LINNAEACEAE (CAPRIFOLIACEAE Subfam. Linnaeoideae) [Draft]

忍冬科  add pinyin name

Qin-er Yang (杨亲二); Sven Landrein; Peter F. Stevens

Shrubs erect or creeping. Leaves opposite, sometimes whorled, simple, estipulate, with an interpetiolar line. Inflorescence a cyme (or a reduced cyme of 1 or 2 flowers). Flowers bisexual, ± zygomorphic. Perianth 4- or 5-merous. Corolla imbricate in bud; nectary consisting of ± compact glandular hairs, inside corolla tube. Stamens 4, adnate to corolla tube (adnate for 1/3–1/2 with corolla), alternating with corolla lobes. Ovary inferior, 3- or 4-loculed, placentation axile; 1 or 2 locules with a single fertile ovule and 1 or 2 locules with numerous sterile ovules; style long; stigma capitate. Fruit an achene crowned with persistent sepals (deciduous in Linnaea). Seeds 1 or 2; embryo small, straight; endosperm copious.

Six genera and ca. 16 species: mostly in China and Japan; five genera and 12 species in China (??#endemic).

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

1a.  Petioles of opposite leaf pairs dilated and connate at base, enclosing axillary buds; corolla hypocrateriform, with a cylindrical tube and 4 or 5 spreading lobes; cymes 2- or 3-flowered; flowers fragrant ..  1. Zabelia

1b.  Petioles of opposite leaf pairs not dilated, axillary buds exposed; corolla infundibuliform or bilabiate with 5 spreading lobes; cymes 1- or 2-flowered; flowers not fragrant except in Abelia chinensis.

2a.  Ovary 4-locular with two fertile locules; 2 bracts accrescent and becoming wing-like in fruit  4. Dipelta

2b.  Ovary 3-locular with one fertile locule; bracts not wing-like in fruit.

3a.  Two bracts shield-like and covered with sticky glandular hairs; flowers  paired at apices of branchlets        5. Linnaea

3b. Bracts not as above.

4a.     Bracts and bracteoles covered with bristles, accrescent and becoming hard and covering base of ovaries in fruit ....................................................................................  3. Kolkwitzia

4b.     Bracts and bracteoles without bristles, non-accrescent in fruit ......................  2. Abelia

 

1. ZABELIA (Rehder) Makino, Makinoa. 9: 175. 1948; cf. Hisauchi & Hara, J. Jap. Bot. 29: 143. 1954.

Abelia sect. Zabelia Rehder. Pl. Wilson 1(1): 125. 1911

Shrubs deciduous. Vigorous reiteration shoots often with 6 longitudinal grooves and retrorse stiff hairs. Leaves opposite, margin entire or dentate (or sometimes lobed on reiteration shoots), shortly petiolate, estipulate. Petioles of opposite leaf pairs dilated and connate at base, enclosing axillary buds. Inflorescence a terminal cyme often clustered and forming a thyrse; cymes usually with 8 bracts at base of ovaries (only 6 bracts in Zabelia sect. Biflorae). Calyx of 4 or 5 sepals, persistent, spreading, narrowly oblong to elliptic. Corolla white, pale rose, or sometimes reddish, hypocrateriform and ± zygomorphic, 4- or 5-lobed; corolla tube usually without distinct swelling at base, glandular inside. Stamens included, didynamous, inserted at base or middle of corolla tube; anthers yellow, introrse. Ovary usually 3-locular, 2 locules with 2 series of sterile ovules and 1 locule with a single fertile ovule; style filiform; stigma green, capitate, viscous. Fruit a leathery achene, oblong, crowned with persistent calyx lobes; seed subterete, testa membranous; endosperm fleshy.

Four species: Afghanistan, China, Japan, NW India, Korea, Kyrgyzstan, Nepal, Russia (Far East); two species in China.

This group was first published by Rehder (in Sargent, Pl. Wilson. 1: 118. 1911) as Abelia sect Zabelia. It was described in honor of Zabel who was the first to delimit the genus Abelia into sections (Mitt. Deutsch. Dendrol. Ges. volume??: 33. 1893). The phylogeny of this group was further studied by Makino and supported by the studies of Ikuse and Kurosawa ‘Notes on Sect. Zabelia Rehder of the genus Abelia’ (J. Jap. Bot. 29(4): 11. 1954), and later by Fukuoka ‘Phylogeny of the Tribe Linnaeeae’ (Acta Phytotax. Geobot. 23: 82. 1968). The genus Zabelia was then segregated from Abelia based on pollen, wood anatomy, inflorescence structure, and karyology (Makino, Makinoa. 9: 175. 1948, invalid publication; Hisauchi and Hara in J. Jap. Bot. 29: 143. 1954).

