antiloga |
| + |
Corolla tube 13-15 mm; calycophylls oblong-elliptic or oblanceolate. |
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18 M. lotungensis |
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| 36 (34) |
Corolla tube ca. 12.8 mm. |
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2 M. breviloba |
| + |
Corolla tube 18-21 mm |
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(37) |
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| 37 (36) |
Leaves 5.5-7.5 × 3-4.7 cm; petioles 0.5-0.6 cm. |
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4 M. chingii |
| + |
Leaves 10-17 × 2.5-6 cm; petioles 0.5-1.6 cm. |
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Schizomussaenda dehiscens |
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| 38 (33) |
Corolla with tube ca. 32 mm and lobes ca. 4 mm; Hainan. |
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20 M. membranifolia |
| + |
Corolla with tube 11-25 mm and lobes 2-6 mm; widespread, including Hainan |
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(39) |
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| 39 (38) |
Corolla lobes ca. 2 mm |
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(40) |
| + |
Corolla lobes 3.5-6 mm; widespread |
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(41) |
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| 40 (40) |
Corolla tube ca. 22 mm; Assam (not known from China, included here for reference). |
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M. parryorum (see comment above)
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| + |
Corolla tube 13-15 mm; Hainan. |
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18 M. lotungensis |
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| 41 (39) |
Inflorescences subcapitate, mostly unbranched; flowers 3-20, sessile to subsessile (though fruit may be stipitate) |
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(42) |
| + |
Inflorescences branched, at least to one order, i.e., tripartite; flowers 5 to numerous, sessile to long pedicellate |
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(43) |
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| 42 (41) |
Corolla tube 22-25 mm; stems generally without axillary short shoots. |
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6 M. densiflora |
| + |
Corolla tube 11-20 mm; stems often with axillary short shoots with small leaves. |
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25 M. pubescens |
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| 43 (41) |
Stems strigose; near sea level to 1400 m. |
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7 M. divaricata |
| + |
Plants tomentose to glabrescent; 1200-1400 m. |
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28 M. simpliciloba |
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List of lower taxa
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