CAPRIFOLIACEAE (sensu stricto) [Draft]

忍冬科 

Qin-er Yang (杨亲二);

Shrubs or woody vines, rarely small trees or herbs. Leaves opposite, rarely verticillate, simple or pinnately compound; stipules absent or very small. Inflorescence basically cymose or with pairs of flowers sometimes joined toward bases of ovaries. Flowers bisexual, regular or irregular. Calyx 4- or 5-toothed or lobed; tube adnate to ovary. Corolla epigynous, gamopetalous; lobes 5, spreading, sometimes 2-lipped, mostly imbricate in bud. Stamens (4 or)5, arising from corolla tube, alternating with lobes; anthers free, 2-celled, opening by longitudinal slits, mostly introrse. Carpels 3–5, united; ovary inferior, 1–5-loculed; placenta axile; ovules 1 to many per locule, pendulous; style solitary; stigma capitate or lobed. Fruit a berry or a achene-like drupe, rarely a capsule. Seeds 1 to many; embryo small, straight; endosperm copious.

About thirteen genera and ca. 500 species: mostly in temperate regions of eastern North America and eastern Asia; twelve genera and ca. 200 species in China.

Hsu Ping-sheng, Hu Jia-qi & Wang Han-jin. 1988. Caprifoliaceae. In: Hsu Ping-sheng, ed., Fl. Reipubl. Popularis Sin. 72: 1–284.

add note about former and modern circumscriptions.

Doubtful taxa: Actinotinus Oliver, Actinotinus sinensis Oliver 1888. Icon. Plant. 18 t. 1740 = Chimerous material; Aesculus chinensis var. wilsonii leaf associated with an inflorescence fragment of Viburnum plicatum var. tomentosum (see FOC Hippocastaneaceae draft and Malécot, V. 2004. La diversité des Viburnum. Hommes et plantes 49: 22–27).

 

1a.     Herbs perennial; leaves connate-perfoliate, otherwise pinnate ..............................  1. Triosteum

1b.     Shrubs or woody vines, rarely small trees or prostrate subshrubs; leaves not connate, otherwise undivided.

2a.    Woody vines .................................................................................................  5. Lonicera

2b.    Shrubs or small trees.

3a.     Leaves triplinerved; flowers 6 or 7 in a head, forming a terminal panicle .  2. Heptacodium

3b.     Leaves pinnativeined.

4a.     Flowers paired on peduncle, calyx tubes often ± connate .........................  5. Lonicera

4b.     Calyx tubes of paired flowers separate.

5a.    Branchlets hollow; fruit a many seeded berry .................................  4. Leycesteria

5b.    Branchlets solid ....................................................................  3. Symphoricarpos

 

1. TRIOSTEUM Linnaeus, Sp. Pl. 1: 176. 1753.

莛子藨属  ting zi biao shu

Herbs perennial, with underground rhizomes. Stem erect, unbranched. Leaves simple, opposite, obovate, entire, undulate, or emarginate to deeply divided. Inflorescences in clusters on a terminal axis or in axils of leaves. Calyx 5-lobed; lobes short or long and foliaceous, persistent. Corolla whitish, yellow, or purple, tubular-campanulate, base gibbous on one side; lobes 5, unequal, imbricate, bilabiate, upper labium 4-lobed, lower labium entire. Stamens 5, inserted in corolla tube; anthers introrse. Ovary 3–5-loculed; ovule 1 per locule, pendulous; styles filiform; stigmas discoid, 3–5-lobed. Fruit a berry-like drupe, subglobose, leathery or fleshy; pyrenes cartilaginous; seeds 2 or 3, narrowly orbicular; endosperm fleshy, embryo small.

About six species: C and E Asia, North America; three species in China.

 

1a.     Inflorescence in clusters on a terminal axis or in axils of leaves; leaves entire, or slightly repand to sinuous or slightly lobed ..................................................................................................  3. T. sinuatum

1b.     Inflorescence in clusters on a terminal axis.

2a.    Leaves entire, base connate and perfoliate ............................................  1. T. himalayanum

2b.    Leaves pinnatifid, base not connate .......................................................  2. T. pinnatifidum

 

1. Triosteum himalayanum Wallich in Roxburgh, Fl. Ind. 2:180. 1824.

穿心莛子藨  chuan xin ting zi biao

Echium connatum H. Léveillé; Triosteum erythrocarpum H. Smith; T. fargesii Franchet.

Herbs perennial. Stem 40–60 cm tall, rarely 2-branched at apex at anthesis, densely hirsute and glandular hairy. Leaves usually 9 or 10 pairs; leaf blade obovate-elliptic to obovate-oblong, 8–16 × 5–15 cm, abaxially relatively densely long hirsute on veins and mixed glandular hairy, adaxially long hirsute, base connate and perfoliate, apex acute. Cymes 2–5-whorled, clustered at apex of stem or sometimes forming a spike on branches. Calyx lobes triangular-orbicular, hirsute and glandular hairy, constricted between tube and lobes. Corolla yellow-green, tube purple-brown inside, ca. 1.6 cm, ca. 3 × as long as calyx, outside glandular-hairy; tube curved at base, gibbous on one side. Stamens inserted at middle of corolla tube; filaments yellowish, long, slender; anthers yellow, oblong. Fruit red, subglobose, 10–12 cm in diam., crowned with a short beak consisting of persistent calyx lobes and constricted calyx tube, hirsute and glandular hairy. Fl. Jun–Jul, fr. Aug–Sep.

Mountain slopes, needle-leaved forests, streamsides, grasslands; 1800–4100 m. ??Henan, Hubei, Hunan, Shaanxi, Sichuan, Yunnan, Xizang [??Bhutan, India (Sikkim), Nepal].

2. Triosteum pinnatifidum Maximowicz, Bull. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 3, 27: 476. 1881.

莛子藨  ting zi biao

Triosteum intermedium Diels & Graebner; T. rosthornii Diels & Graebner.

Herbs perennial. Stem to 60 cm tall, 2-branched at apex at anthesis, white hirsute and glandular hairy. Leaves pinnatifid, subsessile; leaf blade obovate to obovate-elliptic, 8–20 × 6–18 cm, adaxially sparsely hirsute, more densely so on veins and margin, base cuneate to broadly cuneate, apex acuminate; leaves at base of stem sometimes not divided. Cymes opposite, each 3-flowered; peduncule absent, sometimes inflorescence subtended by ovate, entire bracts, clustered into a short spike at apex of stem or branches. Calyx tube hirsute and glandular hairy; lobes triangular, ca. 3 mm. Corolla yellow-green, narrowly campanulate, ca. 1 cm; tube curved at base, shallowly gibbous on one side, glandular hairy; lobes orbicular, short, inside purplish punctate. Stamens inserted below middle of corolla tube; filaments short; anthers oblong. Styles long pubescent at base; stigmas cuneate-capitate. Fruit white, ovoid, ca. 10 cm, crowned with persistent calyx lobes, hirsute and glandular-hairy. Fl. May–Jun, fr. Aug–Sep. 2n = ca. 18*, 36.

Dark needle-leaved forests, sunny places on streamsides; 1800–2900 m. Gansu, Hebei, Henan, Hubei, Ningxia, Qinghai, Shaanxi, Shanxi, Sichuan [Japan].

3. Triosteum sinuatum Maximowicz, Bull. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 3, 15: 373. 1871.

腋花莛子藨  ye hua ting zi biao

Herbs perennial. Stem to 60 cm tall, densely hirsute and glandular hairy. Leaves orbicular-ovate or oblong, ca. 14 × 5 cm, sparsely adpressed-hirsute, densely ciliate on veins and margin, ca. 1/3 from base abruptly narrowed and spatulate-ovate or oblong, base sometimes amplexicaul, margin entire or slightly repand to sinuous or slightly lobed, apex acuminate. Cymes axillary, 1–3-flowered. Calyx lobes lanceolate or narrowly elliptic, 6–10 mm, subequaling tube, outside hirsute or glandular hairy. Corolla yellow-green, inside purple-brown. Stamens: 2 longer, other 3 equal in length; anthers narrowly elliptic. Styles filiform; stigmas exserted, globose. Fruit greenish, globose, 1–1.5 cm in diam, sessile, densely hirsute and glandular-hairy. Fl. Jun, fr. Sep–Oct. 2n = 18.

Forests, streamsides; 800–900 m. Jilin, Liaoning, Xinjiang [Japan].

2. HEPTACODIUM Rehder in Sargent, Pl. Wilson. 2: 617. 1916.

七子花属  qi zi hua shu

Shrubs or small trees, deciduous. Winter buds perulate. Leaves opposite, entire, triplinerved, estipulate. Flowers sessile, forming a terminal panicle composed of numerous whorled head-like cymes, with a pair of cymes of 3 flowers and a single terminal flower in each whorl. Involucre orbicular or ovate, large, persistent, including 10 bracts and bracteoles often decussate and densely sericeous, with 4 outside, 6 inside. Calyx tube turbinate, densely setaceous, 5-lobed; lobes narrowly elliptic, enlarged and persistent after anthesis. Corolla white, tubular-infundibuliform; tube slightly curved, base not equal-sided, 5-lobed, slightly 2-labiate; lobes narrowly elliptic, upper labium erect, 3-lobed; lower labium spreading or revolute, 2-lobed. Stamens 5; filaments inserted at middle of corolla tube, longer than corolla lobes; anthers narrowly elliptic. Ovary 3-loculed, 2 locules with numerous sterile ovules, remaining locule with 1 fertile ovule; styles hairy, stigmas discoid. Fruit an achene-like drupe, narrowly ellipsoid, leathery, crowned with persistent and enlarged calyx lobes, 3-loculed, 2 locules vacant and compressed, remaining locule with 1 seed; seed subterete, compressed in upper part, pericarp membranous; endosperm fleshy, embryo shortly terete, at base of seed.

One species; endemic to China.

1. Heptacodium miconioides Rehder in Sargent, Pl. Wilson. 2: 618. 1916.

七子花  qi zi hua

Heptacodium jasminoides Airy-Shaw.

Plants to 7 m tall. Young branches red-brown, sparsely pubescent; bark of stem gray-white. Petiole 1–2 cm; leaf blade ovate or oblong-ovate, 8–15 × 5–9 cm, thickly leathery, abaxially sparsely pubescent on veins, base obtuse or slightly cordate, apex long caudate. Inflorescence a panicle, nearly pyramidal, 8–15 × 5–9 cm; branches spreading, upper ones ca. 1.5 cm, lower ones 2.5–4 cm; small inflorescence capitate, bracteoles variously shaped, unequal in size, outermost pair emarginate. Flowers fragrant. Calyx lobes 2–2.5 mm, equaling tube, densely hirsute. Corolla 1–1.5 cm, densely downward pubescent. Fruit 1–1.5 × ca. 3 mm, 10-striate, sparsely hirsute-sericeous, persistent calyx with conspicuous midvein; seeds 5–6 mm. Fl. Jun–Jul, fr. Sep–Nov.

* Cliffs, scrub, forests; 600–1000 m. Anhui, Hubei, Zhejiang.

This is a rare species.