1a.     Calyx and corolla lobes 5; inflorescence a terminal or subterminal cyme with 1 terminal and 2 axillary flowers (sometimes more due to supernumerary flowers axillary to bracteoles); cymes often clustered in a thyrse; bracts and bracteoles leaf-like at base of short pedicels; sepals long ciliate in fruit ............  1. Z.triflora

1b.     Calyx and corolla lobes 4; inflorescence a reduced terminal cyme consisting of paired axillary flowers (occasionaly more due to supernumerary flowers axillary to bracteoles); bracts and bracteoles reduced and located at base of ovaries; sepals not long ciliate in fruit (occasionally sparsely ciliate)  2. Z. biflora

 

Zabelia sect. Zabelia Fukuoka, Acta. Phytotax. Geobot. 43: page?? 1968.

1. Zabelia triflora (R.Brown ex. Wallich) Makino Makinoa 9: 175. 1948; cf. Hisauchi & Hara in J. Jap. Bot. 29: 144. 1954.

醉鱼草状六道木  zui yu cao zhuang liu dao mu

Abelia triflora R. Brown. ex Wallich, Pl. Asiat. Rar. 1(1): 14, t. 15. 1829; A. angustifolia Bureau ex Franchet; A. buddleioides W. W. Smith; A. buddleioides var. divergens W. W. Smith; A. buddleioides var. intercedens Handel-Mazzetti; A. buddleioides var. stenantha Handel-Mazzetti; Zabelia buddleioides (W. W. Smith) Hisauchi & H. Hara; Z. buddleioides var. divergens (W. W. Smith) Golubkova; Z. buddleioides var. stenantha (Handel-Mazzetti) Hisauchi & H. Hara; Z. stenantha (Handel-Mazzetti) Golubkova.

Shrubs deciduous, 1–2 m tall. Young branchlets with retrorse stiff hairs, becoming gray and glabrous. Petiole very short, to 2 mm, hispid. Leaf blade ovate to lanceolate, 15–70 × 5–20 mm, both surfaces glabrous but long hispid on margin and veins abaxially, base cuneate, margin entire to occasionally serrate or lobed, apex acute. Inflorescence a terminal cyme often clustered and forming a thyrse; cyme with 8 bracts at base of ovaries. Pedicels short or nearly absent. Bracts leaf-like, lanceolate to obovate; bracteoles linear to subulate, ca. 4 mm, hispid. Ovary narrowly ovoid, hirsute. Calyx of 5 linear sepals, 4–10 × to 1 mm, ciliate with stiff hairs, midvein conspicuously raised. Corolla white, sometimes tinged red, hypocrateriform, 10–20 mm, nearly 2 × as long as sepals; lobes 5, spreading, suborbicular; tube densely villous inside, with sparsely adpressed hairs outside. Stamens 4, didynamous, included; filaments short, hispid; anthers oblong. Styles filiform, exceeding stamens; stigma capitate. Achene terete, striate, crowned with 5 slightly enlarged and long ciliate persistent sepals. Fl. May, fr. Jun–Aug.

Forests, scrub, grasslands; 1800–3500 m. SW Sichuan, SE Xizang, NW Yunnan [Afghanistan, NW India, Nepal, Pakistan].

The Chinese specimens were previously named as Abelia buddleioides by W. W. Smith (Notes Roy. Bot. Gard. Edinburgh 9: 75. 1915–16). The type specimen Forrest 12636 from Yunnan is similar to the type of Abelia triflora var. parvifolia (Clarke) Hisauchi & H. Hara from Pakistan. Nevertheless, the group shows much variation in the size and shape of the leaves and sepals.

Zabelia sect. Biflorae (Rehder) Fukuoka, Acta. Phytotax. Geobot. 43: page?? 1968.

Abelia Gruppe biflorae Zabel.

