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FOC | Family List | FOC Vol. 19 | Caprifoliaceae

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

忍冬属 ren dong shu

Lonicera tangutica

Credit: Harvard University Herbaria

Caprifolium Miller; Euchylia Dulac; Xylosteon Miller.

Shrubs erect or dwarf, rarely small trees, sometimes climbers, deciduous or evergreen. Branches hollow or solid with white or brown pith; winter buds with 1 to several pairs of scales, rounded or acutely 4-angular, inner scales sometimes accrescent and reflexed. Accessory buds sometimes present, occasionally terminal buds reduced and substituted by 2 lateral buds. Leaves opposite, rarely whorled, margin entire, rarely dentate or divided; leaves usually estipulate, occasionally with interpetiolar stipules or a swollen interpetiolar line; sometimes 1 or 2 pairs of leaves below inflorescence connate and forming involucral bracts. Inflorescence thyrsoid, terminal or axillary, cymes opposite and usually reduced to paired flowers, rarely 1-, sometimes 3-flowered. Inflorescence occasionally pedunculate; cymes sessile, sometimes forming a capitulum, or cymes pedunculate with a pair of bracts and 2 pairs of bracteoles; bracts usually small, sometimes leaflike; bracteoles usually free, sometimes ± fused and cupular occasionally enclosing ovaries, sometimes absent. Paired flowers with free or partially to completely fused ovaries. Calyx 5-lobed, rarely 4-lobed, sometimes truncate, base occasionally with a collarlike emergence. Corolla white, yellow, reddish, or purple-red, often changing color after anthesis, campanulate, funnelform, regularly or subregularly 5(or 4)-lobed, or bilabiate and upper lip 4-lobed; tube long or short, often shallowly to deeply gibbous on ventral side toward base, rarely spurred. Nectary of compact sessile glandular hairs on ventral side toward base of corolla tube, occasionally in 5 regular lines, rarely swollen at base of style. Stamens 5; anthers dorsifixed. Ovary 2 or 3(-5)-locular; style slender, hairy or glabrous; stigmas capitate. Fruit a berry, red, blue-black, black, green, or white sometimes pruinose, bracteoles occasionally accrescent in fruit and enclosing paired berries. Seeds 1 to numerous, smooth, pitted or granular, with rounded embryo.

About 180 species: N Africa, Asia, Europe, North America; 57 species (23 endemic) in China.

See Rehder, Rep. (Annual) Missouri Bot. Gard. 14: 27-232, t. 1-20. 1903.

The identities of Lonicera confusa 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. Aug. 1st Agric. Coll. 18(2): 7. 1995), and L. subrotundata C. Y. Yang & J. H. Fan (J. Aug. 1st Agric. Coll. 18(2): 8. 1995) cannot be determined because of the unavailability of the type specimens for examination or the original publications. One North American species, L. sempervirens Linnaeus, is occasionally cultivated.

Species Complexes

6-8. Lonicera tangutica species complex

Shrubs, deciduous, to 1-4 m tall. Branches with solid pith. Winter buds with 2-4 pairs of scales. Petiole 1-5 mm; leaf blade obovate to lanceolate, 0.5-8.5 × 0.2-2.5 cm, both sur­faces glabrous or pubescent, base cuneate, apex obtuse to acute. Inflorescences axillary, paired flowers; peduncle usually nod­ding, 0-3 cm, slender, glabrous, rarely pubescent; bracts nar­row, sometimes leaflike, usually exceeding or rarely shorter than ovaries; bracteoles sometimes present, separate or fused, minute to 1/4 as long as ovaries, often ciliate. Neighboring 2 ovaries fused completely or at least to middle, elliptic or ob­long, ca. 2 mm, glabrous or occasionally pubescent. Calyx cu­pular, to 2 mm, truncate or with ovate to triangular lobes, gla­brous to pubescent, sometimes ciliate. Corolla white, yellow, or pink, sometimes tinged purple, tubular-funnelform, 8-13 mm; tube glabrous, occasionally with sparse stiff hairs, nectary swol­len or gibbous on ventral side at base of corolla tube, sometimes forming a spur shorter than ovaries; lobes suberect, orbicular-ovate, 1-2 mm; mouth glabrous or with sparse spreading stiff hairs. Stamens inserted at middle of corolla tube; anthers par­tially exserted, glabrous or ciliate. Style exserted by 1-3 mm. Berries orange, red, purple, or black, 5-6 mm in diam.; seeds brownish, ovoid or oblong, 2-2.5 mm, smooth. Fl. May-Aug, fr. Jul-Sep.