3. SYMPHORICARPOS Duhamel du Monceau, Traité Arb. Arbust. 2: 295. 1755.

毛核木属  mao he mu shu

Shrubs, deciduous. Winter buds with 2 pairs of scales. Leaves opposite, entire or sinuous, shortly petiolate, estipulate. Flowers clustered or solitary in axils of leaves at apices of lateral branches, forming a spike or a panicle. Calyx cupular, 4- or 5-lobed. Corolla reddish or white, campanulate to infundibuliform or hypocrateriform, 4- or 5-lobed, regular; tube slightly gibbous at base, inside long pubescent. Stamens 4 or 5, inserted in corolla tube, included or slightly exserted; anthers introrse. Ovary 4-loculed, 2 locules with several sterile ovules, remaining 2 each with 1 ovule; styles slender; stigmas capitate or slightly 2-lobed. Fruit a berry-like drupe with 2 pyrenes, white, red or black, globose, ovoid, or ellipsoid; pyrenes ovoid, ± compressed; seeds with endosperm, embryo small.

Sixteen species: 15 in North America and Mexico, one species in China (the one endemic).

1. Symphoricarpos sinensis Rehder in Sargent, Pl. Wilson. 1: 117. 1911.

毛核木  mao he mu

Shrubs erect, 1–2.5 m tall. Young branches red-brown, slender, pubescent. Petiole 1–2 mm; leaf blade abaxially gray-white, adaxially green, rhombic-ovate to ovate, 1.5–2.5 × 1.2–1.8 cm, both surfaces glabrous, triplinerved, lateral veins inconspicuous, base cuneate or broadly cuneate, margin entire, apex acute or obtuse. Flowers small, sessile, solitary in axils of short, small, subulate bracts, forming a short, small, terminal spike; lower bracts foliaceous and longer. Calyx tube ca. 2 mm; lobes 5, ovate-lanceolate, ca. 1 mm, apex acute, glabrous. Corolla white, campanulate, 5–7 mm; lobes ovate, slightly shorter than tube, glabrous. Stamens 5, inserted at middle of corolla tube, equaling or slightly exceeding corolla; anthers white, ca. 2 mm. Styles 6–7 mm, glabrous; stigmas capitate. Fruit blue-black, ovoid, ca. 7 mm, crowned with a small beak, pruinose; drupelets 2, densely long pubescent. Fl. Jul–Sep, fr. Sep–Nov.

* Scrub; 600–2200m. S Gansu, Guangxi, W Hubei, Shaanxi, E Sichuan, N Yunnan.

4. LEYCESTERIA Wallich in Roxburgh, Fl. Ind. 2: 181. 1824.

鬼吹箫属  gui chui xiao shu

Shrubs or subshrubs, deciduous. Young branches often [??]vacant. Leaves simple, opposite, margin entire or serrate, rarely shallowly divided; stipules present or absent. Inflorescence spicate or thyrsoid-cymose, terminal or axillary, sometimes clustered and capitate, often with conspicuous leaf-like bracts. Calyx lobes 5. Corolla white, pink, or purple-red, sometimes orange-yellow, infundibuliform, regular; lobes 5. Stamens 5; anthers dorsifixed. Ovary 5–8(–10)-loculed, with numerous ovules per locule; styles long, slender; stigmas peltate or capitate. Fruit a berry, with persistent calyx; seeds minute, numerous.

About eight species: Sino-Himalayan region; five species in China (one endemic).

??Leycesteria crocothyrsos Airy-Shaw.

 

1a.     Flowers 2 per whorl.

2a.    Stipules absent; leaves subentire or dentate, petioles 5–18 mm; filaments and ovary glabrous; bracts shorter than ovary ....................................................................................  4. Leycesteria gracilis

2b.    Stipules present; leaves serrate except at base, petioles 1–2 mm; filaments and ovary pubescent; bracts longer than ovary ..................................................................  5. Leycesteria glaucophylla

1b. Flowers 6 per whorl.

2a.    Spikes with only 1 whorl of flowers; corolla sparsely glandular hairy  2. Leycesteria sinensis

2b.    Inflorescence with 2 to several whorls of flowers; corolla outside pubescent or glabrous.

3a.     Stipules absent; abaxial leaf surfaces, petioles, and inflorescences sparsely adpressed pubescent or subglabrous; corolla outside pubescent ......................................  1. Leycesteria formosa

3b.     Stipule present; abaxial leaf surfaces, petioles, and inflorescences densely tomentose; corolla outside glabrous .................................................................................  3. Leycesteria stipulata

 

1. Leycesteria formosa Wallich in Roxburgh, Fl. Ind. 2: 182. 1824.

鬼吹箫  gui chui xiao

Leycesteria formosa var. brachysepala Airy-Shaw; L. formosa var. glandulosissima Airy-Shaw; L. formosa var. stenosepala Rehder; L. limprichtii H. Winkler.

Shrubs, 1–5 m tall, often sparsely or densely dark red short glandular hairy throughout. Branchlets, petioles, peduncles, bracts, and calyx lobes adpressed pubescent. Petiole 5–12(–15) mm; leaf blade ovate-lanceolate or ovate-oblong to ovate, 4–13 × 2–6 cm, papery, abaxially sparsely adpressed-pubescent or subglabrous, adaxially pubescent, more densely so on midvein, base rounded to subcordate or broadly cuneate, margin often entire, sometimes remotely dentate or irregularly sinuate, apex long caudate to acuminate. Spikes terminal or axillary, 1- to several whorled, with 6 flowers per whorl; peduncle (8–)10–25(–30) mm; bracts leaf-like, green, purplish, or purple-red; bracteoles very small, less than 1 mm. Calyx tube oblong, 3–4 mm, densely glandular hairy; limb 5-lobed; lobes orbicular-ovate or lanceolate to linear-lanceolate, 1–9 mm, sometimes 2 longer and remaining 3 shorter. Corolla white or pink, sometimes purple-red, infundibuliform, (1.2–)1.4–1.8 cm, outside pubescent; lobes orbicular-ovate, ca. 5 mm. Stamens subequaling corolla; anthers oblong. Styles slightly exceeding corolla; stigmas peltate; ovary 5-loculed. Fruit red, turning black-purple, ovoid or subglobose, 5–7 mm in diam., with persistent calyx lobes; seeds minute, numerous, brownish, broadly ellipsoid to oblong, slightly compressed, 1.2–1.5 mm. Fl. (May–)Jun–Sep(–Oct), fr. (Aug–)Sep–Oct. 2n = 18.

Forests, forest margins, scrub; 1100–3500 m. W Guizhou, W Sichuan, Xizang, Yunnan [Bhutan, India, ??Kashmir, Myanmar, Nepal, Pakistan].

2. Leycesteria sinensis Hemsley, Hooker’s Icon. Pl. 27: t. 2633. 1900.

华鬼吹箫  hua gui chui xiao

Shrubs, 1–2 m tall. Branchlets subherbaceous, glabrous. Petiole ca. 5 mm, pubescent with sparse coarse hairs and glandular hairs; leaf blade ovate-lanceolate or ovate, 2.5–6.5 cm, papery, abaxially sparsely pubescent on veins, ciliate, adaxially pubescent, base rounded or subtruncate, margin dentate, apex long caudate to mucronate. Spikes terminal on branchlets, 1-whorled, composed of 6 flowers; peduncle 5–10 mm, sparsely glandular hairy; bracts 6, leaf-like, both sides minutely glandular hairy, lowest pair largest, broadly ovate, 1–2 cm, upper 2 pairs ovate to lanceolate, 8–10 mm; bracteoles lanceolate or oblanceolate, smaller, shorter than calyx tube, shortly glandular hairy. Calyx tube 5–6 mm, with dense, brown-yellow glandular hairs; limb shallowly cupular in lower part, 5-lobed; lobes lanceolate to linear-lanceolate, (2–)3–5(–7) mm, sparsely minutely glandular hairy and glandular ciliolate. Corolla pink, infundibuliform, 1.3–1.5 cm, outside sparsely minutely glandular hairy; lobes orbicular-ovate, ca. 3 mm. Stamens subequaling corolla lobes. Ovary 5-loculed; stigmas peltate, slightly exserted from corolla. Fruit red, ovoid-orbicular; seeds minute, numerous, subrounded, ca. 1 mm in diam. Fl. May, fr. Sep.

* Forests, forest margins; 1900–2700 m. Yunnan.

3. Leycesteria stipulata (J. D. Hooker & Thomson) Fritsch in Engler & Prantl, Nat. Pflanzenf. 4(4): 169. 1891.

 绵毛鬼吹箫  mian mao gui chui xiao

Lonicera stipulata J. D. Hooker & Thomson, J. Linn. Soc., Bot. 2: 165. 1858; Pentapyxis stipulata (J. D. Hooker & Thomson) J. D. Hooker ex C. B. Clarke.

Shrubs, ca. 3 m tall. Branchlets pubescent, abaxial leaf surfaces, petioles, and inflorescences densely gray-white lanate. Stipules conspicuous, suborbicular, 6–8 mm, abaxially lanate, adaxially pubescent, persistent; petiole 8–10 mm; leaf blade oblong-lanceolate, 8–15(–20) × 3.5–6 cm, thickly papery, glabrous and shiny adaxially, veinlets reticulate, conspicuously impressed, midvein and lateral veins raised abaxially, base rounded, inequilateral, margin slightly involute, remotely shallowly dentate, apex acuminate to long acuminate. Spikes axillary, short, 1–3-whorled, with 6 flowers and 6 brown leaf-like bracts per whorl; bracts orbicular-ovate to lanceolate, exceeding calyx tube; peduncle short, 8–12 mm. Corolla white, ca. 1.5 cm, glabrous. Fruit lanate, ca. 1.9 cm; seeds brownish-yellow, ellipsoid or subglobose, ca. 1 mm. Fl. Mar–Apr, fr. Oct–Nov.

Broad-leaved forests, grassy places; 1300–2000 m. Yunnan [Bhutan, India, Myanmar].

4. Leycesteria gracilis (Kurz) Airy-Shaw, Hooker’s Icon. Pl. 32: t. 3166. 1932.

 纤细鬼吹箫  qian xi gui chui xiao

Lonicera gracilis Kurz, J. Asiat. Soc. Beng. 39(2): 77. 1870.

Shrubs, 1.5–3(–4) m tall. Stipules absent; petiole 6–10 mm, pubescent; leaf blade oblong-lanceolate or oblong-ovate, 7–12(–18) × 2.5–6.5 cm, thickly papery, abaxially strongly glaucous, sparsely pubescent on midvein and lateral veins, adaxially sparsely glandular dentate and sparsely ciliate, pubescent at base of midvein, base rounded, truncate or subcordate, margin slightly involute, apex acuminate to long caudate. Spikes terminal or axillary, 1–3-whorled, with 2 flowers, 1 bract, and 2 bracteoles per whorl; bracts and bracteoles ovate-lanceolate or lanceolate, ca. 1/3 or 1/2 as long as calyx tube, sparsely pubescent and glandular ciliate. Calyx tube 5–6 mm, narrower and slender at apex, sometimes pubescent; limb 2.5–3.5 mm, connate and shallowly cupular in lower part; lobes linear-lanceolate to lanceolate, 1.5–2 mm, often glandular ciliolate. Corolla white, infundibuliform, 1.5–2 cm; lobes ovate or orbicular-ovate, 5–7 mm. Stamens slightly shorter than corolla. Ovary (5–)7- or 8(–10)-loculed, with numerous ovules. Fruit red, turning blue-purple, oblong or ellipsoid, 1–1.3 cm; seeds minute, numerous, subrounded, ca. 1 mm in diam. Fl. (Sep–)Oct–Nov(–Dec), fr. (Mar–)Apr–May.

Forests, thickets; 2000–3800 m. Xizang, Yunnan [Bhutan, India, Myanmar, Nepal].