2. Zabelia biflora (Turczaninow) Makino, Makinoa 9: 175. 1948; cf. Hisauchi & Hara, J. Jap. Bot. 29: 143. 1954.

六道木  liu dao mu

Abelia biflora Turczaninow, Bjull. Moskovsk. Obsc. Isp. Prir. Otd. Biol. 10: 152. 1837; A. anhweiensis Nakai; A. biflora var. coreana (Nakai) C. F. Fang; A. biflora f. minor (Nakai) C. F. Fang; A. biflora var. minor Nakai; A. brachystemon (Diels) Rehder; A. davidii Hance; A. hersii Nakai; A. onkocarpa (Graebner) Rehder; A. zanderi (Graebner) Rehder; Linnaea dielsii Graebner; L. onkocarpa Graebner; L. umbellata Graebner & Buchwald; L. zanderi Graebner; Zabelia brachystemon (Diels) Golubkova; Z. dielsii (Graebner) Makino.

Shrubs deciduous, 2–3 m tall. Young branchlets reddish brown, becoming gray. Petiole 4–7 mm, sparsely hispid. Leaf blade narrowly ovate or obovate to lanceolate, 30–80 × 5–30 mm, abaxially glabrous but with stiff hairs on veins, adaxially sparsely pubescent when young, base cuneate to obtuse, margin entire or with 1–6 pairs of teeth, apex acute to long acuminate. Inflorescence a reduced terminal cyme consisting of paired axillary flowers (occasionally more due to supernumerary flowers axillary to bracteoles); peduncles 0–12 mm; cyme with 6 bracts at base of ovaries (2 bracts and 4 bracteoles); bracts 1–6 mm, bracteoles 1–6 mm. Sepals 4, ovate-lanceolate or obovate. Corolla white to reddish, 4-lobed; lobes orbicular, 1/3 or 1/5 as long as tube; tube pubescent inside. Stamens 4, didynamous, included. Ovary to 8 mm, with sparse, stiff hairs; styles long, equalling corolla, slender; stigma capitate, not exserted from corolla tube. Achene 1–1.5 cm, crowned with 4 persistent and slightly enlarged sepals. Fl. Apr–Jun, fr. Aug–Sep.

Scrub, forests, grasslands; 800–3700 m. Anhui, SE Gansu, Hebei, Henan, Hubei, Hunan, Liaoning, ??Nei Mongol, Shaanxi, Shanxi, Sichuan, Yunnan, Zhejiang [Japan, Korea, Russia (Far East)].

Zabelia sect. Biflorae was revised by Nakai (1937) who distinguished 11 species (J. Jap. Bot. 13(8): 1–12). We currently recognize only one species in this section.

Most of the differences between taxa were based on the length of the peduncles, bracts, and bracteoles, which are highly variable.

2. ABELIA R. Brown in Abel, Narr. J. China, App. B, 376. 1818.

六道木属  liu dao mu shu

Shrubs deciduous or semi-evergreen. Vegetative buds exposed, ovate-orbicular, small, with several pairs of scales. Leaves opposite, rarely 3- or 4-whorled, shortly petiolate with an interpetiolar line, estipulate. Leaf margin entire to dentate or crenate-serrate. Inflorescence a reduced cyme of 1 or 2 flowers; conflorescence sometimes a thyrse; cymes either with 6 bracts (Abelia sect. Serratae and Abelia sect. Abelia) or 4 bracts (Abelia sect. Uniflorae) at base of ovaries. Sepals 2–5, spreading, narrowly oblong, elliptic, or spatulate, persistent. Corolla 5-lobed, infundibuliform or bilabiate, white, yellow, pink, or red. Corolla tube gibbous ventrally at base containing a nectary of dense glandular hairs. Stamens didynamous, adnate to corolla tube, included or exserted; anthers introrse. Ovary narrowly oblong, 3-locular, 2 locules with 2 series of sterile ovules, 1 locule with a single fertile ovule; style filiform; stigma capitate. Fruit an oblong, leathery achene, crowned with persistent sepals. Seed subterete, testa membranous; endosperm fleshy.

Five species: China, Japan; five species in China (two endemic).

1a.     Flowers solitary within conflorescence; ovary with 4 bracts at the base.

2a.    Sepals usually 5* .........................................................................................  4. A. forrestii

2b.    Sepals usually 2* ........................................................................................  5. A. uniflora

1b.     Flowers paired within conflorescence; ovaries with 6 bracts at base.

