Chinese Flora of Cornaceae L. s. l.

This work has been published in: Xiang, Q.-Y. & Boufford, D. E. 2005. Cornaceae, Mastixiaceae, Toricelliaceae, Helwingiacaee, Aucubaceae. Pp. 206–234 in: Wu, Z. Y. & Raven, P. H. (eds.), Flora of China. Vol. 14 (Apiaceae through Ericaceae). Science Press, Beijing, and Missouri Botanical Garden Press, St. Louis.

The following manuscript is not the final version of the publication. It may contain information slightly different from the published version.

Cornaceae
Shan Zhu Yu Ke
(Jenny) Qiu-yun and David E. Boufford

(Excluding Aucuba, Helwingia, Toricellia, Nyssa, Alangium, Davidia, Mastixia)

Trees or shrubs, rarely rhizomatous herbs, mostly deciduous, sometimes evergreen, hermaphroditic (but dioecieous in the African species). Old branches terete, pithed, often with conspicuous lenticels and leaf scars. Young branches round, or slightly 4-ridged. Nodes slightly swollen. Leaves opposite, rarely alternate or whorled, exstipulate, petiolate (rarely sessile); blade simple, entire, pinnately veined (rarely parallelly veined), often pubescent, sometimes pappilate. Hairs often two-armed; arms equal or unequal, appressed (T-shaped) or elevated (V-shaped, Y-shaped, or pseudo-filiform). Inflorescences compound cymes, paniculate, corymbose, umbellate, or capitulate, terminal, rarely axillary; bractes minute, non-petaloid and early deciduous, or 4-folds (rarely 6), showy or non-showy. Flowers small, typically 4- merous, polypetalous. Calyx small, tubulate, fused to ovary, minutely 4-dentate or truncate. Petals 4, creamy white to yellow (rarely dark redish purple or partially dark redish purple), valvate. Androecium 4-staminate, around a fleshy floral disk; stamens alternate with petals; anthers dehiscing via longitudinal slits. Gynoecium two carpellate, rarely 1, 3, or 4 carpellate. Ovary inferior, typically 2-locular, rarely 1, 3, or 4-locular. Ovules pendulous, solitary per locule. Style 1, columnar. Stigma capitate, disciform, punciform, or truncate, sometimes slightly 2-lobed. Fruit a drupaceous berry, white, blue, red, or black, distinct or fused into a fleshy syncarpous compound fruit; fruit stone bony, 1-2 (--4)-seeded. Endosperm oily, cotyledons 2, leafy.

Monogeneric. Widespread in temperate regions, extending to tropical and circumboreal areas of the Northern Hemisphere; with one species in tropical Africa and 1 or 2 species in South America.

Woody species produce hard wood used for farming tools; fruits of some species are sources of food or industrial oil. Cornus mas is cultivated in China for medicinal uses. Many species are widely cultivated as ornamentals, such as Cornus stolonifera Maximum., C. alba L., C. controversa Hemsl., C. mas L., C. nuttallii Audub., C. florida L., C. kousa Hance, and C. canadensis L.