5. Leycesteria glaucophylla (J. D. Hooker & Thomson) C. B. Clarke in J. D. Hooker, Fl. Brit. India 3: 16. 1880.

西域鬼吹箫  xi yu gui chui xiao

Lonicera glaucophylla J. D. Hooker & Thomson, J. Linn. Soc., Bot. 2: 165. 1858; Leycesteria glaucophylla var. thibetica (H. J. Wang) J. F. Huang; Leycesteria thibetica H. J. Wang.

Shrubs, 1–2 m tall. Branchlets of current year herbaceous, sparsely pubescent and shortly glandular hairy; branchlets of previous year subglabrous. Stipules variously shaped, broadly obovate, reniform, suborbicular, or ovate, 3–5(–10) mm, densely pubescent abaxially, persistent or deciduous; petiole short, ca. 4 mm, pubescent; leaf blade ovate-lanceolate to oblong-lanceolate, 4.5–10.5 × 2–6 cm, papery, abaxially densely gray-white pubescent, adaxially sparsely pubescent and minutely glandular hairy, base rounded, inequilateral, margin slightly involute, sparsely serrate but entire at base, apex acuminate, long acuminate, or mucronate. Inflorescence axillary, 1- or 2-whorled, with 2 flowers per whorl; peduncle short, 5–8 mm; bracts 2, leaf-like, ovate to narrowly ovate, bracteoles 4, ovate to narrowly ovate. Calyx tube 1–2 mm, outside sparsely glandular hairy; lobes narrowly lanceolate or linear-lanceolate, 4–6 mm. Corolla greenish or white; lobes ovate, ca. 4 mm. Stamens slightly shorter than corolla. Ovary 5-loculed, with numerous ovules. Fruit yellow-green, ellipsoid or ovoid-ellipsoid, 1.3–1.4 × ca. 1 cm, long strigose and shortly glandular hairy, 5-loculed, many seeded per locule; seeds yellowish-brown, ovoid-ellipsoid or ovoid, small, ca. 1 mm. Fl. Mar–Sep, fr. Jun–Nov.

Tsuga forest margins; 1800–2600 m. SE Xizang [Bhutan, India, Myanmar, Nepal].

5. LONICERA Linnaeus, Sp. Pl. 1: 173. 1753.

忍冬   ren dong shu

Shrubs erect or dwarf, rarely small trees, sometimes climbers, deciduous or evergreen. Branchlets with white or black-brown pith, sometimes vacant; winter buds with 1 to several pairs of scales, inner scales sometimes enlarged and reflexed, sometimes terminal buds reduced and substituted by 2 lateral buds, very rarely with adventitious buds. Leaves opposite, very rarely verticillate, papery to thickly papery or leathery, margin entire, very rarely dentate or divided, estipulate or very rarely with interpetiolar stipules or linear swellings, sometimes leaves 1 or 2 pairs under inflorescence connate. Flowers usually terminal in pair on axillary peduncles, or sessile and verticillate at apices of branchlets, 3–6 per whorl; each flower pair with a pair of bracts and bracteoles; bracts small or large, leaf-like; bracteoles sometimes united and cupular or urceolate and wrapping calyx tube, rarely absent. Two neighboring calyx tubes separate or partly to completely united; limb 5-lobed or sometimes margin of mouth shallowly undulate or annulate, very rarely with cap-like emergence. Corolla white, yellow, reddish, or purple-red, campanulate, cylindric or infundibuliform, regularly or subregularly 5(or 4)-lobed, or bilabiate and upper labium 4-lobed; tube long or short, often swollen on one side at base or shallowly or deeply gibbous, very rarely long calcarate. Stamens 5; anthers dorsifixed. Ovary 2- or 3(–5)-loculed; styles slender, hairy or glabrous; stigmas capitate. Fruit a berry, red, blue-black, or black, with few to numerous seeds; seeds with very rounded embryo.

About 200 species: North America, Europe, Asia, N Africa; ca. 91 species in China [??#endemic].

The identities of Lonicera confusa de Candolle var. glabrocalyx Miau & X. J. Liu (Acta Sci. Nat. Univ. Sunyatseni 28(4):78. 1989), L. fengkaiensis Miau & X. J. Liu (Acta Sci. Nat. Univ. Sunyatseni 28(4):78. 1989), L. stenantha Pojarkova var. angustifolia C. Y. Yang & J. H. Fan (J. August 1st Agri. College 18(2): 7. 1995), and L. subrotundata C. Y. Yang & J. H. Fan (J. August 1st Agri. College 18(2): 8. 1995), cannot be determined because of the unavailability of the type specimens for examination. One species, L. sempervirens Linnaeus, is occasionally cultivated.

??What about the following names in their respective provincial floras:

??Lonicera apodantha Ohwi, Acta Phytotax. Geobot. 3: 84. 1934 – Taiwan.

??Lonicera caerulea var. salicifolia Dip. – Nei Mongol.

??Lonicera chrysantha f. villosa Rehder – Liaoning.

??Lonicera chrysantha f. crassipes (Nakai) S. L. Tung in Y. L. Chou, Ligneous Fl. Heilongjiang 524. 1986 = Lonicera chrysantha var. crassipes Nakai – Liaoning.

??Lonicera chrysantha var. linearifolia S. W. Liu & T. N. Ho, Fl. Qinghai. 3: 511. 1996.

??Lonicera caprifolium Linnaeus, Sp. Pl. 1: 173. 1753 – Sichuan.

??Lonicera dielsii – Guizhou.

??Lonicera engleriana – Guizhou.

??Lonicera caucasica subsp. govaniana (Wallich ex de Candolle) H. Hara - ?S Tibet.

??Lonicera cyanocarpa var. porphyrantha Marquand & Shaw, J. Linn. Soc., bot. 48: 187. 1929 – SE Xizang.

??Lonicera henryi var. fulvovillosa Ohwi, Acta Phytotax. Geobot.3: 85. 1934.Taiwan: J. Ohwi 2317 (type).

??Lonicera heterophylla var. alpina (Krassn.) Rehder, Syn. Genus Lonicera110. 1903.Sinkiang.

??Lonicera hispida var. bracteata (Royle) Rehder ex Airy-Shaw, Bot. Mag. 157: t. 9360. 1934 – Xizang, China.

??Lonicera hispida var. hirsutior Regel, Bull. Soc. Imp. Naturalistes Moscou1: 4. 1867.Sinkiang: A. Regel.

??Lonicera hispida var. setosa Hook. f. & Thoms., J. Linn. Soc., Bot.2: 166..Tibet: Ward 5988.

??Lonicera hypoglauca var. heterotricha – Fujian.

??Lonicera infundibulum var. rockii Rehder, J. Arnold Arbor.9: 122. 1928.W Szechuan: 12050.

??Lonicera insularis var. lutescens Skvortzow, China J.23: 389. 1935.Manchuria.

??Lonicera insularis var. mandshurica Skvortzow, China J.23: 389. 1935.Manchuria.

??Lonicera japonica var. aureo-reticulata – Jiangsu.

??Lonicera japonica f. macrantha Matsuda, Bot. Mag. (Tokyo) 26: 307. 1912.Chekiang: Chang 54.

??Lonicera japonica f. macrantha Matsuda, Bot. Mag. (Tokyo) 26: 308. 1912.Chekiang: Honda 1278-1281.

??Lonicera japonica var. sinica Rehder, Mitt. Deutsch. Dendr. Ges. 22: 264. 1913.Hunan: 11794, 11200.

??Lonicera koehneana var. poponanthera Hand.-Mazz., Symb. Sin. 7: 1047. 1936.Szechuan: 2483.

??Lonicera korolkowii var. zabelii Rehder, Syn. Genus Lonicera 131. 1903. Sinkiang: A. Regel, Apr. 15, 1879.

??Lonicera macroptera – Guizhou.

??Lonicera maximowiczii var. sachalinensis Fr. Schmidt – Liaoning.

??Lonicera orientalis var. setchuensis Franch., J. Bot. (Morot)10: 311..Shensi: Giraldi 1465, 1466; Szechuan: Farges.

??Lonicera parvifolia – Guizhou.

??Lonicera pseudoproterantha var. longifolia Pamp., Nuov. Giorn. Bot. Ital., n.s.18: 139. 1911.Hupei: Silvestri 3200, 3200a.

??Lonicera pseudoprotherantha var. intermedia Pamp., Nuov. Giorn. Bot. Ital., n.s.18: 139. 1911.Hupei: Silvestri 3199, 3199a.

??Lonicera rupicola f. glabrata Suessenguth, Repert. Spec. Nov. Regni Veg.42: 43. 1937.Kansu: Pater J. Trippner 406.

??Lonicera ruprechtiana var. lanceolata Skvortzow, China J.23: 388. 1935.Manchuria: Kirin & Tengtien.

??Lonicera standishii var. monbeigii W. W. Sm., Notes Roy. Bot. Gard. Edinburgh10: 48. 1917.Yunnan: Monbeig 135 (type).

??Lonicera tricalysioides C. Y. Wu ex P. S. Hsu & H. J. Wang, Acta Phytotax. Sin. 17(4): 77. 1979.

??Lonicera trichogyne Rehder in Sargent, Pl. Wilson. 1: 131. 1911.

??Lonicera trichogyne var. aequipila Hand.-Mazz., Oesterr. Bot. Z.83: 234. 1934.Shansi: Licent 2132.

??Lonicera umbellata – Guizhou.

 

1a.     Flowers solitary, 1- to several verticillate at apices of branchlets, 3–6 per whorl, peduncle present or absent; leaves 1 or 2 pairs under inflorescence connate and discoid at base, very rarely distinct.

2a.    Corolla regular or slightly irregular, not labiate; stamens inserted in upper part of corolla tube.

3a.     Corolla inside and outside yellow, 2.5–3.5 cm, outside sparsely long stiffly hairy and glandular; calyx tube glandular hairy in upper part, lobes stiffly ciliate ................................  87. L. subaequalis

3b.     Corolla outside red-yellow, inside red, ca. 2 cm, outside glabrous; calyx tube not glandular hairy, lobes not ciliate .............................................................................................  88. L. carnosifolia

2b.    Corolla labiate; stamens inserted at base of labia.

4a.     Corolla 5–9 cm; leaves abaxially shortly stiffly hairy or at least densely barbate with transverse short stiff hairs on midvein in lower part, very rarely glabrous ............................  91. L. tragophylla

 

4b.     Corolla not more than 2.5 cm; leaves abaxially glabrous or adpressed pubescent, not barbate with stiff hairs on midvein.

5a.     Leaves 1 or 2 pairs under inflorescence connate and discoid at base; calyx tube glabrous; leaves not ciliate along margin ....................................................................  89. L. yunnanensis

5b.     Leaves 1 pair under inflorescence different from other leaves in shape, but not connate and discoid; calyx tube densely villous and minutely glandular hairy; leaves long ciliate along margin  90. L. ciliosissima

1b.     Flower pairs at apex of peduncle; two opposite leaves not connate at base, and not discoid.

6a.    Climbers, if shrubs prostrate, then leaves leathery.

7a.     Corolla tube long calcarate; two neighboring calyx tubes of a flower pair connate; fruit red       63. L. calcarata

7b.     Corolla tube not calcarate; two neighboring calyx tubes of a flower pair distinct; fruit black or blue-black.

8a.     Shrubs prostrate; leaves leathery, apex obtuse to rounded, sometimes mucronate, or slightly emarginate ....................................................................................................  64. L. crassifolia

8b.     Climbers.

9a.    Young branches, petioles, and peduncle spreading yellow-brown hispid and glandular hairy; leaves at least with two series of rigid ciliae along margin ..........................  72. L. nubium

9b.    Plant pubescent or stiffly hairy, never hispid, sometimes glabrous.

10a.   Leaves or at least young ones abaxially velutinous, not spaced among hairs.