3a.    Corolla infundibuliform, with conspicuously exserted stamens and style; conflorescence forming a dense terminal panicle in autumn; paired flowers opening consecutively, flowers fragrant  1. A. chinensis

3b.    Corolla bilabiate, stamens inserted; flowers usually at ends of short axillary branches during spring; paired flowers opening simultaneously, flowers not fragrant.

4a.     Sepals usually 2 .......................................................................................  2. A. serrata

4b.     Sepals usually 5 ..................................................................................  3. A. spathulata

 

*5a. Abelia x grandiflora (A. uniflora x A. chinensis) is a commonly cultivated hybrid with sepals varying from 2–5.

Abelia sect. Abelia

1. Abelia chinensis R. Brown in Abel, Narr. J. China, App. B, 376. 1818.

糯米条  nuo mi tiao

Abelia aschersoniana (Graebner) Rehder; A. chinensis var. ionandra (Hayata) Masamune; A. hanceana M. Martens ex Hance; A. ionandra Hayata; A. rupestris Lindley; Linnaea aschersoniana Graebner; L. chinensis (R. Brown) A. Braun & Vatke; L. rupestris (Lindley) A. Braun & Vatke.

Shrubs deciduous to semi-evergreen, to 2 m tall, highly branched. Young branchlets reddish brown, slender, pubescent. Leaves opposite, sometimes in whorls of 3; leaf blade ovate, 2–5 × 1–3.5 cm, abaxially sparsely pubescent, densely white villous on midvein and lateral veins at base, base rounded or cordate, margin remotely crenate-serrate, apex acute to long acuminate. Conflorescence a dense terminal panicle. Flowers fragrant, with 6 bracts at base of ovaries (2 supernumerary bracts, 2 bracts, 2 bracteoles); bracts oblong or lanceolate. Ovary cylindric, slightly compressed, pubescent, longitudinally striate. Calyx of 5 sepals, elliptic, 5–6 mm, turning red at fruiting stage. Corolla 5-lobed, white to pink, infundibuliform, 10–12 mm, ca. 2 × as long as sepals, outside pubescent. Stamens and style long exserted; filaments slender, equal in length, inserted at base of the corolla tube. Stigma capitate. Achene crowned with persistent and slightly enlarged sepals. Fl. Aug–Sep, fr. Oct–Nov. 2n = 32.

Mountains; 200–1500m. Fujian, Guangdong, Guangxi, Guizhou, ??Henan, Hubei, Hunan, ??Jiangsu, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [Japan] ?Commonly cultivated.

Abelia sect. Serratae (Graebner) Rehder

2. Abelia serrata Siebold & Zuccarini, Fl. Jap. 1: 76. 1835.

黄花六道木  huang duo liu dao mu

Shrubs deciduous, to 3 m tall. Branchlets brown, pubescent. Leaf blade ovate, ca. 5 × 2.5 cm, both surfaces sparsely pubescent, more densely so on veins, base cuneate, margin entire or remotely serrate, ciliate, apex acute to acuminate. Flowers paired; ovaries with 6 bracts at base (2 bracts and 4 bracteoles); peduncle 2–3 mm; bracts lanceolate, 2–3 mm. Ovary 8–10 mm, pubescent. Calyx of usually 2 oblong sepals, ca. 10 × 6 mm, apex sometimes lobed. Corolla bilabiate, yellow or yellow-green, with orange markings on lower lip; corolla tube ca. 18 mm, inside villous. Stamens 4, didynamous, partially adnate to corolla tube, slightly exserted. Style filiform, slightly exceeding stamens; stigma capitate. Achene crowned with usually 2 slightly enlarged sepals. Fl. May, fr. Sep.

Forests; ca. 900 m. Zhejiang (Yongjia) [Japan].

3. Abelia spathulata Siebold & Zuccarini, Fl. Jap. 1. 77. 1835.

温州六道木  wen zhou liu dao mu

Shrubs deciduous, to 3 m tall. Branchlets brown, glabrous. Petiole to 4 mm; leaf blade ovate, ca. 6 × 3 cm, both surfaces sparsely pubescent, base rounded, margin entire to remotely serrate and sinuate, apex acuminate to caudate. Flowers paired; ovaries with 6 bracts at base (2 bracts and 2 bracteoles); peduncle 4–9 mm; bracts lanceolate, 2–3 mm. Calyx reddish; sepals usually 5, oblong-lanceolate. Corolla to 25 mm, bilabiate, upper lip 2-lobed, lower lip 3-lobed, pink or white tinged with yellow, villous inside and with orange markings on lower lip; corolla tube constricted in middle. Stamens 4, didynamous; filaments partially adnate to corolla tube. Style filiform, equalling corolla tube; stigma capitate. Achene glabrous or sparsely pubescent, crowned with usually 5 persistent and slightly enlarged sepals. Fl. May, fr. Jun–Oct.