Forests, forest margins, grasslands, scrub, mountains; 800-4500 m. Anhui, Gansu, Guizhou, Hebei, Henan, W Hubei, W Hunan, Ning­xia, E Qinghai, Shaanxi, Shanxi, Sichuan, Taiwan, Xizang, NW Yunnan [Bhutan, India (Sikkim), Nepal]. This is a highly variable species complex; further studies are needed for a satisfactory taxonomic treatment.

19-23. Lonicera hispida species complex

Shrubs, deciduous, to 3 m tall, occasionally dwarf. Winter buds with a pair of narrow keeled outer scales, covering inner scales and sometimes longitudinally sulcate. Young branches often pruinose, minutely scabrid to stiffly hairy and glandular hairy, very rarely glabrous. Petiole 1.5-6 mm; leaf blade usually ovate to oblong, sometimes elliptic, 0.6-9 × 0.4-3.5 cm, sub­glabrous, with short stiff hairs or adpressed villous, base cune­ate to slightly cordate, margin hirsute-ciliate, apex acute to ob­tuse and mucronulate. Inflorescences axillary paired flowers, and at base of new shoots; peduncle stout, sometimes slightly compressed, 0.5-2 cm; bracts broadly ovate, 1-4 cm; bracteoles absent. Paired ovaries free, glabrous to hirsute, often glandular hairy. Calyx cupular, 1-4 mm, to 5 mm at fruiting, truncate or lobed; lobes broad and hirsute-ciliate. Corolla yellow-green or dark purple, funnelform, subregular, 13-35 mm; tube shallowly to deeply gibbous toward base, glabrous to puberulent outside and inside at upper part; lobes erect, ovate, 5-10 mm. Stamens and style usually subequaling or longer than corolla, occa­sionally shorter. Style stiffly hairy on lower half. Berries red, black-brown, or blue-black and pruinose, ovoid, 1-1.5 cm, 3-8-seeded; seeds deep brown, irregularly triangular-oblong, 4-5 mm, smooth. Fl. May-Jul, fr. Jul-Sep. 2n = 18*.

Forests, scrub, alpine grasslands, rocky places, slopes, high moun­tains; 1700-4800 m. Gansu, Hebei, ?Henan, Ningxia, Qinghai, Shaanxi, Shanxi, Sichuan, Xinjiang, Xizang, Yunnan [Afghanistan, Bhutan, India (Sikkim), Kashmir, Kazakhstan, Kyrgyzstan, Mongolia, Nepal, Paki­stan; SW Asia (Iran)]. Lonicera sublabiata P. S. Hsu & H. J. Wang (Acta Phytotax. Sin. 17(4): 78. 1979), based on a type specimen collected in Lixian, Sichuan, by Z. He (12715) and kept in the Shanghai Museum of Natural History (SHM), could not be located; it has a short hispid style, a bilabiate corolla (though the 5 lobes are all located on the same side so it must be the result of a deliberate split of the corolla), and it seems to flower with the emerging leaves. Nevertheless, it does not seem to differ signifi­cantly from L. hispida species complex.

