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96. Wendlandia Bartling ex Candolle, Prodr. 4: 411. 1830.

水锦树属 shui jin shu shu

Authors: Tao Chen & Charlotte M. Taylor

Shrubs or trees, unarmed; branches sometimes flattened. Raphides absent. Leaves opposite or occasionally in whorls of 3, without or rarely with domatia, with margins rarely denticulate (apparently where scabrous projections form triangular thickenings), infrequently with venation not visible abaxially; stipules persistent or caducous as a whole or by fragmentation, interpetiolar, triangular to pandurate or leaflike (i.e., generally ovate and narrowed to a stipitate base), entire or rarely bilobed, erect and flat to longitudinally folded and/or spreading to reflexed. Inflorescences terminal, cymose, thyrsoid, or paniculiform, many flowered, sessile to pedunculate, bracteate. Flowers sessile or pedicellate, bisexual, monomorphic, often fragrant. Calyx limb 5-lobed. Corolla white, purple, or red, tubular, salverform, or funnelform, glabrous or pubescent inside, with top portion of tube often reflexed at anthesis; lobes (4 or)5, imbricate in bud, strongly reflexed to revolute at anthesis. Stamens (4 or)5, inserted in corolla tube near throat, partially to fully exserted; filaments short to developed; anthers dorsifixed, sometimes with connective prolonged in short apical and/or basal appendages. Ovary 2(or 3)-celled, ovules numerous in each cell on peltate axile placentas; stigma bifid or rarely clavate and shortly emarginate at apex (Wendlandia pendula), exserted. Fruit capsular, subglobose, loculicidally dehiscent across apical portion into 2 valves with valves later sometimes splitting septicidally, papery to woody, with calyx limb persistent; seeds numerous, small, compressed, sometimes narrowly winged; testa membranous, reticulate-striate; endosperm fleshy.

At least 90 species: mainly in tropical and subtropical Asia and a few in the Pacific region; 31 species (21 endemic) in China.

The flowers seem to open generally all at once on an individual plant and perhaps in the population. They are reported in several species to be strongly fragrant. W. C. Chen (in FRPS 71(1): 191. 1999) described the fruit as rarely septicidal, but this has not been reported by other authors and has not been seen on the specimens studied.

Wendlandia has been studied by several authors, notably in a comprehensive monograph by Cowan (Notes Roy. Bot. Gard. Edinburgh 16: 233-313, t. 232-235. 1932, with supplemental notes shortly afterward; Notes Roy. Bot. Gard. Edinburgh 18: 183-188. 1934) and then in treatments of the Chinese species by F. C. How (Sunyatsenia 7(1-2): 32-62. 1948) and later W. C. Chen (Acta Phytotax. Sin. 21: 277-284, 386-403. 1983).

Many Wendlandia species seem to be morphologically variable with much of the variation difficult to partition into distinct clusters, as discussed by Cowan (loc. cit. 1932). Cowan also evaluated characters used by various authors to distinguish species and concluded that many were variable and/or incorrectly interpreted, although he used several of these to distinguish infraspecific taxa. With more species known now, there also seems to be variation within species in some of the characters that Cowan considered reliable to distinguish species. W. C. Chen (loc. cit. 1983) used Cowan’s characters to delimit some of his new species, but circumscribed other species to include a relatively wide range of variation in some of the same features (e.g., W. pingpienensis).

Cowan recognized four series and four subseries, distinguished by stigmas, anther, and stipule morphology as well as habit. These taxa were accepted by W. C. Chen in FRPS (loc. cit. 1999: 195, 196, 200, 202, 208, 218, 221); however, the classification following Cowan’s treatment of several Chinese Wendlandia species described by recent authors is problematic because flowers are needed but are unknown for these. The key published by W. C. Chen (loc. cit. 1999: 192-195) closely follows that of Cowan except some leads were numbered incorrectly while others appear to be missing, and the updated information on morphological variation and geographic distribution was not added to the key so could be problematic to use. The key to Wendlandia species here follows that of W. C. Chen, to outline for reference the species distinctions in that treatment; however, it has been augmented with the new morphological and distributional information.

In addition to the species treated here, F. C. How (loc. cit.: 43) reported Wendlandia ternifolia Cowan provisionally from China based on Tsang 21937, Liang 67941, and Zoo 69290. However, W. C. Chen in FRPS (loc. cit. 1999: 192) commented that study of Tsang 21937 showed the calyx to have stiff pubescence, which is inconsistent with the protologue description of W. ternifolia, while the other two specimens cited have no flowers in adequate condition for identification. Therefore, W. ternifolia was excluded from the Chinese flora; no new or alternative identification was given by Chen for these specimens.