The circumscription and relationships among members of Cornaceae have been extremely controversial. A total of 17 diverse genera have been ascribed to the family at one time or another. The family has been defined to consist of a single genus Cornus to as many as 15 genera by various authors (e.g., Takhtajan, Sistema Magnoliofitov, 1987; Harms, Ber Deutsch. Bot. Ges., 15: 1. 1898; Cronquist, Integr. Syst. Classif. Flowering Plants, 1988; See Xiang et al., Ann. Missouri Bot. Gard. 80: 723, 1993; Xiang et Soltis in Bourfford & Ohba, Sino-Japanese flora-its characteristics and diversification, p123, 1998). Recent phylogenetic analyses of the chloroplast gene rbcL sequences (Xiang, Harvard Pap. Bot., 4: 527, 1999; Xiang et al., Ann. Missouri Bot. Gard. 80: 723, 1993; Xiang and Soltis in Boufford and Ohba, Sino-Japanes flora-its characteristics and diversification, p123, 1998) suggested that nine genera (Aralidium, Aucuba, Corokia, Garrya, Griselinia, Helwingia, Kaliphora, Melanophylla, and Toricellia) previously placed in Cornaceae by different authors were not closely related to Cornus and should be removed from Cornaceae. Evidence from other studies, including pollen morphology and wood anatomy (Furguson and Hiduex, Proc. IV Int. Palynol. Conf., Lucknow 1: 240, 1980; Li & Chao, Quart., J. Taiwan Mus. 7: 119, 1954; Noshiro et Baas, IAWA J. 19: 43, 1998; also see Eyde, Bot. Rev. 54: 1, 1988) also support the removal of these genera from Cornaceae. Various broad phylogenetic analyses of rbcL and other chloroplast gene sequences identified a strongly supported monophyletic Cornales consisting of Cornus, Alangium, Camptotheca, Davidia, Nyssa, Diplopanax, Mastixia, Curtisia, Hydrangeaceae, Loasaceae, Grubbiaceae, and Hydrostachyaceae (Chase et al., Ann. Missouri Bot. Gard. 80: 528, 1993; Xiang & Soltis in Boufford and Ohba, Sino-Japanes flora-its characteristics and diversification, p123, 1998; Xiang, Harvard Pap. Bot. 4: 527, 1999; Albach et al., in press; Olmstead et al., Mol. Phyl. Evol. 16: 96, 2000; Soltis et al., Bot. J. Linn. Soc. 133: 381, 2000; Savolainen et al., Kew Bull. 55: 257, 2000; Savolainen et al., Syst. Biol. 49: 306, 2000). Phylogenetic analyses of rbcL and matK sequences for the Cornales suggested a Cornaceae consisting of Cornus and Alangium, a Grubbiaceae consisting of Curtisia and Grubbia (Xiang, Harvard Pap. Bot. 4: 527, 1999; Xiang et al., Amer. J. Bot. 85:, 285, 1998; Xiang et al., Mol. Phyl. Evol., in press). The monophyly of Cornaceae of Eyde (Bot. Rev. 54: 1. 1988) that consist of Cornus, Nyssa, Davidia, Camptotheca, Mastixia, and Diplopanax was not strongly supported by these molecular data.

The Cornaceae of Hu and Soong (in Fang Wen-bei & Hu Wen-kuang, eds., Fl. Reipubl. Popularis Sin. 56: 10-108. 1990) in the Chinese version of Flora of China included Mastixia, Cornus s. l., Aucuba, Helwingia, and Toricellia. Given that the latter three genera are allied with higher asterids in broad phylogenetic analyses, they are, therefore, removed from the Cornaceae in this treatment. These genera will be recoganized as three monotypic families following Takhtajan (Sistema Magnoliofitov, 1987). Based on molecular data, Mastixia is a close relative, but not the sister of Cornus. Its relationships to Cornus and other close relatives of Cornus remain unclear. Thus, here we also treat Mastixia as a family separate from Cornaceae. Although molecular data has suggested that Alangium is a member of Cornaceae, the genus has been treated as a separate family in a different volume in Flora of China. Therefore, the Cornaceae in this treatment include only Cornus L. s. l.

1. CORNUS L., Sp. Pl. 117. 1753.
Shan Zhu Yu Shu

Benthamia Lindl.; Benthamidia Spach, Bothrocaryum (Koehne) Pojark.; Chamaepericlimenum Aschers. & Graebn.; Chamaepericlymenum Hill; Cornella Rydberg; Cynoxylon Raf.; Dendrobenthamia Hutch.; Eukrania Raf.; Macrocarpium (Spach) Nakai; Svida Opiz,;Telukrania Raf.

Shrubs, trees, or rarely herbs, precocious or non-precocious (postcocious?). Young shoots pubescent or rarely glabrous. Hairs curly or straight, elevated or appressed. Stem sympodial, rarely monopodial. Winter buds terminal or axillary, mixed or separate, covered or exposed. Leaves long elliptic, elliptic, oblong, or ovate; glabrous to densely pubescent; lateral veins actinodromous, often elevate on lower surface; petioles slightly furrowed above. Inflorescence preformed or formed in the current year; bracts protective or non-protective. Petals 4, free, oblong to round, spreading, 1.5-5 mm long, valvate. Sepals 4, fused; teeth absent or minute, or various triangular. Filaments filiform or awn shaped, longer than the style, longer or shorter than the petals. Anthers whitish or yellow, rarely blue, red, or purplish, 2-locular, elliptic to long-elliptic or long oblong. Ovary small, obovate, crowned by a pulvate disk. Fruit globose, ovoid, oblong, or ellipsoid, crowned by the persistent calyx, disk, and style; stones globose, ovoid, ellipsoid, oblong, or asymmetric, smooth or ribbed on surface, rarely pitted at apex.

About 55 species among 9 subgenera: North America, Central America, Asia, a few scattered in Europe, South America and Africa. 25 species of six subgenera in China.