Forests; 700–900 m. Zhejiang (Wenzhou) [Japan].

Abelia sect. Uniflorae (Graebner) Rehder

4. Abelia forrestii (Diels) W. W. Smith, Notes Roy. Bot. Gard. Edinburgh 9: 76. 1916.

细瘦六道木  xi shou liu dao mu

Linnaea forrestii Diels, Notes Roy. Bot. Gard. Edinburgh 5: 178. 1912; Abelia gracilenta W. W. Smith; A. gracilenta var. microphylla W. W. Smith; A. microphylla (W. W. Smith) Golubkova.

Shrubs deciduous, to 2 m tall. Branchlets red-brown, becoming gray, densely pubescent. Leaf blade narrowly elliptic to lanceolate, 3–7 × 1–2 cm, papery, base obtuse, margin entire, apex acute. Inflorescence a cyme reduced to a solitary flower; conflorescence an axillary thyrse with 1–flowers, ?fragrant; peduncles 3–4 mm; ovary with 4 linear bracts at base (1 pair of bracts and 1 pair of bracteoles). Calyx reddish in fruit; sepals 5, oblanceolate, 6–8 × 1–2 mm, 3-veined, sparsely pubescent. Corolla white to rose, bilabiate, outside pubescent or glandular hairy, inside sparsely villous; corolla tube to 45 mm, expanded in upper part, gibbous at base; lobes 5, orbicular, 4–5 mm; lip long pubescent. Stamens 4; filaments ca. as long as corolla; anthers elliptic, to 2 mm. Style sparsely pubescent; stigma capitate, slightly exserted. Achene to 7 mm, crowned with 2 persistent and enlarged sepals. Fl. May–Sep, fr. Oct.

* Sunny places on mountain slopes, scrub; 1900–3300 m. SW Sichuan, NW Yunnan.

5. Abelia uniflora R. Brown in Wallich, Pl. Asiat. Rar. 1. 15. 1830.

二翅六道木  er chi liu dao mu

Abelia engleriana (Graebner) Rehder; A. graebneriana Rehder; A. longituba Rehder; A. macrotera (Graebner & Buchwald) Rehder; A. mairei H. Léveillé; A. myrtilloides Rehder; A. parvifolia Hemsley; A. schischkinii Golubkova; A. schumannii (Graebner) Rehder; A. tereticalyx (Graebner) Rehder; A. verticillata H. Léveillé; Linnaea engleriana Graebner; L. koehneana Graebner; L. macrotera Graebner & Buchwald; L. parvifolia (Hemsley) Graebner; L. schumannii Graebner; L. tereticalyx Graebner & Buchwald.

Shrubs deciduous, to 4 m tall. Branchlets red-brown, glabrous. Leaves highly variable in shape and size, ovate, orbicular, or lanceolate, 1–8 × 0.5–3.5 cm, abaxially densely white pubescent on midvein and base of lateral veins, adaxially sparsely pubescent, base cuneate, margin subentire or remotely serrate, apex obtuse to acuminate. Inflorescence a cyme reduced to a solitary flower; conflorescence an axillary thyrse with 1–3 flowers. Corolla white to purplish pink, bilabiate, gibbous at base of tube, 5-lobed, 25–50 mm, outside pubescent, inside villous on lip; upper lip 2-lobed, lower lip 3-lobed, bearded and with reticulate orange markings. Ovary pubescent, with 4 ovate to lanceolate bracts at base (1 pair of bracts and 1 pair of bracteoles). Calyx of 2 oblong to elliptic sepals, 10–15 mm, ca. 1/3 as long as corolla tube. Stamens 4, didynamous; filaments partially adnate to corolla tube. Style ca. as long as corolla tube; stigma capitate. Achene 6–15 mm, pubescent, crowned with 2 persistent and slightly enlarged sepals. Fl. Apr–Jun, fr. Aug–Oct.

* Thickets, forests; 200–2000 m. Fujian, Gansu, Guangxi, Guizhou, Henan, Hubei, Hunan, Shaanxi, Sichuan, Yunnan.