1 Most or all flowers with well-developed pedicels 2-6 mm; stipules triangular to broadly triangular, acute, cuspidate, or narrowly spatulate   (2)
+ Flowers sessile to shortly pedicellate, with pedicels up to 2 mm; stipules variously shaped, generally triangular, spatulate, ovate, suborbicular, leaflike, or pandurate   (7)
2 (1) Shrubs or trees; petioles 0.5-1.5 cm; corolla lobes 1-2.5 mm; anthers 0.8-2 mm, subsessile or with filaments less than 1 mm   (3)
+ Low shrubs or sometimes trees up to 12 m tall; petioles 0.1-0.6 cm; corolla lobes 3.5-8 mm; anthers 2-3.5 mm, exserted on filaments 4-6 mm (W. ser. Montigenae Cowan)   (5)
3 (2) Corolla with lobes ca. 1 mm, less than half as long as tube; anthers ca. 1 mm.   12 W. laxa
+ Corolla with lobes 1.5-2.5 mm, half as long as tube to ± equal to tube; anthers 1.2-2 mm   (4)
4 (3) Leaves abaxially with secondary and higher order venation well marked; corolla white to yellow, with tube 1.5-4 mm.   8 W. formosana
+ Leaves abaxially with secondary and higher order venation hardly visible; corolla purplish red, with tube 2.3-3 mm.   16 W. longipedicellata
5 (2) Shrubs or trees 1-12 m tall; leaves elliptic-oblong, oblong-lanceolate, oblanceolate, or elliptic, 3-14.5 cm, glabrous on both surfaces; stipules spatulate to lanceolate or leaflike (i.e., ovate and stipitate); Hainan.   18 W. merrilliana
+ Leaves elliptic-lanceolate, lanceolate, narrowly lanceolate, ovate, or suborbicular, 0.8-3 cm, pubescent on one or both surfaces; stipules triangular, lanceolate-triangular, or ovate; mainland   (6)
6 (5) Leaves acute, shortly acuminate, or obtuse at apex, with secondary veins ca. 3 pairs, visible abaxially; calyx lobes linear-oblong, spatulate, or oblanceolate.   15 W. longidens
+ Leaves obtuse at apex, veins not visible abaxially; calyx lobes triangular or lanceolate.   28 W. subalpina
7 (1) Stipules triangular, ovate, broadly triangular, or leaflike, at apex acute, cuspidate, ligulate, or aristate, erect or slightly spreading with age   (8)
+ Stipules triangular, ovate, suborbicular, leaflike, or pandurate, with apex obtuse, rounded, shortly acuminate, 2-lobed, or ligulate and usually spreading to reflexed   (17)
8 (7) Leaves subsessile or shortly petiolate, petioles up to 3 mm   (9)
+ Leaves petiolate, petioles 3-25 mm   (10)
9 (8) Leaves opposite or ternate, ovate to ovate-lanceolate, 1.8-4 cm wide, drying papery, margins not thickened nor edged with a vein; stigma clavate.   22 W. pendula
+ Leaves opposite, narrowly elliptic to narrowly lanceolate, 0.4-1.2 cm wide, drying thinly leathery, marginally edged with a thickened vein; stigma deeply bilobed.   25 W. salicifolia
10 (8) Stipule apex prolonged, ligulate, 5-7 mm.   9 W. grandis
+ Stipule apex acute, cuspidate, or shortly acicular, 0.5-2 mm   (11)
11 (10) Calyx lobes narrowly triangular to linear, 1-1.2 mm, equal or unequal on an individual flower with at least one of lobes longer than 1 mm   (12)
+ Calyx lobes lanceolate, triangular, or spatulate, 0.3-1.2 mm, equal or subequal   (13)
12 (11) Petioles 0.8-1.2 cm; stipules ca. 5 mm; corolla lobes ca. 1 mm.   6 W. cavaleriei
+ Petioles 0.8-1.5 cm; stipules 2-3 mm; corolla lobes 1.5-2.2 mm.   13 W. ligustrina
13 (11) Leaves strigillose to puberulent on both surfaces; corolla tube 8.5-11 mm.   1 W. aberrans
+ Leaves strigillose, puberulent, tomentulose, villous, or glabrescent on both surfaces; corolla tube 1.5-4 mm   (14)
14 (13) Calyx lobes 0.3-0.5 mm.   8 W. formosana
+ Calyx lobes 0.8-1 mm   (15)
15 (14) Inflorescences smaller, 5-6 × 3-6 cm; calyx glabrous or sparsely villosulous; leaves sparsely pubescent to glabrous abaxially.   4 W. brevipaniculata
+ Inflorescences larger, 9-17 × 6-22 cm; calyx glabrous to densely pilosulous, hirtellous, or villosulous; leaves glabrous to strigillose, tomentose, or villous abaxially   (16)
16 (15) Inflorescences 9-17 × 9-22 cm; corolla outside with tubes and lobes variously glabrous, strigillose, and/or villosulous (including glabrous on tube and densely villosulous on lobes in subsp. affinis).   29 W. tinctoria
+ Inflorescences 9-12 × 6-11 cm; corolla outside glabrous on tube and densely villosulous or hirtellous on lobes.   31 W. villosa