27-28. Lonicera alpigena species complex

Shrubs, deciduous, to 4 m tall. Winter buds with 2-5 pairs of outer scales; inner scales accrescent, enlarged and sometimes reflexed. Branches glabrous to densely hairy with mixed glan­dular hairs. Petiole 3-20 mm; leaf blade obovate or ovate-ellip­tic to ovate-lanceolate, 1.5-17 × 0.8-6 cm, with dense or sparse stiff hairs and sparse glands or glabrous, base cuneate to sub­cordate, margin ciliate, apex apiculate to long acuminate. Inflo­rescences axillary, paired flowers; peduncle 0.5-9 cm, usually thickened toward apex; bracts linear-subulate, to 10 mm, or sometimes minute or leaflike and ovate-lanceolate, to 15 mm; bracteoles separate or sometimes fused into a cupule, ovate to oblong, to 1 mm, ciliate. Neighboring 2 ovaries free to fused, glabrous or densely glandular hairy. Calyx lobes ovate to orbic­ular, to 1 mm, sometimes forming a collarlike emergence at base, apex irregularly lobed, sometimes glandular ciliate. Co­rolla bilabiate, purple-red, very rarely white or turning from white to yellow, 1-1.5 cm, outside sparsely pubescent and glan­dular hairy or glabrous; tube 4-7 mm, inside puberulent, deeply gibbous above slender base; upper lip erect, 4-lobed to 1/4; lower lip recurved. Stamens subequaling corolla; filaments and style hirsute on lower part or glabrous. Berries red or black, glo­bose, ca. 1 cm in diam.; seeds ellipsoid, 5-6 mm, shallowly pitted to smooth. Fl. Apr-Jun, fr. Jun-Oct.

Needle-leaved and broad-leaved mixed forests, scrub, grassy slopes; 900-4000 m. Anhui, S Gansu, W Henan, Hubei, Jiangxi, S Ningxia, E Qinghai, S Shaanxi, Shanxi, Sichuan, Xinjiang, Xizang, NW Yunnan, Zhejiang [Afghanistan, Bhutan, India, Kashmir, ?Myanmar, Russia; Europe]. Lonicera alpigena Linnaeus occurs in Europe; further studies are necessary to better understand the relationships within this species com­plex in Europe and Asia. The type specimen of Lonicera jilongensis P. S. Hsu & H. J. Wang (Acta Phytotax. Sin. 17(4): 76. 1979) has suffered extensive insect dam­age, but it seems to belong here.

31-33. Lonicera nigra species complex

Shrubs, deciduous, to 4 m tall. Winter buds with several pairs of outer acute, keeled scales; inner scales sometimes accrescent and enlarged. Branches with white or sometimes brown pith. Young branches and peduncles often puberulent and glandular hairy. Petiole 2-10 mm; leaf blade oblong to elliptic-lanceolate, 1.5-10 × 1-3 cm, glabrous throughout but abaxially often white hairy on midvein, base cuneate to rounded, apex acute to acuminate. Inflorescences axillary, paired flowers; peduncle 0.5-3 cm; bracts lanceolate-linear, ca. 2 mm; bracte­oles free, fused into 2 pairs or into an entire or lobed cupule, 1/3 to almost as long as ovaries, glandular ciliate. Neighboring 2 ovaries free, fused in lower half or completely fused. Calyx lobes triangular or linear-lanceolate, 1-2 mm, sometimes with a collar emergence at base, glandular ciliate. Corolla bilabiate, purplish, purple-red, or white becoming yellow, 9-13 mm; tube ca. 5 mm, shallowly gibbous at base, outside puberulent or gla­brous, inside puberulent; upper lip crenulate; lower lip reflexed. Stamens and style exserted from corolla tube; filaments gla­brous or hairy at base. Style hairy below middle part or throughout. Berries black, often pruinose, globose, 5-7 mm in diam.; seeds oblong or ovoid, 3-7 mm, granular-raised and coarse. Fl. Apr-Jul, fr. Aug-Oct. 2n = 18.