The classification of the genus has long been debated. The ranks and circumscriptions of subgroups within Cornus vary considerably among taxonomists. The current treatment remains a broad sense of the genus as defined by Linnaeus and represents a synthesis of Ferguson (J. Arn. Arb. 47: 100, 1966), Murrell (Syst. Bot. 18: 469, 1993), and Xiang (Acta Phytotax. Sin. 25: 125, 1987). This treatment agrees with recent molecular phylogenetic analyses for Cornus.


  1a. Perennial rhizomatous herbs; leaves appearing whorled at summit; inflorescence 
        minute corymbose cymes subtended by 4 white, petaloid bracts...............................................................6. Subg. Arctocrania
  1b. Trees or shrubs; leaves opposite or alternate; inflorescences paniculate, 
        corymbose, umbellate, or capitular cymes.
        2a. Inflorescence capitulate, subtented by 4 large, showy, petaloid bracts; 
              fruit a compound drupaceous berry ..................................................................................................5. Subg. Syncarpea
        2b. Inflorescences not capitulate, bracts non-showy; fruit distinct.
              3a. Inflorescences umbellate, precocious; bracts 4, rounded, ca. 3-4 mm; fruit 
                    oblong, red to blackish red…......................................................................................................4. Subg. Cornus
              3b. Inflorescence paniculate or corymbose cymes, postcocious(?);bracts along 
                    branches, minute, linear or branched; fruits globose, ovoid, or rarely ellipsoid, 
                    white, blue, or black.
                4a. Leaves alternate; apex of fruit stone conspicuously pitted. ..........................................................2. Subg. Mesomora
                4b. Leaves opposite, rarely subopposite; apex of fruit stone not pitted.
                      5a. Bracts at the base of inflorescence branches, often persistent to anthesis; 
                            leaves evergreen, leathery; style cylindric; stigma small, subcapitate or punciform; 
                            fruit stones (2--) 3--4 locular; anthesis fall and winter...........................................................1. Subg. Yinquania
                      5b. Bracts displaced distal and adnate to the inflorescence branches, early 
                            deciduous; leaves paperous or rarely leathery; style cylindric or clavate; stigma 
                            capitate, disciform, rarely punciform; fruit stones 2-locular; anthesis spring 
                            and early summer, rarely fall.................................................................................................3. Subg. Kraniopsis
  
1. Subgen. Yinquania (Zhu) Murrell, Syst. Bot. 18: 476. 1993.

Chang Yuan Ye Lai Mu Ya Shu

Yinquania Zhu, Bull. Bot. Res. 4(4): 121. 1984. Cornus subsect. Oblongifoliae Wanger.; Swida sect. Oblongifoliae (Wanger.) W. K. Hu.

Evergreen trees or shrub. Buds terminal or axillary, narrow awn-shaped, pubescent with gray short hairs. Leaves typical opposite, rarely sub-opposite at some nodes, leathery; blade narrow elliptic to oblong-elliptic or oblong lanceolate; abaxial surface glabrous, or papillate and pubescent with grayish appressed short two-armed hairs, or densely pubescent with long, dense soft hairs. Paniculate cymes terminal; bracts small to minute, green, sometimes leaf-like, at base of inflorescence branches, and often persistent to at least anthesis. Calyx tube conspicuously 4-dentate; teeth ovate triangular. Petals white, long elliptic. Anthers purplish yellow, elliptic. Ovary (2--)3--4-locular. Style cylindric. Stigma small, subcapitate to punciform. Fruit ellipsoid, oblong, or globose, 2-4-seeded, purple red ripen into black; stones bony, non-pitted, pointed at one or both ends. 2n = 22.

One species, 3 varieties, from eastern Himalaya to Central China. All in China.