11a.   Young branches densely pubescent, and spreading yellowish brown long stiffly hairy, with hairs longer than 2 mm, or young branches subglabrous.

12a.   Corolla shorter than 3 cm; leaves abaxially gray-white tomentose with both long and short soft hairs .................................................  86. L. similis (var. omeiensis)

12b.   Corolla longer than 4 cm.

13a.   Calyx lobes triangular, to 1 × 1 mm; leaves abaxially gray-white to gray-yellow tomentose with short soft hairs .......................................  86. L. similis

13b.   Calyx lobes triangular-lanceolate to triangular, longer than wide; leaves stiffly hairy, and gray-white or yellow-white tomentose mainly with short stiff hairs  77. L. macrantha (var. heterotricha)

11b.   Young branches tomentose-pubescent or strigillose, but not spreading long stiffly hairy; hairs not longer than 2 mm.

14a.   Plant often with gray-white indumentum; reticulate veins of leaves adaxially not impressed, abaxially obviously raised and beehive-like.

15a.   Bracts sessile, not leaf-like, 2–4 mm; calyx tube glabrous; young branches (or their apices), petioles, and peduncles densely tomentose with short stiff hairs  83. L. macranthoides

15b.   Bracts stipitate, leaf-like, longer than 4 mm; calyx tube sparsely pubescent in upper half; young branches, petioles, and peduncles densely gray-white retrosely pubescent ............................................................................  84. L. buchananii

14b.   Young branches, petioles, and inflorescence with yellow-brown or gray yellow-brown indumentum; reticulate veins of leaves abaxially not raised and thus not beehive-like.

16a.   Leaves or at least old ones adaxially with reticulate veins obviously impressed and thus obviously wrinkled; corolla longer than 2.5 cm .....  85. L. rhytidophylla

16b.   Leaves adaxially with reticulate veins not obviously impressed and thus not wrinkled; corolla shorter than 2.5 cm.

17a.   Leaves abaxially with veins raised; flower pairs several to more than 10, axillary or terminal in a corymb, very rarely solitary in leaf axils; peduncle 3–15 mm; filaments and styles hairy at bases ........................................  67. L. inodora

17b.   Leaves with veins not raised; flower pairs often solitary in leaf axils or few clustered at apices of branchlets; peduncle very short or absent; filaments and styles glabrous ......................................................................  69. L. buddleioides

10b.   Leaves abaxially glabrous, stiffly hairy, or pubescent, but not velutinous, spaced among hairs.

18a.   Labia of corolla very short, ca. 1/8 as long as corolla tube ......  82. L. bournei

18b.   Labia of corolla at least 2/5 as long as corolla tube.

19a.   Calyx tube densely pubescent ..........................................  75. L. confusa

19b.   Calyx tube glabrous.

20a.   Bracts 2–3 cm, leaf-like, ovate; peduncle conspicuous; young branches densely spreading stiffly hairy ................................................  74. L. japonica

20b.   Bracts smaller, not leaf-like, if so, then peduncle very short or nearly absent.

21a.   Bracts much shorter than calyx tube, 1–2 mm, triangular.

22a.   Leaves papery; branchlets, petioles, and peduncle densely gray-white pubescent; corolla 2–3.5 mm ............................................  73. L. dasystyla

22b.   Leaves leathery, glabrous; corolla 8–12 cm.

23a.   Corolla ca. 8 cm, slender; leaves 4–6.5 cm ..  80. L. longituba

23b.   Corolla (9–)10–12 cm; leaves 7–15 cm  81. L. hildebrandiana

21b.   Bracts slightly shorter than or exceeding calyx tube.

24a.   Corolla shorter than 3 cm.

25a.   Peduncle 4–18(–23) mm; calyx lobes glabrous or only ciliate along margin; styles hairy at least below middle part ........  65. L. acuminata

25b.   Peduncle often shorter than 5 mm, sometimes nearly absent; calyx lobes hairy outside and along margin.

26a.   Styles densely hairy throughout; calyx lobes linear-lanceolate or linear    66. L. trichosepala

26b.   Styles glabrous.

27a.   Bracts much exceeding calyx lobes in length, sometimes leaf-like; peduncle very short or nearly absent; petioles 2–5 mm; leaves on both surfaces often shortly stiffly hairy only on midvein; calyx lobes subtriangular ..................................  68. L. pampaninii

27b.   Bracts nearly as long as calyx lobes; peduncle to 7 mm, petioles to 10 mm; leaves on both surfaces densely ferruginous stiffly hairy; calyx lobes linear .....................................  70. L. ferruginea

24b.   Corolla 3–14 cm.

28a.   Leaves abaxially with sessile or very shortly stipitate mushroom-shaped orange or orange-red glands; young branches densely gray-yellow or gray-white pubescent .....................................  76. L. hypoglauca

28b.   Leaves without glands, or with stipitate glandular hairs.

29a.   Young branches and abaxial surfaces of leaves ± hairy.

30a.   Young branches and abaxial surfaces of leaves densely spreading yellow stiffly hairy, hairs shorter than 2 mm  71. L. fulvotomentosa

30b.   Young branches densely pubescent, and spreading yellow-brown stiffly hairy, hairs longer than 2 mm ...........  77. L. macrantha

29b.   Plant subglabrous.

31a.   Pistils and stamens not exserted from corolla; leaves ovate-oblong to ovate-lanceolate ..............................  78. L. calvescens

31b.   Pistils and stamens exserted from corolla; leaves oblong to oblong-lanceolate ........................................  79. L. longiflora

6b.    Shrub erect, very rarely branches prostrate, never climbing; if shrub prostrate, then leaves membranous, not leathery.

32a.   Branchlets with black-brown pith, later becoming hollow with disappearance of pith.

33a.   Shrubs prostrate; leaves 1–2 cm ......................................................  62. L. prostrata

33b.   Shrubs erect; leaves longer than 2 cm.

34a.   Bracteoles ± connate, ca. 1/2 as long as or equaling calyx tube, apex ± truncate; peduncle shorter than 1 cm, very rarely exceeding petioles.

35a.   Calyx limb 5-toothed; teeth broadly triangular or lanceolate, apex acute  60. L. maackii

35b.   Calyx limb 2-sect or laciniate only on one side, with very short triangular teeth  61. L. trichosantha

34b.   Bracteoles distinct, 1/4–1/2 as long as calyx tube; peduncle usually longer than 1 cm, much exceeding petioles.

36a.   Winter buds large, ovate-lanceolate, with 5 or 6 pairs of outer scales; scales densely white ciliate along margin; calyx tube glandular, sometimes sparsely pubescent  59. L. chrysantha

36b.   Winter buds small, ovoid, often with 2 or 3(or 4) pairs of scales; scales glabrous or ciliate along margin; calyx tube glabrous.

37a.   Corolla tube ca. as long as or slightly shorter than labia ..........  56. L. tatarica

37b.   Corolla ca. 1/2 as long as labia.

38a.   Leaves oblong-obovate or ovate-oblong to oblong-lanceolate, (3–)4–6(–10) cm, apex acuminate to acute; calyx lobes ovate-triangular to triangular-lanceolate, ca. 1/2 as long as calyx tube, not ciliate ..............................................  57. L. ruprechtiana

38b.   Leaves obovate to elliptic, 2–4.5 cm, apex obtuse and mucronate, or acute; calyx lobes broadly ovate or suborbicular, ca. 1/8 as long as calyx tube, often ciliate  58. L. brevisepala

32b.   Branchlets with white, solid pith.

39a.   Corolla tube not swollen at base, longer than 5 equal or subequal corolla lobes.

40a.   Leaves often verticillate, 3 or 4 per whorl, or some opposite; calyx lobes lanceolate; corolla 5-lobed .....................................................................................................  7. L. rupicola

40b.   Leaves opposite; calyx lobes ovate to ovate-triangular or suborbicular, if lanceolate or subulate, then corolla 4-lobed.

41a.   Stigmas exserted from corolla tube.

42a.   Branchlets not spine-like at apices after leaves fall; two neighboring calyx tubes of flower pair 1/2 or nearly completely connate ..................................  6. L. tomentella

42b.   Branchlets usually spine-like at apices after leaves fall; two neighboring calyx tubes of flower pair distinct.

43a.   Anthers nearly as long as filaments; plant ca. 60 cm tall ......  9. L. spinosa

43b.   Anthers obviously shorter than filaments; plant to 1.5 m tall .  10. L. alberti

41b.   Stigmas inserted.

44a.   Two neighboring calyx tubes of flower pair distinct.

45a.   Bracteoles distinct; corolla campanulate, lobes ca. 1/2 as long as tube; leaves abaxially adpressed pubescent ..................................................  1. L. codonantha

45b.   Bracteoles connate and cupular; corolla tubular, lobes ca. 1/7 as long as tube; leaves glabrous ......................................................................  2. L. tubuliflora

44b.   Two neighboring calyx tubes of flower pair connate.

46a.   Branchlets usually spine-like at apices after leaves fall; anthers slightly exserted from corolla tube ........................................................................  8. L. minuta

46b.   Branchlets not spine-like at apices after leaves fall; anthers inserted.

47a.   Corolla outside strigillose; leaves strigillose on both surfaces  3. L. minutifolia

47b.   Corolla outside glabrous; leaves glabrous or very rarely sparsely strigillose abaxially or only on midvein.

48a.   Young branches with straight minute coarse hairs or glabrous; leaves obovate to obovate-oblong or oblanceolate, sometimes suborbicular, broadly elliptic to oblong, 0.5–2 cm; corolla 5-lobed ........................................  4. L. myrtillus

48b.   Young branches pubescent; leaves narrower and longer, ovate-oblong to oblong or linear-oblong, (1–)2–5 cm; corolla 4- or 5-lobed ..  5. L. angustifolia

39b.   Corolla tube ± swollen or obviously saccate at base.

49a.   Winter buds with only 1 pair of outer scales, if with several outer scales, then bracteoles connate and cupular, outside densely glandular hairy.

50a.   Bracteoles connate, urceolate or cupular, completely or partly surrounding two neighboring calyx tubes of flower pair.

51a.   Bracteoles connate and urceolate, glabrous, becoming fleshy and not dehiscent when fruit matures; corolla tubular-infundibuliform, regular or subregular  36. L. caerulea

51b.   Bracteoles connate, cupular or urceolate, outside hairy, not becoming fleshy, dehiscent when fruit matures; corolla labiate.

52a.   Bracteoles connate and urceolate, completely surrounding two neighboring calyx tubes of flower pair, outside densely straight stiffly hairy when young, dehiscent when fruit matures .................................................................................  37. L. ferdinandii

52b.   Bracteoles connate and cupular, not completely surrounding two neighboring calyx tubes of flower pair, outside densely glandular hairy, dehiscent before fruit matures    38. L. hypoleuca

50b.   Bracteoles not as above.

53a.   Winter buds with a pair of connate, cap-like, longitudinally wrinkled outer scales.

54a.   Styles short, ca. 1/2 as long as corolla tube.

55a.   Flowers opening at same time as leaves; flowering in June; corolla yellowish green, nearly labiate; styles minutely stiffly hairy ................  39. L. sublabiata

55b.   Flowers opening before leaves; flowering in March–April; corolla white, reddish, or rose, regular or subregular; styles glabrous.

56a.   Calyx limb shorter than calyx tube; leaves often undulate or toothed  40. L. setifera

56b.   Calyx limb ca. 2 × longer than calyx tube; leaves entire  41. L. fragilis

54b.   Styles ca. as long as or longer than corolla tube.