17 (7) Anthers linear-lanceolate, 1.3-2 mm, exserted from corolla on developed filaments; corolla 7-9.4 mm; capsule 2.5-3 mm in diam. (W. ser. Euexsertae Cowan p.p., W. subser. Orbiculares Cowan).   27 W. speciosa
+ Anthers lanceolate or elliptic, 0.5-1.7 mm, partially included, sessile or with filaments less than 1 mm; corolla 2-13.5 mm; capsule 1-2.5 mm in diam. (fruit unknown in W. augustinii, W. erythroxylon, W. myriantha, W. parviflora, W. pubigera) (W. ser. Wendlandia p.p., W. subser. Paniculatae Cowan)   (18)
18 (17) Leaves abaxially densely strigose or sericeous along midrib; inflorescences smaller, 4-10.5 × 3-10 cm; corolla 11.5-13.5 mm, red or purple; capsule 2-2.5 mm in diam.   3 W. bouvardioides
+ Leaves abaxially glabrous, glabrescent, or sparsely to densely strigillose, puberulent, pilosulous, tomentose, hispidulous, or hirtellous; inflorescences often larger, 4-30 × 4-25 cm; corolla 2-6 mm, white, pale green, pale yellow, or yellowish green; capsule 1-2 mm in diam. (unknown in W. augustinii, W. erythroxylon, W. myriantha, W. parviflora, W. pubigera)   (19)
19 (18) Corolla tube 1-1.5 mm, with lobes equal to, longer than, or sometimes slightly shorter than tube; leaves 5-17 × 2-8.5 cm   (20)
+ Corolla tube 2-5 mm, with lobes clearly shorter than tube; leaves 3-26 × 1.5-14 cm   (21)
20 (19) Leaves glabrescent or sparsely strigillose to pilosulous abaxially; Guangdong, Guangxi.   5 W. brevituba
+ Leaves densely ferruginous pubescent abaxially; Yunnan.   21 W. parviflora
21 (19) Leaves drying leathery, abaxially with secondary veins plane and tertiary venation hardly or not visible   (22)
+ Leaves drying papery to leathery, abaxially with secondary veins raised and tertiary venation easily visible and usually also raised (i.e., most common condition)   (23)
22 (21) Calyx densely hirtellous.   14 W. litseifolia
+ Calyx glabrous to sparsely strigillose.   20 W. oligantha
23 (21) Calyx glabrous to sparsely puberulent, with lobes 1.2-2 mm, entire to ciliate   (24)
+ Calyx with hypanthium portion and limb glabrous, glabrescent, or pilosulous, puberulent, tomentose, hirtellous, pilose, strigose, or strigillose, with lobes 0.3-1 mm   (26)
24 (23) Calyx lobes 1.2-2 mm; corolla tube 2-3.5 mm; Guangdong, Hainan.   10 W. guangdongensis
+ Calyx lobes 1.2-1.5 mm; corolla tube 2.5-4 mm; Guangxi, Taiwan   (25)
25 (24) Leaves entire or scabrous to denticulate marginally; stipules entire; Taiwan.   7 W. erythroxylon
+ Leaves entire, smooth or ciliolate marginally; stipules entire or 2-lobed; Guangxi.   19 W. myriantha
26 (23) Leaves abaxially moderately to densely hirtellous, strigillose, pilosulous, pilose, or strigose with pubescence on lamina spreading and/or partly obscuring its surface   (27)
+ Leaves abaxially glabrous to sparsely strigillose or hispidulous on lamina and strigillose to moderately hispidulous or hirtellous on principal veins   (28)
27 (26) Leaves scaberulous on lamina adaxially; stipules usually equal to or only slightly wider than stem; corolla inside with short trichomes confined to lower to upper part of tube; Yunnan.   26 W. scabra
+ Leaves sparsely hirtellous, hispidulous or scaberulous adaxially; stipules usually twice or more as wide as stem; corolla hirsute inside upper part of tube and throat.   30 W. uvariifolia
28 (26) Leaves abaxially glabrous on lamina and sparsely strigillose and/or hirtellous on principal veins; Taiwan.   17 W. luzoniensis
+ Leaves abaxially glabrous to hispidulous or strigillose; mainland   (29)
29 (28) Stipules with apex ligulate, obtuse, often longitudinally folded.   9 W. grandis
+ Stipules with apex oblanceolate to suborbicular, folded to usually flat   (30)
30 (29) Corolla tube 4-5 mm; Yunnan   (31)
+ Corolla tube 2-3 mm   (32)
31 (30) Stipules with apical portion 3-4 mm wide; leaves with secondary veins 7-9 pairs.   2 W. augustinii
+ Stipules with apical part broad, ca. 10 mm wide; leaves with secondary veins 7-13 pairs.   23 W. pingpienensis
32 (30) Leaves smaller, 7-10.5 × 2-4 cm, abaxially glabrous on lamina and sparsely pubescent only on veins; secondary veins 7 or 8 pairs; Yunnan.   11 W. jingdongensis
+ Leaves larger, 14.5-18 × 6-7 cm, abaxially glabrous to sparsely pubescent on lamina with pubescence denser along veins; secondary veins 10 or 11 pairs; Guangxi.   24 W. pubigera

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