This species shows variation in leaf and sepal shape, ovary length, and corolla size. Several forms have been recognized as separate species in the past. The holotype of Abelia uniflora is held in Cambridge University Herbarium (CGE) and the name was validly published by R. Brown within the description of Zabelia trilfora.

5a. Abelia x grandiflora (Rovelli ex André) Rehder, Cycl. Amer. Hort. 1: 1. 1900.

二翅六道木 

Abelia rupestris Lindley f. grandiflora Rovelli ex André, Rev. Hort. 58: 488. 1886; A. rupestris hybrida Rovelli ex Schaedtler; A. floribunda hybrida hort. ex Zabel; A. multiflora hybrida hort. ex Zabel; A. x rupestris hort. ex Späth; A. uniflora hort.; Linnaea biflora x rupestris Koehne.

Shrubs semi-evergreen, 1–1.5 m tall. Branchlets reddish brown, becoming gray, pubescent. Leaves sometimes whorled on reiteration shoots (3- or 4-whorled). Leaves adaxially glossy green, sometimes coppery tinged, ovate, to 4.5 cm, glabrous or with tuft of hairs on veins abaxially, base cuneate, margin remotely serrate with unequal teeth, apex acute. Inflorescence a cyme reduced to a solitary flower; conflorescence a large terminal panicle; peduncle 2–4 mm; flower with 4 bracts at base of ovary. Sepals varying from 2–5, reddish, often partly united, lanceolate, apex acute. Corolla white, sometimes tinged with pink, infundibuliform to slightly bilabiate, gibbous at base, ca. 20 mm, slightly fragrant, hirsute bearded with long hairs on lower lip. Stamen nearly as long as corolla tube; filaments partially adnate to corolla, generally inserted but sometimes slightly protruding, glabrous. Ovary 2–8 mm, slender, minutely pilose; style slightly exserted, 17–18 mm, glabrous; stigma capitate. Achene 8–10 mm, slender, sparsely pilose or glabrous. Achene with persistent sepals at apex. Fl. Jun–Oct; fr. Sep–Nov.

??distribution

3. KOLKWITZIA Graebner, Bot. Jahrb. Syst. 29: 593.1901.

实属  wei shi shu

Shrubs deciduous. Vegetative buds with several pairs of pubescent scales. Leaves opposite, shortly petiolate with an interpetiolar line, estipulate. Inflorescence a reduced cyme of 1 or 2 flowers; conflorescence a thyrse; cymes variable, either with 6 bracts (paired flowered cyme) or 4 bracts (single flowered cyme) at base of ovaries; ovaries of paired flowers connate, flowering consecutively, each crowned with a long narrow beak; bracts densely hairy, adnate to ovaries, accrescent and becoming bristly in fruit. Calyx of 5 narrow, sparsely pubescent sepals. Corolla bilabiate, 5-lobed; lobes spreading. Stamens 4, didynamous, partially adnate to corolla tube. Ovary 3-locular, 2 locules with 2 series of sterile ovules, 1 locule with a single fertile ovule. Achenes single or connate in pairs, bristly, each crowned with persistent sepals.

One species: endemic to China.

1. Kolkwitzia amabilis Graebner, Bot. Jahrb. Syst. 29: 593. 1901.

  wei shi

Kolkwitzia amabilis var. tomentosa Pampanini, Nuovo Giorn. Bot. Ital. 17(4):721. year??

Shrubs erect, to 3m tall. Branchlets red-brown, hirsute, becoming glabrous. Petiole 1–2 mm; leaf blade elliptic to ovate, 3–8 × 1.5–2.5 cm, both surfaces sparsely pubescent, densely spreading pubescent and ciliate on veins and margin, base rounded or broadly cuneate, margin entire, rarely shallowly dentate, apex acute to acuminate. Conflorescence a thyrse; peduncle 10–15 mm; bracts lanceolate, tightly adnate to base of ovary. Sepals 5, lanceolate, to 5 mm, pubescent. Corolla whitish, tinged pink, 15–25 mm, outside pubescent, base very narrow, abruptly enlarged above middle. Lower lip 3-lobed, bearded, with orange reticulate markings, upper lip 2-lobed. Ovary bottle-shaped with a long neck; style pubescent; stigma capitate, inserted. Achenes densely yellow bristly, apex elongated, crowned with persistent sepals. Fl. May–Jun, fr. Aug–Sep. 2n = 32*.