57a.   Stamens not exceeding corolla tube; corolla 3–4 cm; calyx limb very wide and large, 4–5 mm .............................................................  47. L. stephanocarpa

57b.   Stamens exceeding corolla tube; corolla 1.5–3 cm; calyx limb not very wide.

58a.   Corolla tube ca. 1/2 as long as corolla lobes; calyx lobes unequal in size    42. L. anisocalyx

58b.   Corolla tube longer than corolla lobes; calyx lobes equal in size or absent.

59a.   Shrubs erect, 1–2(–3) m tall; calyx limb not obviously toothed.

60a.   Young branches, petioles, abaxial surfaces of leaves, peduncle, and bracts densely minutely stiffly hairy; calyx tube glabrous; fruit blue-black  43. L. cyanocarpa

60b.   Leaves at least abaxially not minutely stiffly hairy; calyx tube often hispid and glandular hairy; fruit red .................................  44. L. hispida

59b.   Shrubs dwarf, prostrate, ca. 20 cm tall; calyx limb obviously toothed.

61a.   Styles hairy below middle part; branchlets and petioles pubescent; bracts outside pubescent .......................................  45. L. oreodoxa

61b.   Styles glabrous; branchlets and petioles minutely stiffly hairy and glandular hairy; bracts outside glabrous ......................  46. L. semenovii

53b.   Winter buds with several pairs of distinct, decussate scales, sometimes lowest pair ca. as long as bud body, covering other scales, but never longitudinally wrinkled.

62a.   Corolla subregular or slightly not regular, but not labiate; two neighboring calyx tubes of flower pair distinct; hairs without a nodular base.

63a.   Flowers opening before leaves; corolla lobes ca. 2 × longer than corolla tube; stamens exceeding corolla lobes .........................................  48. L. praeflorens

63b.   Flowers opening at same time as or after leaves; corolla tube ca. 2 × longer than corolla lobes; stamens not exceeding corolla lobes.

64a.   Flower pair and its two bracts all developed; calyx limb 1–2 mm, obtusely toothed .................................................................................  49. L. elisae

64b.   One flower of flower pair and its bract all reduced; calyx limb shorter than 0.5 mm, truncate or shallowly undulate ............................  50. L. subhispida

62b.   Corolla labiate; two neighboring calyx tubes of flower pair connate, if distinct, then plant with dense minute stiff hairs with nodular bases.

65a.   Two neighboring calyx tubes of flower pair 1/2 or completely connate.

66a.   Young branches and petioles densely minutely stiffly hairy and strigose; leaves 1–2.2 cm ..............................................................  54. L. mucronata

66b.   Young branches and petioles glabrous or sparsely strigose; leaves 3–7(–8.5) cm  55. L. fragrantissima

65b.   Two neighboring calyx tubes of flower pair distinct.

67a.   Corolla tube narrowly infundibuliform, equaling or subequaling corolla lobes in length; leaves 2–6.5 cm ..................................................  53. L. altmannii

67b.   Corolla tube narrowly cylindric, 1.5–2 × as long as corolla lobes; leaves 0.6–2 cm.

68a.   Leaves abaxially gray velutinous, often long stiffly hairy on veins  51. L. cinerea

68b.   At least old leaves abaxially glabrous or only strigillose on veins  52. L. humilis

49b.   Winter buds with several to numerous pairs of outer scales; bracteoles distinct or connate, sometimes absent, if connate and cupular, then not glandular hairy outside.

69a.   Calyx limb with cap-like emergence.

70a.   Leaves papery, 5–10(–13.5) cm, ovate-lanceolate to linear-lanceolate, apex acuminate, abaxially densely white villous on midvein .................  32. L. gynochlamydea

70b.   Leaves papery, thinly leathery, or leathery, 1–6(–15) cm, suborbicular to lanceolate or linear-lanceolate, apex obtuse or rounded, very rarely acute, abaxially not villous on midvein.

71a.   Leaves with midvein obviously raised, base usually cuneate  35. L. pileata 

71b.   Leaves with midvein impressed or at least flat adaxially, base often rounded to broadly cuneate.

72a.   Leaves papery, adaxially not nitid, margin obviously rigidly ciliate; corolla lobes ca. 1/2 as long as tube .....................................................  33. L. virgultorum

72b.   Leaves thinly leathery or leathery, adaxially nitid, margin glabrous or sparsely rigidly ciliate; corolla lobes 1/4–1/2 as long as tube ...............  34. L. ligustrina

69b.   Calyx limb without cap-like emergence.

73a.   Corolla with 5 subequal lobes; if corolla labiate, then winter buds not 4-angular, inner scales neither accrescent nor reflexed when young branches elongate.

74a.   Corolla labiate; labia nearly as long as tube.

75a.   Leaves membranous, both surfaces adpressed pubescent  15. L. microphylla

75b.   Leaves leathery, both surfaces with stiff hairs with nodular bases  16. L. oiwakensis

74b.   Corolla lobes subequal, or slightly unequal, but absolutely not labiate, shorter than tube.

76a.   Anthers with short stiff hairs .....................................  13. L. serreana

76b.   Anthers glabrous.

77a.   Bracts ovate-lanceolate; fruit black ....................  11. L. kawakamii

77b.   Bracts subulate to lanceolate or linear-lanceolate; fruit red.

78a.   Young branches, leaves, peduncle, bracts, and flowers glabrous; peduncle shorter than 1 cm .....................................................  14. L. litangensis

78b.   Young branches, leaves, peduncle, bracts, and flowers ± hairy; peduncle longer ......................................................................  12. L. tangutica

73b.   Corolla labiate; winter buds 4-angular, otherwise inner scales accrescent and reflexed when young branches elongate.

79a.   Winter buds not 4-angular, inner scales accrescent and reflexed when young branches elongate.

80a.   Two neighboring calyx tubes of flower pair distinct.

81a.   Bracteoles 2 pairs, distinct; winter buds with more than 3 pairs of outer scales.

82a.   Leaves sparsely or densely stiffly hairy and sparsely glandular on both surfaces; petioles 3–6(–8) mm; winter buds with 5 pairs of outer scales  17. L. webbiana

82b.   Leaves subglabrous; petioles 5–12 mm; winter buds with 3 pairs of outer scales .................................................................  18. L. heterophylla

81b.   Bracteoles connate and cupular; winter buds with 2 pairs of outer scales.

83a.   Branchlets of current year, petioles, and peduncle densely white pubescent; leaves oblong, obovate-oblong, or ovate-oblong, 2.5–5.5 × 1–2.4 cm; petioles 3–6 mm ..............................................................  21. L. jilongensis

83b.   Branchlets of current year, petioles, and peduncle shortly glandular hairy; leaves triangular-ovate to rhombic-ovate, 2.5–5.3 × 2.5–5.3; petioles 1.5–2.5 cm        22. L. oblata

80b.   Two neighboring calyx tubes of flower pair ca. 1/2 or completely connate.

84a.   Bracteoles distinct; bracts leaf-like, ovate-lanceolate to ovate-oblong, 6–17 mm; young branches, petioles, and peduncles white stiffly hairy and glandular hairy; peduncle 3–4(–5) cm .............................................  19. L. fargesii

84b.   Bracteoles connate and cupular, with 4 shallow, orbicular lobes; bracts not leaf-like, subulate, 3–5 mm; young branches, petioles, and peduncles sparsely glandular hairy, glabrescent; peduncle (5–)8–23 mm ..................  20. L. hemsleyana

79b.   Winter buds 4-angular, inner scales not obviously accrescent when branchlets elongate.

85a.   Peduncle (or fruit stalk) usually ca. as long as or slightly longer than petioles; fruit red or black.

86a.   Leaves powdery-white, often strigillose, 1–2.5 cm; fruit black; young branches, petioles, and peduncles glabrous ................................  25. L. retusa

86b.   Leaves abaxially not powdery-white, pubescent, 2–8.5 cm; fruit red.

87a.   Calyx tube glandular in upper part; corolla tube ca. as long as or slightly shorter than labia; young branches, petioles, and peduncle densely pubescent  23. L. modesta

87b.   Calyx tube not glandular; corolla tube short, ca. 1/2 as long as labia; young branches, petioles, and peduncle sparsely pubescent or glabrous  24. L. graebneri

85b.   Peduncle (or fruit stalk) often obviously longer than petioles; fruit black.

88a.   Two neighboring calyx tubes of flower pair distinct or ca. 1/2 connate.

89a.   Leaves glabrous; flowers white, later turning yellow  28. L. nervosa

89b.   Leaves pubescent on both surfaces or abaxially white barbate only on midvein; flowers purplish or purple-red.

90a.   Leaves abaxially white barbate only on midvein; peduncle (1.5–)2–3 cm     26. L. nigra

90b.   Leaves sparsely pubescent on both surfaces; peduncle usually shorter than 1.5 cm ...........................................................  27. L. lanceolata

88b.   Two neighboring calyx tubes of flower pair more than 1/2 connate.

91a.   Calyx lobes lanceolate, equaling or slightly exceeding tube  29. L. kansuensis

91b.   Calyx lobes broadly triangular to triangular-lanceolate, shorter than tube.

92a.   Leaves abaxially initially gray-white velutinous, glabrescent, margin not ciliate; calyx lobes narrow and long, triangular-lanceolate  30. L. tatarinowii

92b.   Leaves abaxially sparsely strigose or subglabrous, margin ciliate; calyx lobes short, broadly triangular .........................  31. L. maximowiczii

 
1. Lonicera codonantha Rehder, J. Arnold Arbor. 22: 578. 1941.

钟花忍冬  zhong hua ren dong

Shrubs, deciduous, to ca. 1 m tall. Young branches reddish brown, together with petioles densely adpressed yellowish brown puberulous to subglabrous; old branches gray-brown. Winter buds with ca. 4 pairs of scales. Leaves very shortly petiolate; leaf blade dark green adaxially, broadly ovate-elliptic or ovate-elliptic, 1–2 × 0.6–1.2 cm, papery, abaxially densely adpressed puberulous, midvein impressed [abaxially or adaxially??], sparsely puberulous [midvein or adaxially??], base rounded or subcuneate, margin revolute, apex rounded or obtuse and mucronate. Flowers at bases of branchlets of current year; peduncle ca. 2 mm, glabrous; bracts ovate-lanceolate or obovate, equaling or slightly exceeding calyx tube, abaxially puberulous, sparsely ciliate; bracteoles separate, suborbicular, ca. 1/3 as long as calyx tube, glabrous. Neighboring 2 calyx tubes separate, subglobose, ca. 1 mm; limb shorter than calyx tube; calyx lobes ovate, glabrous, apex slightly obtuse. Corolla white, tinged with pink, campanulate, outside glabrous; tube 6–7 mm, inside puberulous above middle; lobes ca. 1/2 as long as tube. Stamens inserted in lower 1/3 of corolla tube, slightly exceeding middle of corolla in length, glabrous. Styles shorter than stamens, glabrous; ovary 3-loculed, with numerous ovules per locule. Fruit unknown. Fl. Jul–Aug.

* Sunny rocky slopes; ca. 3500 m. SW Yunnan (Zhenkang).