Forests, scrub, coniferous forests, forest margins, grasslands; 1500-4000 m. W Anhui, Gansu, NE Guizhou, W Henan, W Hubei, Jilin, Ningxia, Qinghai, Shaanxi, S Shanxi, Sichuan, Xizang, Yunnan [Bhutan, India, Korea, Nepal; Europe]. Lonicera govaniana Wallich ex Candolle (Prodr. 4: 337. 1830) and L. orientalis Lamarck (Encycl. 1: 731. 1785) belong to L. caucasica Pal­las (Fl. Ross. 1(1): 57. 1784). Further studies are necessary to determine if L. caucasica, predominantly from Asia Minor and S Asia, differs from L. nigra in Europe and China.

34-36. Lonicera maximowiczii species complex

Shrubs, deciduous, to 2 m tall. Winter buds with several pairs of outer acute, keeled scales; inner scales sometimes ac­crescent and enlarged. Branches sparsely puberulent to glabres­cent. Petiole 2-8 mm; leaf blade ovate or broadly ovate to ob­long-lanceolate, 4-12 × 1.5-3.5 cm, abaxially sparsely strigose, minutely tomentose, or glabrous, adaxially sparsely strigose or glabrous, base cuneate to rounded, margin often ciliate, apex acute to acuminate. Inflorescences axillary, paired flowers; pe­duncle 1-2.5 cm, glabrous or sparsely hairy; bracts subulate, 1-3 mm; bracteoles fused into a cupule, 1/3 to as long as ovaries. Neighboring 2 ovaries more than half to completely fused, very rarely completely free. Calyx lobes triangular to linear, 1-1.5 mm. Corolla bilabiate, dark purple to purple-red or white be­coming yellow, ca. 1 cm, outside glabrous; tube slightly gib­bous toward base, ca. 4 mm, inside densely hairy; upper lip 4 lobed, lobes 1-2 mm; lower lip recurved. Stamens and style exserted from corolla tube; filaments hairy at base. Style hairy throughout. Berries red, ovoid-orbicular; seeds yellowish brown, oblong, 4-5 mm, granular and coarse. Fl. May-Jul, fr. Aug-Sep.

Forests, forest margins, scrub; 400-2400 m. S Gansu, Hebei, Hei­longjiang, ?Henan, Jilin, Liaoning, ?Nei Mongol, Ningxia, Shaanxi, Shandong, N Sichuan [Japan, Korea, Russia].

47-51. Lonicera macrantha species complex

Climbers, semievergreen to deciduous. Branches solid, often becoming hollow. Branches, petioles, and peduncles with ± dense spreading, yellow-brown long stiff hairs and minute glandular hairs, grayish pubescent, yellow-brown velutinous, or sometimes glabrous. Petiole 3-15 mm; leaf blade ovate or ob­long to lanceolate, 2-14 × 1-5 cm, abaxially hirsute mixed with short glandular hairs, pubescent, shortly white velutinous, glau­cous with large sessile orange glands, or occasionally subgla­brous, veins reticulate and raised, adaxially hirsute-hairy on veins and midvein, veins conspicuously impressed, sometimes wrinkled, base rounded to subcordate, occasionally broadly cu­neate, margin ciliate and revolute, apex acute to acuminate, sometimes obtuse and mucronate. Flowers fragrant, paired and in racemes toward apex of branchlets, often paniculate with leaflike to subulate involucral bracts; peduncle 1-15 mm, some­times to 40 mm in lower part of inflorescence; peduncle, bracts, bracteoles, and calyx lobes stiffly hairy and glandular hairy; bracts lanceolate to linear, 2-7 mm, occasionally leaflike to 14 mm; bracteoles orbicular-ovate, ca. 1 mm. Neighboring 2 ovaries free, ca. 2 mm, glabrous or sometimes sparsely to densely hirsute or with sessile glands. Calyx lobes lanceolate, 1-2 mm, ciliate. Corolla bilabiate, white, later yellow, (1.5-)3-7 cm, outside with spreading hairs and minute glandular hairs, glabrous, or strigose; tube (1-)2-5 cm, inside densely puberu­lent, not gibbous at base; upper lip 4-lobed, lobes ca. 5 mm; lower lip recurved. Stamens and style slightly exceeding co­rolla, glabrous or sometimes pubescent at base. Berries white or black, sometimes pruinose, 1 of paired ovaries sometimes aborting, globose or ellipsoid, 7-12 mm; seeds few, ovoid, compressed, 5-7 mm, shallowly pitted and furrowed. Fl. Mar-Jul, fr. Jul-Dec.