1. Cornus oblonga Wall., in Roxb., Fl. Ind. 1:432. 1820.
Chang Yuan Ye Lai Mu
Yinquania muchuanesis Z. Y. Zhu, Bull. Bot. Res., Harbin 4(4): 123. 1984; Swida muchuanensis (Z. Y. Zhu) Holub, Preslia 70: 114. 1998. Swida oblonga (Wall.) Sojak, Novit. Bot. & Sem. Hort. Bot. Univ. Carol Prag. 10. 1960; Ardisia discolor Levl. In Fedde, Repert. Sp. Nov. 10: 373, 1912.
Evergreen trees 2--16 m. Barks smooth, grayish brown to blackish gray. Current year branches more or less 4-angled, glabrous to densely pubescent; old branches with sparse rounded lenticels and semicircular leaf scars. Leaves 6--13 x 1.6-4 cm, narrow elliptic, oblong elliptic, or lanceolate elliptic, acute or caudate at apex, cuneate at base, slightly revolute at margin, 4-5(--6)-veined; veins elevated below. Paniculate cymes ternimnal, 6-6.5 cm x 6-8 cm, pubescent with white or brown or mixed white and brown hairs. Flowers ca. 8 mm in diam., pedicelate. Calyx teeth ca. 2--3 mm, taller than the disk. Stamens longer or subequal to petals; filaments ca 5.00 mm. Style 2.5--2.8 mm. Fruit ellipsoid, ca 4--6 x 6--7 mm, black when mature; stones ca. 6.0 x 3.8 mm, inconspicuously ribbed. Fl. Sept--Jan., Fr. Apr--June.

In slope thickets, sparse stream forests, or evergreen forests, elev. 1000-3700 m. Guizhou, Hubei, Sichuan, Xizang, Yunnan [Vietnam, Myanmar, Pakistan, northern India, Bhutan, Sikkim, Nepal, Thailand, and Kashmir region].
Fruits may be used as a source of industrial oil or used medicinally as replacement of "Zao Pi" (the flesh of fruits of Cornus officinalis and C. chinensis). Bark contains essential oil and tannins and used in folk remedies to treat arthritis and injuries.

Yinquania muchuanesis Z. Y. Zhu was restricted to Muchuan county of Sichuan, and was recognized as a new species based on its smaller size (2--4 m in height) and subglobose shape of fruit, compared to C. oblonga which is typically a tree of 2--16 m and bearing ellipsoid fruits. Given that the size of Y. muchuanesis falls within the range of C. oblonga and subglobose fruits are also found in C. oblonga from other areas, this species is tentatively treated as part of C. oblonga. More evidence is needed to determine the status of this species.

  1a. Leaves glabrous on both surfaces, smooth by touch............................................................1c. C. oblonga var. glabrescens
  1b. Leaves pubescent.
        2a. Leaves pubescent with grayish, short, appressed hairs on abaxial surface; 
              rough by touch............................................................................................................1a. C. oblonga var. oblonga
        2b. Leaves densely pubescent with long and soft hairs on abaxial surface, soft 
              by touch......................................................................................................................1b. C. oblonga var. griffithii
1a. Cornus oblonga Wall. var. oblonga
Chang Yuan Ye Lai Mu

Leaves often elliptic to oblong, papillate abaxially. Inflorescence and leaf abaxial surface and petioles pubescent with short, grayish, appressed hairs.

In evergreen or evergreen-deciduous mixed forests or thickets along streams; elev. 800-3700 m.

Guizhou, Hubei, Yunan, Sichuan, Xizang [ Pakistan, Nepal, India, Myanmar, Thailand, Bhutan, Sri Lanka, Kesmia, and Vietnam].

1b. Cornus oblonga Wall. var. griffithii C.B. Clark in Hook. f., Fl. Brit. Ind. 2: 745. 1879.
Mao Ye Lai Mu
Cornus oblonga Wall. forma pilosula Li in J. Arn. Arb. 25: 311, 1944; Swida oblonga (Wall.) Sojak var. griffithii (Clarke) W. K. Hu.

Leaves often long elliptic, more or less scurfy abaxially. Inflorescence and leaf abaxial surface and petioles densely pubescent with elevated, long and soft hairs.

In slope thickets or in evergreen forests; elev. 850 - 3000m.

Guizhou, Hubei, Sichuan, Yunan, Xizang [Bhutan, India].

1c. Cornus oblonga Wall. var. glabrescens Fang et W.K.Hu, J. Sichuan Univ., Not. Sci. 3; 156, pl. 2, fig. 2. 1980.
Wu Mao Chang Yuan Ye Lai Mu
Swida oblonga (Wall.) Sojak var. glabrescens (Fang et W. K. Hu) Fang et W, K, Hu, Bull Bot. Res. 4(3): 102. 1984.

Leaves smaller, 7--9 x 3--5 cm, long elliptic, seldom lanceolate; abaxial surface glabrous, smooth, and non-papillate.

In thickets or mixed forests; elev. 1500 -3400 m.

NE & NW Yunnan, SE Xizang.