* Mountain slopes, roadsides, scrub; 300–1300 m. Anhui, Gansu, ??Hebei, Henan, Hubei, Shaanxi, Shanxi.

This species is rare in the wild but widely cultivated.

4. DIPELTA Maximowicz, Bull. Acad. Imp. Sci. Saint-Pétersbourg, sér. 3, 24: 50. 1877.

双盾木属  shuang dun mu shu

Shrubs erect, deciduous. Vegetative buds with several pairs of scales; branchlets pubescent. Leaves opposite, estipulate, shortly petiolate, slightly pubescent on veins and margin, margin entire or dentate at apex. Inflorescence a reduced cyme of 1 flower (exceptionally 2); conflorescence a thyrse; ovaries with 4 bracts at base; bracts unequal in size, larger 2 tightly adnate to ovary. Sepals 5, linear to lanceolate, ± fused at base. Corolla bilabiate, distinctly narrowed and gibbous at base, upper lip 2-lobed, lower lip 3-lobed. Stamens 4, didynamous, partially adnate to corolla tube, included. Ovary 4-locular, 2 locules with 2 series of sterile ovules, 2 locules each with a single fertile ovule; style slightly shorter than corolla. Fruit an achene, crowned with persistent calyx, with 2 accrescent, membranous, wing-like bracts at base.

Three species: all endemic to China.

1a.  Sepals fused into a tube for at least 2/3 of their length; style villous ........................  3. D. elegans

1b.  Sepals fused for less than 1/2 their length; style glabrous to sparsely hairy.

2a.  Narrow part of corolla tube exserted from sepals; accrescent bracts peltate, obscuring sepals at anthesis      1. D. floribunda

2b.  Narrow part of corolla tube not exserted from sepals; accrescent bracts reniform, not obscuring sepals at anthesis ..................................................................................................  2. D. yunnanensis

 

1. Dipelta floribunda Maximowicz, Bull. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 3, 24: 51. 1877.

双盾木  shuang dun mu

Dipelta floribunda var. parviflora Rehder, J. Arnold. Arbor. 5: 241. 1924; D. wenxianensis Y. F. Wang & Lian, Acta. Bot. Boreal.-Occid. Sin. 14(6): 91 (92 fig). 1994. (as seen on the illustration but type specimen not located).

Shrubs or small trees to 6 m tall, deciduous. Branchlets glandular hairy, becoming glabrous. Petiole 6–14 mm; leaf blade ovate to lanceolate, 4–10 × 1.5-6 cm, midvein white pubescent, base cuneate or obtuse, margin entire, occasionaly remotely 2- or 3-dentate at apex, apex acute or long acuminate. Inflorescence a reduced cyme of 1 flower (exceptionally 2); conflorescence a terminal thyrse on short, lateral branches; peduncles to 10 mm; ovaries with 4 bracts tightly adnate to base; 2 outer bracts larger, peltate, obscuring sepals at anthesis, accrescent to 20 mm wide when mature, dry membranous, with conspicuous veins; 2 inner bracts smaller, unequal, one ovate, other smaller and narrowly elliptic. Calyx 5-lobed, glandular hairy; lobes linear, 5–7 mm. Corolla white to pink, bilabiate, narrowly cylindric at base, flaring and campanulate in upper part, 25–40 mm, lower lip with reticulate orange-yellow markings. Ovary pubescent; style filiform, glabrous to sparsely hairy. Achene crowned with persistent calyx, enclosed within 2 accrescent, membranous, wing-like and peltate bracts. Fl. Apr–Jul, fr. Aug–Sep.

* Mixed forests, scrub; 600–2200 m. Gansu, Guangxi, Hubei, Hunan, Shaanxi, Sichuan.

2. Dipelta yunnanensis Franchet, Rev. Hort. 11: 246. 1891.

云南双盾木  yun nan shuang dun mu

Dipelta ventricosa Hemsley.

??· Dipelta yunnanensis var. brachycalyx Handel-Mazzetti, Anz. Akad. Wiss. Wien, Math.-Naturwiss. Kl. 61: 201. 1924. Yunnan: 9950.