Forests of mountain valleys or slopes, scrub, riversides, stream­sides, roadsides; 200-2900 m. S Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hainan, Hubei, Hunan, Jiangxi, Shaanxi, Shanxi, Si­chuan, Taiwan, Xizang, Yunnan, Zhejiang [Bhutan, India, Japan, ?Kash­mir, Myanmar, Nepal, N Vietnam]. These species have long been used medicinally in China.

1 Flowers in 3-flowered cymes forming whorls at ends of branches   (2)
+ Flowers paired (occasionally single), axillary, paniculate or forming thyrses at ends of branches; involucral bracts not connate   (5)
2 (1) Involucral bracts not connate; cymes pedunculate.   14 L. hypoleuca
+ Involucral bracts usually connate; cymes sessile   (3)
3 (2) Corolla funnelform, lobes regular.   55 L. subaequalis
+ Corolla bilabiate   (4)
4 (3) Corolla 1.5-2.5 cm; bracts lanceolate.   56 L. yunnanensis
+ Corolla 5-9 cm; bracts ovate.   57 L. tragophylla
5 (1) Shrubs; corolla actinomorphic, tube not gibbous and nectary of compact sessile glandular hairs in 5 regular lines; branches with solid pith; accessory buds absent   (6)
+ Shrubs or climbers; corolla zygomorphic (sometimes only at base), tube gibbous at base or not and nectary of compact sessile glandular hairs on ventral side toward base of corolla tube; branches solid or hollow; accessory buds often present   (10)
6 (5) Style as long as or longer than corolla tube   (7)
+ Style ca. 1/2 length of corolla tube   (8)
7 (6) Anthers and stigmas exserted; branches often with leafless spinelike apex; berries pale violet to white; dwarf shrubs 30-60 cm tall.   5 L. spinosa
+ Anthers and stigmas not exserted; branches not spinelike at apex; berries blue-black with glaucous bloom; shrubs 200-400 cm tall.   3 L. tomentella
8 (6) Leaves sometimes whorled; flowers at bases of branches; branches often becoming spinelike at apex; stems sympodial; ovary 3-locular.   4 L. rupicola
+ Leaves not whorled; flowers on short lateral branches or along monopodial stems; branches not spinelike at apex; ovary 2-locular   (9)
9 (8) Ovaries fused above middle or throughout; corolla tubular-campanulate, lobes usually more than 1.5 mm; flowers 4- or 5-merous.   2 L. angustifolia
+ Ovaries free; corolla narrowly tubular, lobes usually ca. 1.5 mm; flowers 5-merous.   1 L. tubuliflora
10 (5) Climbers, sometimes sprawling   (11)
+ Upright shrubs, sometimes compact or prostrate with upright shoots   (20)
11 (10) Corolla with a ca. 12 mm spur toward base, neighboring 2 ovaries connate.   42 L. calcarata
+ Corolla not spurred, sometimes shallowly gibbous, neighboring 2 ovaries free   (12)
12 (11) Corolla less than 3 cm   (13)
+ Corolla more than 3 cm   (15)
13 (12) Corolla tube broadly funnelform, shallowly gibbous toward base.   45 L. acuminata
+ Corolla tube narrowly funnelform, not gibbous   (14)
14 (13) Corolla densely yellow-brown strigose; leaves abaxially densely yellow-brown strigose, margins conspicuously ciliate.   46 L. ferruginea
+ Corolla with spreading or adpressed hairs and minute glandular hairs, glabrous, or strigose; leaves abaxially not densely yellow-brown strigose, margins not conspicuously ciliate.  