2. Subg. Mesomora
Raf., Alsograph. Am. 58, 1838.
Deng Tai Shu Ya Shu

Macrocarpium subsect Bothrocaryum Koehne. Gartenflora 45: 285, 1896; Bothrocaryum (Koehne) Pojark, Cornus sect. Bothrocaryum (Koehne) Harms ex Nakai; Cornus sect. Mesomora (Raf.) Nakai.

Deciduous trees or shrubs. Winter buds false terminal or axillary, ovate or cornical, glabrous or sparsely pubescent. Leaves alternate, paperous; blade broad ovate to elliptic-ovate, pubescent with minute, 2-armed, appressed hairs abaxially. Corymbose cymes terminal; bracts displaced distal and adnate to branches, minute, early deciduous. Calyx tube miutely 4-dentate; teeth triangular. Petals white, oblanceolate. Anthers yellowish, elliptic. Ovary 2-locular. Style cylindric. Stigma small, capitate. Fruit globose, 2-seeded, dark purple red ripen into black; stones bony, rectangularly pitted at the summit.

Two species in subtropical and north temperate regions of eastern Asia and eastern North America. One species in China.

1. Cornus controversa Hemsl. ex Prain, Bot. Mag. 125, sub t. 8261. 1909; Kew Bull. 331. 1909.
Deng Tai Shu
Swjda (Swida) controversa (Hemsl.) Sojak., Novit. Bot. & Del. Sem. Hort. Bot. Univ. Carol Prag. 10. 1960; Swida controversa (Hemsl.) Mold., Phytologia, 17: 114. 1968; Bothrocaryum controversum (Hemsl.) Pojark, Not. Syst. Herb. Inst. Bot. Acad. Sci. URSS, 12: 170, fig. 1. 1950; B. longipetiolatum (Hayata) Pojark., Not. Syst. Herb. Inst. Bot. Acad. Sci. URSS, 12: 170, 1950; Cornus brachypoda Miq., Ann. Mus. Bot. Lugd.-Bat. 2: 160, 1865; C. controversial Hemsl. var. angustifolia Wanger. in Engl., Pflanzenreich, 41(IV. 229): 50, 1910; C. longipetiolata Hayata, Econ. Pl. Formes, 4: 11, 1914; C. obovata Thunb., Mus. Nat. Acad. Upsal. App. 17: 3, 1809, nom nud.; Cornus sanguinea auct. Non Linn.: Thunb., Fl. Jap. 62. 1784; C. macrophylla auct. non Wall.: Hemsl., J. Linn. Soc. Bot. 23: 345, 1886, pro parte; C. ignorata auct. non Koch: Franch. & Savatier, Enum. Pl. Jap. 1: 196, 1875.

Trees 3--13(--20) m;barks smooth, dark gray or yellowish gray; branches more or less horizontal; current year branches purplish turning greenish, glabrous or pubescent; older branches greenish, with conspicuous semicircular leaf scars and rounded lenticels. Winter buds ovate or cornical, purplish, 3--8 mm long, glabrous, covered by several alternate, overlapped scales. Leaves 5--13 x 3--9 cm, broad ovate or broad elliptic ovate, acute or acuminate at apex, subrounded at base; abaxial surface gray-green, sparsely pubescent with appressed hairs and papillate; veines 6-7(--9), elevate and slightly purplish below. Corymbose cymes terminal, 5--14 cm in diam, pubescent with appressed hairs. Flowers 8--9 mm in diam.; buds nearly rounded, shortly pedicelate. Calyx teeth ca 0.5 mm, taller than the disk. Stamens longer than petals; filaments 4--5 mm, whitish. Style 2--3 mm, glabrous. Fruit globose, 6--7 mm in diam, purplish red to blue-black; stones globose, 5--6 mm in diam., inconspicuously 8-ribbed. Fl. May--June, fr. July--Sept. 2n = 20.

In broad-leaved or mixed broad-needle-leaved forests at 250--2600 m. Anhui, Gangsu, Guangdong, Guangxi, Guizhou, Hebei, Henan, Hubei, Hunan, Jiangxi, Jiangsu, Liaoning, Shandong, Shangxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Korea, Japan, northern India, Nepal, Sikkim, Bhutan].

Fruits are source of oil used for lubricating and soap industry. Its large conspicuous white inflorescence, layed branching pattern, and pagoda shape of the tree makes the species a good garden ornamental. Leaves are used to relieve pain and reduce swelling in folk remedies.