2. Lonicera tubuliflora Rehder in Sargent, Pl. Wilson. 1: 129. 1911.

管花忍冬 guan hua ren dong

Shrubs deciduous, to 2 m tall, subglabrous throughout except young branches minutely adpressed puberulous and minutely glandular hairy and young leaves sparsely glandular ciliolate. Petiole shorter than 1 mm; leaf blade blue-green adaxially, oblong or oblong-ovate, 5–12(–17) × 2–3 mm, papery, abaxially glaucous, base broadly cuneate to rounded, apex obtuse. Flowers fragrant, axillary on upper or middle part of young branches; peduncle erect, short, equaling or slightly exceeding petioles; bracts leaf-like, stipitate, linear-oblong, to 5 mm, 2–3 × as long as calyx tube; bracteoles cupular, apex often truncate, 1/2–2/3 as long as calyx tube. Neighboring 2 calyx tubes separate, elliptic, ca. 1.5 mm; calyx lobes ovate or triangular-ovate, unequal in shape, ca. 1/2 as long as calyx tube, glabrous or sometimes sparsely glandular ciliolate, apex obtuse. Corolla white or yellowish, tubular; tube 8–10 mm, 1.5–2.5 mm in diam., margin of mouth slightly constricted, inside villous throughout; lobes spreading, orbicular-ovate, ca. 1/7 as long as tube; anthers slightly above middle of corolla tube. Stigmas below anthers. Ovary 2-loculed. Fruit red or red-brown, ovoid or globose, ca. 4 mm in diam. Fl. Jun–Jul, fr. Sep.

* Scrub; 2100–3100 m. NW Sichuan.

3. Lonicera minutifolia Kitamura, Acta Phytotax. Geobot. 15: 133. 1954.

细叶忍冬  xi ye ren dong

Subshrubs, deciduous, richly branched, to 50 cm tall, both surfaces of leaves, bracts, bracteoles, and margin of calyx lobes orange-red ciliolate. Branchlets of current year purple-brown  or gray-brown, with spreading short coarse hairs; old branches gray-black, divaricate-spreading; outer scales of winter buds ovoid, ca. 1 mm, dorsally carinate, apex obtuse. Leaves clustered at apices of young branchlets; petiole to 1 mm; leaf blade ovate-elliptic, elliptic, or oblong, 5–9 × 3–6 mm, both surfaces densely strigillose and with scattered orange-red small glandular hairs, base obtuse, margin with short coarse hairs, apex rounded or obtuse. Peduncle very short; bracts leaf-like, oblanceolate, 1.2–1.5 mm, stipitate, with same indumentum as leaves; bracteoles cupular, shorter than 1 mm, apex irregularly truncate, ciliolate. Neighboring 2 calyx tubes separate, ca. 1.2 mm; limb shorter than tube; lobes ovate-triangular, very short, margin ciliolate, apex obtuse. Corolla white, tubular, 5–7 mm; tube 3–5 mm, 2–3 mm in diam., outside densely strigillose, inside densely puberulous; lobes suborbicular, subequal, 1.5–2 mm. Stamens inserted in upper part of corolla tube; filaments very short; anthers up to throat of corolla. Styles ca. 2/5 as long as corolla tube. Fruit unknown. Fl. May–Aug.

[??*] Scrub; 3700–4400 m. Xizang [??Nepal].

4. Lonicera myrtillus J. D. Hooker & Thomson, J. Linn. Soc., Bot. 2: 168. 1858.

越桔叶忍冬 yue ju ye  ren dong

Subshrubs deciduous, richly branched, to 3 m tall; leaves, petioles, and bracts (sometimes bracteoles and calyx lobes) often sparsely red-brown glandular ciliolate. Young branches gray-brown or tinged with red-purple, with straight minute coarse hairs or glabrous, sometimes mixed with minute glandular hairs; old branches erect or prostrate, gray-brown or gray-black; winter buds with several pairs of acute scales. Petiole 1–2 mm, glabrous or very rarely with minute coarse hairs; leaf blade highly variable in shape, those of short branches often obovate to obovate-oblong or oblanceolate, sometimes oblong to broadly elliptic or ovate, 0.5–2 cm × 3–8 mm, papery or thickly so, glabrous, very rarely with scattered short coarse hairs abaxially or only on midvein, often with very small rusty punctuate dots abaxially on old leaves, base cuneate, apex obtuse to rounded and often mucronate or cuspidate. Peduncles axillary on lateral short branches, 1–15 mm, glabrous or minutely coarse hairy; bracts leaf-like, often exceeding calyx lobes in length; bracteoles cupular, ca. 1/2 as long as or equaling calyx tube, apex truncate or shallowly dentate, sometimes 2-lobed. Neighboring 2 calyx tubes united above middle or throughout; limb shallowly cupular; lobes triangular or ovate-triangular, often unequal, apex obtuse or sometimes slightly acute. Corolla white, purplish, or purple-red, tubular-campanulate, 6–8 mm, outside glabrous; tube inside puberulous, more densely hairy on throat; lobes orbicular-ovate or suborbicular, 1/4 or 1/2 as long as tube. Stamens and styles included; anthers ca. 1 mm, up to above corolla tube and slightly below throat. Styles shorter; stigmas up to middle of corolla tube or 1/3 of upper part. Fruit purple-red, suborbicular, 4–6 mm in diam.; seeds brownish, ovoid to oblong, compressed, 2–3 mm. Fl. May–Jun(–Jul), fr. Aug–Sep.

Scrub, sparse forests, stony places of river valleys; 2400–4000(–4700) m. SW Sichuan, Xizang, Yunnan [Afghanistan, ??Bhutan, ??Kashmir, N Myanmar].

1a.. Leaves and bracts narrower, obovate or ovate to oblanceolate or lanceolate ......  4a. var. myrtillus

1b.. Leaves and bracts wider, suborbicular or orbicular-ovate to broadly elliptic ...  4b. var. cyclophylla

 

4a. Lonicera myrtillus var. myrtillus

越桔叶忍冬 (原变种) yue ju ye ren dong (yuan bian zhong)

Caprifolium parvifolium Kuntze; Lonicera angustifolia Wallich ex de Candolle var. rhododactyla W. W. Smith; L. myrtillus J. D. Hooker & Thomson var. depressa Rehder.

Leaves and bracts narrower, obovate or ovate to oblanceolate or lanceolate.

Scrub, sparse forests, rocky places; 2400–4000(–4700) m. SW Sichuan, Xizang, Yunnan [Afghanistan, N Myanmar].

4b. Lonicera myrtillus  var. cyclophylla Rehder, J. Arnold Arbor. 22: 579. 1941.

圆叶忍冬yuan ye ren dong

Leaves and bracts wider, suborbicular or orbicular-ovate to broadly elliptic.

* Rocky places; 3000–4000 m. SE Xizang, NW Yunnan.

5. Lonicera angustifolia Wallich ex de Candolle, Prodr. 4: 337. 1830.

狭叶忍冬  xia ye ren dong

Caprifolium angustifolium (Wallich ex de Candolle) Kuntze.

Shrubs, deciduous, 1–3 m tall. Young branches brownish, pubescent or subglabrous. Leaves shortly petiolate; leaf blade ovate-oblong to oblong or linear-oblong, (1–)2–5 × 0.6–1.3 cm, both surfaces glabrous, or abaxially often sparsely pubescent or only hairy on midvein, base cuneate, apex obtuse or acuminate. Peduncle axillary, 0.5–3 cm; bracts linear to linear-lanceolate or oblanceolate, 6–10 mm, often exceeding calyx tube; bracteoles connate and cupular, apex obtuse or sinuous. Neighboring 2 calyx tubes connate nearly throughout; lobes ovate-triangular to narrowly triangular. Corolla white or purplish, tubular-campanulate, outside glabrous, 4- or 5-lobed; lobes ovate-orbicular, 1/5–1/4 as long as tube. Stamens and stigmas included. Fruit red, globose. Fl. May–Jul, fr. Aug–Sep. 2n = 18.

Forests, scrub; 2700–4500 m. S Xizang [??Bhutan, N India, Kashmir, Nepal].

6. Lonicera tomentella J. D. Hooker & Thomson, J. Linn. Soc., Bot. 2: 167. 1858.

毛冠忍冬  mao guan ren dong

Shrubs, deciduous, to 4 m tall. Branchlets of current year, petioles, and peduncle densely yellowish brown tomentose; branchlets of current year brownish, branchlets of previous year purple-brown or gray-brown. Petiole longer than 2 mm; leaf blade ovate-oblong to ovate-lanceolate, 1–2 × 0.6–1.5 cm, abaxially gray-green or glaucous, pubescent or glabrous, more densely so on midvein, adaxially sparsely strigillose, lateral veins 4, 5, 7, or 8 pairs, together with midvein with scattered orange-red minute glands, base cuneate to slightly cordate, margin revolute, apex obtuse. Peduncle very short, less than 2 mm, to 4 mm at fruiting stage; bracts large, leaf-like, ovate to linear-oblong, to 7 mm; bracteoles cupular, ca. 1/2 as long as calyx tube, glabrous. Neighboring 2 calyx tubes united to more than half, ca. 2.5 mm, glabrous; limb ca. 1 mm; calyx lobes ovate, apex obtuse. Corolla purplish red, tubular-infundibuliform, outside strigillose; tube ca. 8 mm, inside strigillose; lobes subequal, orbicular-ovate, ca. 2.5 mm. Stamens inserted in upper part of corolla tube, unequal in length. Stigmas large, up to base of corolla lobes. Fruit blue-black, globose, ca. 6 mm in diam. Fl. Jun–Aug, fr. Sep.

Forests, scrub; 2000–3200 m. Xizang, NW Yunnan [??Bhutan, NE India (Sikkim), ??Nepal].

1a.. Leaves gray-green, densely pubescent; lateral veins 7 or 8 pairs ....................  6a. var. tomentella

1b.. Leaves glaucous, sparsely pubescent or glabrous; lateral veins 4 or 5 pairs .  6b. var. tsarongensis

 

6a. Lonicera tomentella var. tomentella

毛冠忍冬 (原变种) mao guan ren dong (yuan bian zhong)

Caprifolium tomentellum (J. D. Hooker & Thomson) Kuntze; Lonicera tomentella J. D. Hooker & Thomson var. conaensis P. S. Hsu & Y. F. Huang.

Leaf blade gray-green abaxially, densely pubescent, lateral veins 7 or 8 pairs.

Forests, scrub; 2900–3000 m. Xizang, NW Yunnan [NE India (Sikkim)].

6b. Lonicera tomentella var. tsarongensis W. W. Smith, Notes Roy. Bot. Gard. Edinburgh 8: 168. 1921.

察瓦龙忍冬cha wa long ren dong

Leaf blade often glaucous abaxially, sparsely pubescent or glabrous, lateral veins 4 or 5 pairs.

* Forests, scrub; 2000–3200 m. SE Xizang, NW Yunnan.

7. Lonicera rupicola J. D. Hooker & Thomson, J. Linn. Soc., Bot. 2: 168. 1858.

岩生忍冬  yan sheng ren dong

Shrubs, deciduous, to 1.5(–2.5) m tall, sometimes only 10–20 cm tall at high elevations. Young branchlets and petioles white pubescent and minutely glandular hairy, or subglabrous; branchlets slender, often needle-like when leaves deciduous, sometimes elongated and prostrate. Leaves 3(or 4)-verticillate, rarely opposite; petiole to 3 mm; leaf blade linear-lanceolate or oblong-lanceolate to oblong, 0.5–3.7 cm, papery, abaxially white velutinous, sparsely pubescent, or glabrous, adaxially glabrous or minutely glandular hairy, base cuneate to rounded or subtruncate, inequilateral, margin revolute, apex acute, slightly mucronate, or obtuse. Flowers axillary at bases of young branches, fragrant; peduncle very short; bracts, bracteoles, and margins of calyx lobes minutely puberulous and with minute glands; bracts leaf-like, linear-lanceolate to linear-oblanceolate, slightly exceeding calyx lobes; bracteoles cupular, apex truncate or 4-lobed or divided to middle, sometimes completely separate, ca. 1/2 as long as or equaling calyx tube. Neighboring 2 calyx tubes separate, ca. 2 mm, glabrous; lobes narrowly lanceolate, 2.5–3 mm, exceeding tube. Corolla purplish or purple-red, tubular-campanulate, (8–)10–15 mm, outside often puberulous and minutely glandular hairy; tube 1.5–2 × as long as lobes, inside puberulous, particularly so in upper part; lobes spreading, ovate, 3–4 mm, 1/5–1/2 as long as tube. Anthers up to middle of corolla tube. Styles up to half of corolla tube, glabrous. Fruit red, ellipsoid, ca. 8 mm; seeds brownish, oblong, compressed, ca. 4 mm. Fl. May–Aug, fr. Aug–Oct.