L. macrantha species complex

15 (12) Bracts leaflike   (16)
+ Bracts not leaflike, linear or subulate   (17)
16 (15) Bracts ovate to elliptic, 10-30 mm; leaves abaxially sparsely to densely hairy but not obscuring leaf surface.   44 L. japonica
+ Bracts linear-lanceolate, 5-14 mm; leaves abaxially white velutinous with indumentum obscuring leaf surface.  

L. macrantha species complex

17 (15) Corolla 9-12 cm, tube broadly funnelform; berries green when ripe.   53 L. hildebrandiana
+ Corolla 3-9 cm, tube narrowly funnelform; berries white, red, or black when ripe   (18)
18 (17) Corolla 5-9 cm; leaves glabrous; berries white or black.   52 L. longiflora
+ Corolla 3-7 cm; leaves abaxially hirsute mixed with short glandular hairs, pubescent, shortly white velutinous or glaucous with large sessile orange glands, occasionally subglabrous; berries white, red, or black   (19)
19 (18) Corolla 3-4.5 cm, lip less than 1/5 as long as tube; berries red.   54 L. bournei
+ Corolla 3-7 cm, lip more than 1/4 as long as tube; berries white or black.  

L. macrantha species complex

20 (10) Branches with pith becoming brown and later hollow   (21)
+ Branches with solid pith   (25)
21 (20) Peduncles 10 mm or less   (22)
+ Peduncle 10 mm or more   (23)
22 (21) Calyx scarious and campanulate, splitting; corolla 1.2-1.5 cm; winter bud scales hairy inside.   41 L. trichosantha
+ Calyx not scarious, tubular at base but lobed at apex; corolla ca. 2 cm; winter bud scales glabrous inside.   40 L. maackii
23 (21) Inner winter bud scales densely hairy and ciliate outside, ovaries ± glandular hairy.   39 L. chrysantha
+ Inner winter bud scales glabrous to sparsely hairy outside, ovaries glabrous   (24)
24 (23) Upper corolla lip 4-lobed, 2 lateral lobes reaching base and spreading, tube shallowly gibbous toward base; leaf margin ciliate.   37 L. tatarica
+ Upper corolla lip 4-lobed, 2 lateral lobes reaching 1/2-3/4 and slightly spreading, tube deeply gibbous toward base; leaf margin sometimes dentate.   38 L. ruprechtiana
25 (20) Bracteoles fused into a cupule enclosing ovaries at anthesis (note that this can give appearance of fused ovaries and absent bracteoles); bracteoles sometimes accrescent and enclosing paired berries in fruit   (26)
+ Bracteoles absent or shorter than ovaries, free, fused in pairs, sometimes fused into a cupule but not enclosing ovaries   (30)
26 (25) Calyx forming a collarlike emergence at base; bracteoles not accrescent in fruit   (27)
+ Calyx not forming a collarlike emergence at base; bracteoles accrescent in fruit sometimes enclosing paired berries   (28)
27 (26) Corolla bilabiate; style stiffly hairy to apex; leaves usually with tufted hairs at base of midvein abaxially.   11 L. gynochlamydea
+ Corolla lobes regular; style villous at base only; leaves usually without tufted hairs at base of midvein abaxially.   12 L. ligustrina
28 (26) Inflorescence not terminal, paired flowers axillary; corolla lobes regular; cupule glabrous, tightly enclosing paired ovaries and forming a compound fruit.   10 L. caerulea
+ Inflorescence terminal; corolla bilabiate; cupule pubescent, loosely enclosing paired berries, berries often exserted from cupule in fruit   (29)
29 (28) Cupule usually glandular pubescent, accrescent to 1/2 length of berries; inflorescence terminal, of 3-8 whorls; winter buds glabrous inside; seeds shallowly pitted; leaf apex obtuse or rounded.   14 L. hypoleuca
+ Cupule villous, enclosing paired berries, sometimes splitting at maturity; inflorescence terminal, of 1-3 whorls; winter buds densely villous inside; seeds deeply pitted with orange glands; leaf apex acuminate or acute.   13 L. ferdinandi
30 (25) Winter bud acutely 4-angular   (31)
+ Winter bud not acutely 4-angular   (34)
31 (31) Ovaries free or fused only at base; bracteoles free or fused into a cupule; berries black.  