3. Subg. Kraniopsis Raf., Alsograph. Am. 58, 1838.

Lai Mu Ya Shu

Swida Opiz in Berchrold & Opiz, Oekon-tech. Fl. Bohmens 2: 174-180. 1838; Cornus sect. Thelycrania Dumortier. Florula Belgica 83. 1827; Cornus c. Thelycrania Endlicher, Gen.Pl. 798. March. 1839; Thelycrania (Dumortier) Fourreau; Cornus subg. Thelycrania (Dumortier) Endlicher ex Schneider; Cornus sect. Microcarpium Spach. Hist. Nat. Veg. 8: 92. 1839; C. subsect. Amblycaryum Koenhe. Gartenflora 45: 286. 1896; C. sect. Amblycaryum (Koehne) Nakai; Ossea{Lonicer} Nieuwland & Lunell, in Lunell. Am. Midl. Nat. 4: 487. 1916.

Deciduous trees or shrubs, rarely evergreen. Buds small, terminal or axillary, ovate to awn-shaped, pubescent. Leaves opposite, very rarely alternate at some nodes in two species, paperous, rarely leathery; blades ovate to elliptic; abaxial surface pubescent with appressed or elevated hairs, 3--8(--9)-veined. Inflorescences terminal, compact or loose corymbose or paniculate cymes; bracts minute, linear, rarely branched, non-green, deciduous prior to anthesis. Flowers often fragrant. Calyx tubular, minutetly 4-dentated; teeth inconspicuous to long triangular. Petals white, rarely yellowish white, ovate or oblong. Anthers oblong or oblong ovate, yellowish white to blue-gray. Ovary 2-locular. Style cylindric or calvate. Stigma capitate, disciform, or punciform. Fruit globose, wide globose, oblong globose, or subovate, blue black or black when mature, 2-seeded; stones bony, typically not pitted at apex (but inconspicuously pitted in one species). 2n = 22.

About 30 species, mostly North Temperate, a few in subtropical mountains, 1or 2 in South America. 15 in China; in all provinces execept Xingjiang, mostly in SW China.

Many species produce hard wood that is used for argricultural tools; some species are valuable sources of industrial oils or garden ormentals.
  