Shrubs deciduous, to 4 m tall. Branchlets pubescent. Petiole to 5mm; leaf blade elliptic to lanceolate, 5–10 × 2–4 cm, abaxially white villous on midvein, ciliate along margin, adaxially sparsely pubescent, base cuneate to rounded, margin entire or rarely remotely dentate, apex acuminate to long acuminate. Inflorescence a reduced cyme of 1 flower (exceptionally 2); conflorescence a terminal thyrse on short, lateral branches; ovaries with 4 bracts; 2 outer bracts larger, reniform, accrescent in fruit and not obscuring calyx at anthesis; 2 inner bracts smaller, unequal, one ovate, other smaller and narrowly elliptic. Calyx 5-lobed, pubescent; lobes divided to 2/3, splitting to base on one side and spreading in fruit. Corolla white to dark pink, bilabiate, 20–40 mm, very shortly narrowly tubular at base, lower lip pubescent with yellow-orange reticulate markings. Ovary pubescent; style inserted. Achene ovoid with a short neck, pubescent. Achene crowned with persistent calyx, enclosed within 2 accrescent, membranous, wing-like and reniform bracts. Fl. May–Jul, fr. Jul–Nov.

* Mixed forests, scrub; 800–2400 m. Gansu, Guizhou, Hubei, Shaanxi, Sichuan, Yunnan.

3. Dipelta elegans Batalin, Trudy Imp. S.-Peterburgsk. Bot. Sada 14: 174. 1895.

优美双盾木  you mei shuang dun mu

Shrubs deciduous, to 2 m tall. Leaf blade elliptic, 5–10 × 2–5 cm, glandular hairy, pubescent on veins and margin, base acute to rounded, margin entire or remotely dentate in upper 1/2, apex long acute to long acuminate. Inflorescence a reduced cyme of 1 flower; peduncles to 20 mm; conflorescence a terminal thyrse on short, lateral branches; ovaries with 4 bracts; 2 outer bracts larger, peltate, accrescent in fruit and not obscuring calyx at anthesis. Calyx tubular-campanulate, 5-lobed, sparsely hairy; lobes obtuse, ca. 1/3 as long as tube. Corolla bilabiate, densely pubescent, narrowly cylindric at base, flaring and campanulate in upper part, lower lip villous. Ovary pubescent; style pilose to sparsely pubescent; stigmas capitate. Achene with 2 very large persistent peltate bracts to 40 mm wide. Fl. May–Jun, fr. Sep.

* Broad-leaved forests; ca. 2000 m. Gansu, ??Shaanxi, Sichuan.

5. LINNAEA Linnaeus, Sp. Pl. 2: 631. 1753.

北极花属  bei ji hua shu

Subshrubs creeping, evergreen. Branchlets ascending, slender. Leaves opposite, petiolate, estipulate. Inflorescence a reduced cyme of 1 flower; conflorescence of 2 flowers (2 cymes), paired at apices of lateral branchlets; peduncles long, slender; ovary with 2 pairs of bracts adnate to base; 2 outer bracts larger and shield-like, covered with stipitate glandular hairs; 2 inner bracts minute; peduncles, ovaries, and sepals glandular hairy. Sepals 5, deciduous. Corolla infundibuliform, 5-lobed, slightly bilabiate. Stamens 4, didynamous, inserted in corolla tube; anthers included. Ovary 3-locular, 2 locules with 2 series of sterile ovules, 1 locule with a single fertile ovule; style filiform; stigmas capitate. Fruit a 1-seeded achene.

One species: widely distributed in alpine and cold regions of the Northern Hemisphere.

1. Linnaea borealis Linnaeus, Sp. Pl. 2: 631. 1753.

北极花  bei ji hua

Linnaea borealis f. arctica Witrock.

Subshrubs creeping, evergreen, 5–10 cm tall. Stem red-brown, long, slender, sparsely short pubescent. Petiole 3–4 mm; leaf blade orbicular to obovate, abaxially glabrous, adaxially sparsely pubescent, margin with 1–3 pairs of rounded teeth. Flowers fragrant, nodding; peduncles 60–70 mm; inflorescence bracts linear, shortly pubescent; inflorescence peduncles less than 10 mm. Sepals shortly pubescent, narrowly acute. Corolla white to pink, to 10 mm; tube glabrous outside, shortly pubescent inside. Stigma capitate. Achenes nodding, yellow, orbicular. Fl. and fr. Jul–Aug. 2n = 32.

Needle-leaved forests, on trunks, mossy rocks; 700–2300 m. Hebei, Heilongjiang, Jilin, ??Liaoning, Nei Mongol, Xinjiang [widely distributed in other parts of northern temperate regions].