[??*] Alpine scrub and meadows, margins of scree slopes, grasslands at forest margins; 2000–5000 m. Gansu, S Ningxia, Qinghai, Sichuan, Xizang, NW Yunnan [??Bhutan, ??Nepal].

1a.. Leaves abaxially usually white velutinous ........................................................  7a. var. rupicola

1b.. Leaves abaxially glabrous or sparsely pubescent ........................................  7b. var. syringantha

 

7a. Lonicera rupicola var. rupicola

岩生忍冬 (原变种 )  yan sheng ren dong (yuan bian zhong)

Caprifolium rupicolum (J. D. Hooker & Thomson) Kuntze; Lonicera rupicola J. D. Hooker & Thomson subsp. thibetica (Bureau & Franchet) Y. C. Tang; L. rupicola var. thibetica (Bureau & Franchet) Zabel; L. thibetica Bureau & Franchet.

Leaves abaxially usually white velutinous.

* Alpine scrub and meadows, margins of scree slopes, grasslands at forest margins; 2100–5000 m. Gansu, S Ningxia, Qinghai, Sichuan, Xizang, NW Yunnan.

7b. Lonicera rupicola var. syringantha (Maximowicz) Zabel in Beiss. et al., Handl. Laubh–Ben. 462. 1903.

红花岩生忍冬hong hua yan sheng  ren dong

Lonicera syringantha Maximowicz, Bull. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 3, 24: 49. 1878; L. rupicola subsp. syringantha (Maximowicz) Y. C. Tang; L. syringantha var. minor Maximowicz; L. syringantha subsp. syringantha (Maximowicz) Y. C. Tang, nom. invalid??; L. syringantha var. wolfii Rehder; L. wolfii (Rehder) K. S. Hao.

Leaves abaxially glabrous or sparsely pubescent.

* Scrub, forest margins; 2000–4600 m. Gansu, S Ningxia, E Qinghai, Sichuan, Xizang, NW Yunnan.

8. Lonicera minuta Batalin, Trudy Imp. S.-Peterburgsk. Bot. Sada 12: 170. 1892.

矮生忍冬 ai sheng ren dong

Shrubs, deciduous, richly branched, dwarf, to 30 cm tall; branchlets of current year mostly withered in winter; young branches, both surfaces of leaves or at least adaxial surface, petioles, and bracts coarsely hairy. Branchlets yellowish brown, needle-like when leaves deciduous, old branches gray-brown. Leaves opposite; petiole ca. 1 mm; leaf blade linear or linear-oblanceolate, those of short branches often wider and linear-oblong to ovate-oblong, 5–12 × 2–3 mm, midvein raised abaxially, conspicuously impressed adaxially, base cuneate or rounded to subtruncate, margin ± revolute, apex obtuse. Flowers in lower part of branchlets of current year, fragrant; peduncle nearly absent; bracts leaf-like, linear-lanceolate to linear, subequaling calyx lobes; bracteoles cupular, 2-lobed, equaling or slightly exceeding calyx lobes, apex subtruncate, together with calyx lobes ciliate. Neighboring 2 calyx tubes separate, 1.5–2 mm; limb shallowly cupular, subequaling tube; lobes ovate-triangular or narrowly ovate, apex obtuse. Corolla purplish red, tubular-infundibuliform, ca. 1.4 mm; tube ca. 1 cm, inside puberulous, and so at middle and lower parts of lobes; lobes subovate, ± unequal, 3.5–4 mm. Filaments inserted at mouth of corolla tube, very short; anthers ca. 2 mm, slightly exceeding corolla tube. Styles included, ca. 7 mm. Fruit ovoid or subglobose, ca. 7 mm. Fl. May–Jun, fr. Aug–Sep.

* Rocky crevices, sand dunes; 3200–3800 m. W Gansu, Qinghai.

9. Lonicera spinosa Jacquemont ex Walpers, Repert. Bot. Syst. 2: 449. 1843.

棘枝忍冬 ji zhi ren dong

Caprifolium spinosum (Jacquemont ex Walpers) Kuntze; Xylosteum spinosum (Jacquemont ex Walpers) Decaisne.

Shrubs, deciduous, dwarf, to 60 cm tall, often with spine-like, leafless branchlets. Branchlets of current year coarsely hairy; winter buds with several pairs of scales. Leaves opposite; petiole very short, slightly coarse hairy along margin; leaf blade linear to linear-oblong, those of short branches often wider and oblong or obovate, 4–12 × 1–2 mm, glabrous throughout but slightly coarsely hairy along margin, base broadly cuneate to rounded, margin revolute, apex obtuse. Flowers axillary on short branches; peduncle very short; bracts leaf-like, linear to linear-oblong, often exceeding calyx lobes; bracteoles cupular, more than 1/2 as long as calyx tube, apex subtruncate, often 2-lobed. Neighboring 2 calyx tubes separate; limb cupular, ca. 1.5 mm; lobes orbicular-ovate, apex obtuse. Corolla purplish red, later white, tubular-infundibuliform; tube ca. 9 mm, slender; lobes ovate-oblong, ca. 4 mm. Filaments slightly below mouth of corolla tube; anthers exserted from corolla tube, equaling filaments. Styles exserted. Fruit ellipsoid, ca. 5 mm. Fl. Jun–Jul, fr. Sep.

Scrub, scree slopes, often mixed with Caragana plants; 3700–4600 m. Xizang [??Afghanistan, ??India (Sikkim), Kashmir].

10. Lonicera alberti Regel, Trudy Imp. S.-Peterburgsk. Bot. Sada 7: 550. 1881.

沼生忍冬  zhao sheng ren dong

Chamaecerasus alberti (Regel) Carrière; Lonicera spinosa Jacquemont ex Walpers var. albertii (Regel) Rehder.

Shrubs, deciduous, 1–1.5 m tall, glabrous. Branchlets prostrate, slender. Petiole short; leaf blade powdery-green or blue-green, linear-oblong, to 3 cm, margin entire or often basally 1- or 2-dentate, apex slightly obtuse. Peduncle axillary, shorter than flowers. Corolla slightly rose, ca. 1.6 cm. Stamens subequaling corolla; anthers conspicuously shorter than filaments. Fl. Jun, fr. Aug.

Scrub in wetlands; ca. 1700 m. N Xinjiang [Kazakhstan, Kyrgyzstan, Tajikistan].

[No specimens were seen. The description must be rewritten based on specimens].

11. Lonicera kawakamii (Hayata) Masamune, J. Soc. Trop. Agric. 3: 246. 1931.

玉山忍冬  yu shan ren dong

Coprosma kawakamii Hayata, J. Coll. Sci. Imp. Univ. Tokyo 31(1) (Mat. Fl. Formos.): 145. 1911.

Subshrubs, deciduous, to 2 m tall. Branchlets glabrous, covered with remains of petiole bases. Petiole 1–2 mm; leaf blade obovate, very small, 6–16 × 3–5 mm, papery, abaxially sparsely strigillose or subglabrous on veins, adaxially glabrous, midvein and lateral veins slightly raised abaxially, impressed adaxially, lateral veins ca. 4 pairs, base cuneate, margin entire, apex rounded or obtuse. Flowers opening after leaves; peduncle axillary, slightly arched, 2–5 mm, slender, glabrous; bracts ovate-lanceolate, 1–2 mm, exceeding calyx tube; bracteoles orbicular-ovate, ca. 1/4 as long as calyx tube. Neighboring 2 calyx tubes united to more than 2/3; limb very short, undulate-crenate or subtruncate. Corolla yellowish, tubular-infundibuliform, ca. 4 mm, both surfaces glabrous; lobes subregular, ovate, ca. 1/3 as long as tube. Stamens inserted at middle and lower parts of corolla tube; anthers up to throat of corolla. Ovary 2-loculed, ovule 1 per locule; styles slightly exceeding corolla tube, puberulous. Fruit black, globose, ca. 3 mm in diam. Fl. ??, fr. ??.

* Mountains; ca. 1300 m. Taiwan.

No specimens were seen.

12. Lonicera tangutica Maximowicz, Bull. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 3, 24: 48. 1878.

唐古特忍冬  tang gu te ren dong

Caprifolium tanguticum (Maximowicz) Kuntze; Lonicera aemulans Rehder; L. chlamydata W. W. Smith; L. chlamydophora W. W. Smith (1913), not K. Koch (1851); L. cylindriflora Handel-Mazzetti; L. fangii S. S. Chien; L. flavipes Rehder; L. glandulifera S. S. Chien; L. guebriantiana Handel-Mazzetti; L. inconspicua Batalin; L. kungeana K. S. Hao; L. longa Rehder; L. penduliflora Pax & K. Hoffmann; L. saccata Rehder; L. saccata f. calva Rehder; L. saccata var. tangiana (S. S. Chien) P. S. Hsu & H. J. Wang; L. saccata f. wilsonii Rehder; L. schneideriana Rehder; L. serpyllifolia Rehder; L. shensiensis (Rehder) Rehder; L. szechuanica Batalin; L. taipeiensis P. S. Hsu & H. J. Wang; L. tangiana S. S. Chien; L. tangutica var. glabra Batalin; ??L. trichopoda Franchet; L. trichopoda Franchet var. shensiensis Rehder; L. wulingensis Nakai.

Shrubs, deciduous, to 2(–4) m tall. Young branches pubescent or glabrous, sometimes mixed with short glandular hairs; branchlets of previous year brownish; winter buds acuminate or acute; scales 2–4 pairs, ovate or ovate-lanceolate, with short stiff hairs, ciliate, or glabrous, apex acuminate or acute. Petiole 2–3 mm; leaf blade oblanceolate to oblong or obovate to elliptic, 1–4(–8) × 0.5–2 cm, papery, both surfaces pubescent, sometimes subglabrous or glabrous, base attenuate, apex obtuse or ± acute. Peduncle axillary on lower part of young branches, slightly nodding, 1–3(–4.5) cm, slender, pubescent or glabrous; bracts narrow, sometimes leaf-like, slightly shorter than or slightly exceeding calyx lobes; bracteoles separate or united, 1/5–1/4 as long as calyx tube, ciliate or not. Neighboring 2 calyx tubes united to above middle or completely united, elliptic or oblong, 2(–4) mm, glabrous or pubescent; limb cupular, 2/5–1/2 as long as or equaling tube, apex dentate or undulate to truncate, sometimes ciliate. Corolla white, yellow-white, or tinged reddish, tubular-infundibuliform, (8–)10–13 mm, swollen or shallowly gibbous slightly on one side at base of tube, outside glabrous or sometimes with sparse stiff hairs; lobes suberect, orbicular-ovate, 2–3 mm. Stamens inserted at middle of corolla tube. Styles exceeding corolla lobes, glabrous or with sparse spreading stiff hairs on middle and lower parts. Fruit red, 5–6 mm in diam.; seeds brownish, ovoid or oblong, 2–2.5 mm. Fl. May–Jun, fr. Jul–Sep.