  1a. Style cylindric, or slightly expanded at apex, but not clavate. (go to 1b)
        2a. Spreading shrub; fruit creamy white to light blue when mature; fruit stone laterally compressed.......1. C. alba
        2b. Shrubs or trees; fruit blue black to black when mature; fruit stone not laterally compressed.
              3a. Leaves leathery; stigma smaller or subequal to style in diam, puciform or 
                    subcapitate.
                    4a. Leaves broad elliptic; lateral veins 3 (4), conspicuous; small veins horizontal; 
                          flower larger, ca. 9 mm in diam........................................................................................8. C. oligophlebia
                    4b. Leaves elliptic; lateral veins 4--5, thin, inconspicuous; small veins not 
                          so conspicuous; flowers smaller, ca. 7mm in diam............................................... ............2. C. austrosinensis
              3b. Leaf paperous; stigma obviously larger than the style, or rarely smaller 
                    than the style in diam
                            5a. Leaves 3-veined; flowers small, cal 4.5 mm in diam; stigma smaller or equal 
                                  to the style in diam, punciform……........................................................................10. C. parviflora
                            5b. Leaves (2--)3-8(--9)-veined; flowers larger, 7-9 mm in diam; stigma wider 
                                  than the style, not punciform.
                                    6a. Abaxial surface of leaves conspicuously pubescent with more or less curly 
                                          or non-curly soft, elevated hairs (Y-shaped, V-shaped, or pseudo-filiform).
                                          7a. Lateral veins thin and small veins inconspicuous..........................................3. C. bretchneideri
                                          7b. Lateral veins thicker and small veins conspicuous.
                                                8a. Veins 5-7, with dense yellowish hairs; inflorescence broad, loose corymbose 
                                                      cymes, nearly glabrous at anthesis; branches of inflorescences arched inward; 
                                                      flowers sessile or only shortly pediceolate; pedicels thick, up to ca. 2 mm 
                                                      long.................................................................................................  .13. C. ulotricha
                                                8b. Veins 5--8 (9); both veins and interveinal areas of abaxial surface with 
                                                      dense yellowish, grayish, or brown hairs; inflorescences more compact, corymbose 
                                                      or paniculate cymes, densely pubescent at anthesis; branches of inflorescences 
                                                      not arched (straight); flowers longer pedicelated; pedicels slender, 0.5--6 
                                                      mm......................................................................................................12. C. schindleri
                                                                                                                                                                        (including C. fulvescence, C. lixiensis, C. daijingensis, 
                                                                                                                                                                        C. malifolia, C. mombeigii, C. mombeigii subsp. pupolifolia, 
                                                                                                                                                                        C. xanthotricha, C. poliophylla, C. poliophylla var. praelonga, C. scabrida)
                                    6b. Abaxial surface of leaves conspicuously pubescent with short, appressed 
                                          hairs and no elevated hairs except sometimes in the axils of veins.9a. Leaves 3-4(5)-veined.
                                          ........................................................................................................................15. C. wilsoniana
                                          9b. Leaves (5--)6-8(--9)-veined.
                                                10a. Leaves large, 6--17 cm in length; inflorescences large and loose, paniculate 
                                                       or corymbose cymes; filaments thick, equal to petals or slightly exceeding petals; 
                                                       style slightly thickened at apex, appearing subclavate..............................7. C. macrophylla 
                                                                                                                                                                        (incl. C. alpina, C. longipedunculata, brachypoda).
                                                10b. Leaves smaller, 4.5--12 cm; inflorescences smaller and compact, convex 
                                                        corymbose cymes; filaments slender, longer than petals; style cylindric, not 
                                                        subclavate.
                                                        11a. Old branches light yellow; abaxial surface of leaves grayish white, inconspicuously 
                                                                papillate; small veins not conspicuous............................................3. C. bretschneideri
                                                        11b. Old branches red, red-brown, purple-red, or brown; abaxial surface of leaves 
                                                                grayish green, conspicuously papillate; small veins conspicuous.
                                                               12a. Abaxial surface of leaves with blackish streaks; flowers white
                                                                       .............................................................................................9. C. papillosa
                                                               12b. Abaxial surface of leaves without blackish strreaks; flowers white or yellowish
                                                                       .............................................................................................5. C. hemsleyi 
                                                                                                                                                                        (C. alsophila, C. polyantha) 
  1b. Style conspicuously clavate.
        13a. Deciduous shrubs; leaves 4--7 cm long, (2--)3(--4)-veined, lanceolate to 
               elliptic, rarely long elliptic.............................................................................................................11. C. quinquenervis 
                                                                                                                                                                        (C. kweizhouensis, C. paucinervis)
        13b. Deciduous trees; leaves 4--9 (--16) cm long, (3--)4(-5) or 5--8-veined, 
               elliptic, ovate elliptic to broad ovate.
               14a. Leaves 5-8-veined................................................................................................................7. C. macrophylla
               14b. Leaves (3--)4--(5)-veined.
                      15a. Bark dark gray or dark brown; leaf base cunnate to broad cuneate, often 
                              oblique; flowers ca. 7-9.5 cm in diam.
                             16a. Abaxial surface of leaves grayish green, pubscent with fine, appressed 
                                     hairs, 4-veined; petioles 1.2--2.2 cm; infloresecences loose, spreading, with 
                                     branches often arched toward the center; calyx teeth longer than the disk..................6. C. koehneana
                             16b. Abaxial surface of leaves light green, pubescent with thicker appressed 
                                     hairs, rough to touch, (3)4--5-veined; petioles longer, (0.8--)3.5 cm; inflorescence 
                                     dense and compact; branches of inflorescence straight, rarely arched; calyx 
                                     teeth equal to flower disk.......................................................................................14. C. walteri 
                                                                                                                                                                        (C. yunnanensis)
                      15b. Bark light brown; leaf base rounded, rarely oblique; flowers smaller, ca 
                              5.2 cm in diam................................................................................................................4. C. coreana.
 
1. Cornus alba Linn. Mant. 1: 40, 1767.
Hong rue mu

Swida alba Opiz in Seznam, 94, 1852.

Deciduous spreading shrubs up to 3 m. Barks purplish red. Yong branches terete, first pubescent with whitish short hairs, later glabrous and covered by wax-like white powder. Old branches redish white, scattered with grayish white rounded lenticels; leaf-scar semicircular, conspicuous. Leaves paperous, 5--8.5 x 1.8--5.5 cm, elliptic or ovate elliptic, acute at apex, cuneate or broad-cuneate at base, entire or slightly revolute at margin, (4--)5(--6) veined; abaxial surface powdery green, pubescent with white appressed short hairs; axil of veins sometimes with a cluster of long brown hairs; small veins quite conspicuous. Corymbose cymes dense, ca. 3 cm wide, pubescent with white soft hairs. Flowers white or yellowish white, 6-8.2 mm in diam.; pedicels slender, 2-6.5 mm. Calyx lobe sharp triangular, 0.1--0.2 mm, shorter than the disk. Stamens longer than petals; anthers yellowish. Style cylindric, 2.1-2.5 mm; stigma disciform, wider than the style. Fruit oblong globose, slightly laterally compressed, ca. 8 mm long, 5.5-6 mm in diam., creamy white or bluish white at mature; stones subdiamond-shaped, laterally compressed, ca. 5 x 3mm, 3-ribbed on each side. Fl. Jun-Jul, fr. Aug-Oct.