31-33. L. nigra species complex

+ Ovaries half to completely fused, bracteoles fused into a cupule; berries red or black   (32)
32 (31) Peduncle to 7 mm; berries red.   30 L. modesta
+ Peduncles 5-30 mm; berries red or black   (33)
33 (33) Berries red; leaf blade ovate, 4-12 × 1.5-3.5 cm.  

L. maximowiczii species complex

+ Berries black; leaf blade usually obovate, 1-2.5 × 0.7-1.8 cm, abaxially often pruinose, apex retuse to obtuse.  

L. nigra species complex

34 (30) Ovaries connate to ca. 1/2 or more   (35)
+ Ovaries free   (39)
35 (35) Corolla with regular erect lobes; stamens inserted or partially exserted 6-8. L. tangutica species complex   (35)
+ Corolla bilabiate, lobes recurved; stamens fully exserted   (36)
36 (36) Bracteoles present, free or sometimes fused into a cupule; inner winter bud scales long accrescent; peduncle 0.5-9 cm, usually thickened toward apex.  

L. alpigena species complex

+ Bracteoles absent; inner winter bud scales long accrescent or not; peduncle 0.5-1.5 cm, not thickened toward apex   (37)
37 (36) Flowering Apr-Jun; inner winter bud scales not accrescent; berry globose.   9 L. microphylla
+ Flowering Jan-Apr; inner winter bud scales long accrescent; berry oblong   (38)
38 (37) Deciduous shrubs; leaves ovate to lanceolate, acute to acuminate, 3-8.5 cm.   15 L. fragrantissima
+ Evergreen shrubs; leaves broadly obovate, obtuse and mucronate, 1-2 cm.   16 L. mucronata
39 (34) Corolla hypocrateriform, densely hairy at mouth; style ca. 1/2 length of corolla tube; flowers appearing before leaves.   18 L. setifera
+ Corolla not hypocrateriform; style subequaling or longer than corolla tube; flowers appearing before or after leaves   (40)
40 (39) Corolla bilabiate   (41)
+ Corolla funnelform, lobes subregular   (44)
41 (40) Bracteoles absent; bracts leaflike, ovate to lanceolate.   26 L. humilis
+ Bracteoles present, free or fused in pairs or into a cupule; bracts linear-subulate   (42)
42 (41) Prostrate shrubs with evergreen leathery leaves; corolla tube longer than lobes.   43 L. crassifolia
+ Erect shrubs with deciduous leaves; corolla tube shorter than lobes   (43)
43 (43) Bracteoles free or fused in pairs; peduncle 0.5-9 cm, usually thickened toward apex; leaves ovate-elliptic to ovate-lanceolate, longer than wide, apex apiculate to long acuminate.  

L. alpigena species complex

+ Bracteoles fused into a cupule shorter than ovaries; peduncle 0.7-1 cm, not thickened toward apex; leaves broadly rhombic, as long as wide, apex cuspidate and mucronate or obtuse.   29 L. oblata
44 (40) Cymes reduced to a single flower, sometimes with a second partially developed.   25 L. subhispida
+ Cymes of paired flowers   (45)
45 (45) Inner winter bud scales not accrescent and covered by a single outer pair of keeled scales; bracts usually 1 cm or more.  

L. hispida species complex

+ Inner winter bud scales accrescent; bracts usually 1 cm or less   (46)
46 (45) Corolla lobes more than half as long as tube; anthers usually deep pink, exserted; berry globose.   17 L. praeflorens
+ Corolla lobes less than half as long as tube; berry ellipsoid.   24 L. elisae

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