[??*] Forests, grasslands on mountain slopes, scrub at stream sides; 1600–3500(–3900) m. Anhui, Gansu, Guizhou, Hebei, Henan, W Hubei, W Hunan, Ningxia, E Qinghai, Shaanxi, Shanxi, Sichuan, Xizang, NW Yunnan [??Myanmar].

This is the most highly variable species in Chinese Lonicera. Further studies are needed for a satisfactory taxonomic treatment.

13. Lonicera serreana Handel-Mazzetti, Oesterr. Bot. Zeit., Heft. 4, 83: 234. 1934.

毛药忍冬 mao yao ren dong

Lonicera hopeiensis S. S. Chien.

Shrubs, deciduous, to 3(–4) m tall, pubescent throughout except on calyx tube. Branchlets of current year purple-brown, pubescent or glabrous; old branches gray; winter buds acuminate; scales 2 or 3 pairs, lanceolate, apex acuminate or ± acute, ciliate. Petiole very short, 1–3 mm, pubescent; leaf blade obovate to oblanceolate, oblong, or elliptic, (1–)1.5–3.5(–4.5) cm, papery, both surfaces pubescent, base cuneate, apex obtuse or ± acute. Peduncle solitary, axillary on lower part of young branches, slightly nodding, 5–20 mm, pubescent or glabrous; bracts ovate-lanceolate or ovate, subequaling or slightly exceeding calyx tube, ciliate, outside pubescent on midvein; bracteoles 2 or absent, orbicular-ovate to obovate or suborbicular, 1/3–2/3 as long as calyx tube, outside sparsely pubescent or glabrous, apex rounded, obtuse, or acute, ciliate. Neighboring 2 calyx tubes 1/2 to completely united, 2–2.5 mm, glabrous; limb cupular, ½–3/5 as long as tube; lobes small, irregularly undulate, sparsely pubescent or ciliate. Corolla yellow-white, pinkish, or purple, tubular or tubular-infundibuliform, 1–1.3 cm, swollen or shallowly gibbous slightly on one side at base; lobes erect, ovate, 1.5–2.5 mm, outside and along margin sometimes pubescent. Anthers equaling or slightly exceeding corolla lobes, with short stiff hairs. Styles exserted, with sparse stiff hairs throughout or only on middle and lower parts. Fruit red, orbicular, 5–6 mm in diam., with 4 longitudinal ridges. Fl. Jun–Aug, fr. Aug–Sep.

* Scrub, forests; 800–2800 m. Gansu, Hebei, Henan, Ningxia, Shaanxi, Shanxi, Sichuan.

14. Lonicera litangensis Batalin, Trudy Imp. S.-Peterburgsk. Bot. Sada 14: 173. 1895.

理塘忍冬  li tang ren dong

Lonicera farreri W. W. Smith; L. oresbia W. W. Smith; L. rockii Rehder.

Shrubs, deciduous, dwarf, to 1–2 m tall, richly branched, glabrous throughout. Branchlets of previous year brown, internodes extremely shortened, covered with remains of petiole bases; winter buds obtuse at apex; scales 2–4 pairs, apex obtuse or ± acute. Petiole short; leaf blade elliptic or broadly elliptic to obovate, 6–12 × 5–9 mm, papery, base broadly cuneate, apex obtuse or mucronulate. Flowers opening with leaves simultaneously, often axillary on short branches; peduncle very short or nearly absent; bracts leaf-like, ovate to narrowly ovate-lanceolate, large, 5–8 mm, exceeding calyx tube. Neighboring 2 calyx tubes completely united and globose, 2–3 mm; lobes broadly triangular or undulate, apex obtuse. Corolla yellow or yellowish, tubular or narrowly infundibuliform, 1–1.3 cm, outside glabrous, shallowly gibbous at base on one side of tube; lobes erect, orbicular-ovate, ca. 2 mm. Anthers subequaling or slightly exceeding corolla lobes. Ovary 3-loculed, with numerous ovules per locule; styles included or slightly exserted, puberulous on lower part. Fruit red, becoming gray-blue, globose, ca. 8 mm in diam.; seeds ellipsoid or oblong, 4–5 mm. Fl. May–Jun, fr. Aug–Sep.

[??*] Scrub, grasslands, forests, forest margins; 3000–4500 m. Sichuan, Xizang, NW Yunnan [??Bhutan, ??Nepal].

15. Lonicera microphylla Willdenow ex Roemer & Schultes, Syst. Veg. 5: 258. 1819.

小叶忍冬  xiao ye ren dong

Caprifolium microphyllum (Willdenow ex Roemer & Schultes) Kuntze; Xylosteum sieversianum Ruprecht.

Shrubs, deciduous, to 2(–3) m tall. Young branches glabrous or sparsely pubescent; old branches gray-black. Petiole very short; leaf blade adaxially often gray-white, obovate or obovate-elliptic to elliptic or oblong, sometimes oblanceolate, 5–22 × 5–13 mm, papery, both surfaces densely or sparsely adpressed puberulous or sometimes subglabrous, often with scaly glands in vein axils on lower part, base cuneate, [margin??] pubescent-ciliate, apex obtuse or ± acute, sometimes rounded to obtuse and mucronulate. Peduncles in pair, axillary on lower part of young branches, slightly curved or nodding, 5–12 mm; bracts subulate, slightly exceeding calyx limb or to 2 × as long as calyx tube. Neighboring 2 calyx tubes ± completely united, glabrous; limb shallow, short, undulate; lobes inconspicuous. Corolla yellow or white, 7–10(–14) mm, outside with sparse short stiff hairs or glabrous, bilabiate; labia subequaling corolla tube, gibbous at base on one side; lobes of upper labium erect, oblong; lower labium curved. Stamens inserted at base of labium, together with styles slightly exserted; filaments with very sparse, short, stiff hairs. Styles with dense or sparse stiff hairs. Fruit red or orange-yellow, orbicular, 5–6 mm in diam.; seeds yellowish brown, oblong or ovoid-ellipsoid, 2.5–3 mm, smooth. Fl. May–Jun(–Jul), fr. Jul–Aug(–Sep). 2n = 18.

Rocky slopes, grasslands, thickets, sparse forests, forest margins; 1100–3600(–4100) m. Gansu, W Hebei, ??Henan, Nei Mongol, Ningxia, Qinghai, Shanxi, Xinjiang, Xizang [Afghanistan, NW India, Mongolia, Russia].

16. Lonicera oiwakensis Hayata, Icon. Pl. Formosan. 6: 24. 1916.

瘤基忍冬  liu ji ren dong

Shrubs, to 2 m tall. Branchlets and both surfaces of leaves with stiff hairs with nodular bases; branchlets spreading, gray. Petiole 2–3 mm, with stiff hairs; leaf blade elliptic or oblong, sometimes oblanceolate, ca. 2.5 × 1.5 cm, leathery, base rounded or cuneate, margin with short stiff hairs, apex slightly obtuse or rounded. Peduncle on lower part of branchlets of current year, nodding, ca. 1 cm, slender. Bracts linear-lanceolate, ca. 2 × as long as calyx tube, abaxially on midvein and adaxially with short stiff hairs, apex acuminate. Neighboring 2 calyx tubes united to above middle part, obconical, ca. 2 mm, glabrous; limb shallowly cupular, ca. 1 mm, irregularly 5-lobed; lobes obtuse at apex. Corolla white, apex tinged with rose, bilabiate; tube infundibuliform, ca. 7 mm, ca. 2 mm in diam., gibbous at base on one side, outside and inside with short stiff hairs; upper labium reflexed, ca. 7 × 7 mm, both surfaces glabrous, lobes oblong; lower labium ligulate, ca. 8 × 4 mm. Anthers slightly exserted from corolla, ca. 2.5 mm. Ovary 3-loculed, with 4 or 5 ovules per locule; styles ca. 1 cm, glabrous. Fruit unknown. Fl. Apr.

* Forests; 3000–3900 m. Taiwan.

17. Lonicera webbiana Wallich ex de Candolle, Prodr. 4: 336. 1830.

华西忍冬 hua xi ren dong

Shrubs, deciduous, to 3(–4) m tall. Branches often glabrous or with dispersed red glands; winter buds with ca. 5 pairs of outer scales, apex acute, inner scales enlarged and reflexed. Petiole 3–6(–8) mm; leaf blade ovate-elliptic to ovate-lanceolate, 4–9(–18) × 2–8 cm, papery, ciliate, with dense or sparse stiff hairs and sparse glands, base rounded or slightly cordate or broadly cuneate, margin often irregularly undulate or crenulate, apex acuminate or long acuminate. Peduncle 2.5–5(–9) cm; bracts linear, (1–)2–5 mm; bracteoles separate, ovate to oblong, very small, less than 1 mm. Neighboring 2 calyx tubes separate, glabrous or glandular hairy; lobes small, apex obtuse, undulate, or acute. Corolla purple-red, very rarely white or turning from white to yellow, 1–1.5 cm, bilabiate, outside sparsely pubescent and glandular hairy or glabrous; tube very short, base slender, shallowly gibbous, abruptly enlarged upward; upper labium erect, crenate; lower labium ca. 1/3 longer than upper labium, curved. Stamens subequaling corolla; filaments and styles puberulous on lower part. Fruit red, later black, globose, ca. 1 cm in diam.; seeds ellipsoid, 5–6 mm, with small concave dots. Fl. May–Jun, fr. Aug–Sep.

Needle-leaved and broad-leaved mixed forests, scrub, grassy slopes; 1800–4000 m. S Gansu, ??Henan, W Hubei, Jiangxi, S Ningxia, E Qinghai, S Shaanxi, Shanxi, Sichuan, Xizang, NW Yunnan [Afghanistan, Bhutan, Kashmir, ??Myanmar; Europe].

1a.. Peduncle shorter than 5 cm; corolla ca. 1 cm ...............................................  17a. var. webbiana

1b.. Peduncle 5–9 cm; corolla 1.2–1.5 cm ........................................................  17b. var. mupinensis

 

17a. Lonicera webbiana var. webbiana

华西忍冬 (原变种) hua xi ren dong (yuan bian zhong)

Lonicera adenophora Franchet; L. alpigena Linnaeus var. phaeantha Rehder; ??L. heteroloba Batalin; L. tatsienensis Franchet; L. webbiana Wallich ex de Candolle var. lanpinensis Y. C. Tang.

Peduncle shorter than 5 cm. Corolla ca. 1 cm.

Needle-leaved and broad-leaved mixed forests, scrub, grassy slopes; 1800–4000 m. S Gansu, W Hubei, Jiangxi, S Ningxia, E Qinghai, S Shaanxi, Shanxi, Sichuan, Xizang, NW Yunnan [Afghanistan, Bhutan, Kashmir; Europe].

17b. Lonicera webbiana var. mupinensis (Rehder) P. S. Hsu & H. J. Wang, Acta Phytotax. Sin. 17(4): 76. 1979.

川西忍冬  chuan xi ren dong

Lonicera mupinensis Rehder in Sargent, Pl. Wilson. 1: 138. 1911.

Peduncle 5–9 cm. Corolla 1.2–1.5 cm.

* Forests, alpine scrub; 2300–3400 m. W Sichuan.

18. Lonicera heterophylla Decaisne in Jacquemont, Voy. Inde 4: 80. 1844.

异叶忍冬 yi ye ren dong

Caprifolium karelinii Kuntze; Lonicera heterophylla var. karelini (Bunge ex Kirilov) Rehder; L.