In mixed thickets by streams, or in needle-broad leaved mixed forests at 600-1700(--2700 m). Gansu, Hebei, Heilongjian, Jilin, Jiangsu, Jiangxi, Liaoning, Nei Mongol, Qinghai, Shaanxi, Shandong [Korea, Russia, and Europe].

Seeds contain 30% oil for industrial uses; commonly culvated as garden ornamentals.

2. Cornus austrosinensis Fang & W. K. Hu, J. Sichuan Univ., Nat. Sci. ed. 3: 155, Pl. 1, 1-2, 1980.
Hua Nan Lai Mu

Swida (or Svida) austrosinensis (Fang & W. K. Hu) Fang & W. K. Hu.

Shrubs or small trees, 3--6 m. Yong branches terete, sparsely pubescent wih white fine hairs. Old branches yellowish brown, with oblong lenticels (?). Leaves opposite, rarely alternate, thin leathery, 4-8 x 2-4 cm, oblong elliptic, short acuminate at apex, broad cuneate at base, slightly revolute at margin; abaxial surface nearly glabrous and non-papillate, with 4-5 thin veins. Corymbose cymes 5-6 cm wide, sparsely pubescent. Flowers small, white, ca 7 mm in diam.; pedicels slender, 3-4 mm. Calyx teeth sharp triangular, ca. 0.3 mm, subequal to disk. Stamens slightly longer than petals; anthers linear elliptic. Style slender, cylindric, 3.3 mm. Stigma small, capitate, subequal to style in diam. Fruit globose, 5 mm in diam., black when mature; stones vertically compressed globose, 4 x 3 mm. Fl. Jun-July, fr. Dec.

In thickets at ca. 2500 m. Guangdong, Guangxi, Guizhou, Hunan.

There were few herbarium specimens for this species. The leaves were described as thick paperous in the species description, but as thick leathery in the key to species by Fang & W. K.Hu. According to the specimens we examined (including the type), the leaves appeared to be thin leathery. Living specimens were not observed.

3. Cornus bretschneideri L. Henry in Le Jardin, 13: 309, fig. 154, 1899.
Sha Lai

Swida bretschneideri (L. Henry) Sojak; Cornus aspera Wanger. in Fedde, Repert. Sp. Nov. 6: 97, 1908.

Shrubs or small trees 1--6 m. Barks purplish red. Young branches terete, redish, sparsely pubescent with grayish white hairs. Old branches light yellow, glabrous, with or without whitish elliptic lenticels. Leaves opposite, characeous, ovate, elliptic ovate, or oblong, 5-8.5 x 2.5-6 cm, cuspidate to acuminate at apex, broad cuneate to rounded at base; abaxial surface grayish white or powdery white, densely papillose and pubescent with appressed white hairs or occationally with dense yellow-brown curly hairs; lateral veins 5-6(--7), with white long soft hairs in axils; smaller veins inconspicuous. Corymbose cymes 4.5-6 cm wide, pubescent with grayish white, later deciduous short hairs, or rarely with dense curly hairs. Flowers small, white, 5.5-7 mm in diam.; pedicels 1.5-6 mm. Calyx teeth sharp triangular, 0.2-0.25 mm, equal to or slightly longer than the disk. Stamens longer than the petals; anthers yellowish white, ovate-oblong. Style cylindric, 2.3-2.6 mm. Stigma capitate, wider than style. Fruit subglobose, 4-5 mm in diam., bluish black to black; stones ovate globose, ca. 3.5 mm in diam., inconspicuously ribbed. Fl. Jun-Jul, fr. Aug.-Sept.

1a. Abaxial surface of leaves and inflorescence pubescent with short, non-curly hairs………………………..var. bretschneideri.
1b. Abaxial surface of leaves and inflorescence pubescent with dense curly hairs………............................... var. crispa.

3a. Cornus bretschneideri var. bretschneideri
Sha Lai Cornus bretschneideri var. gracilis Wanger. In Engl., Pflanzenreich, 41 (IV. 229): 67, 1910; Swida bretschneideri (L. Henry) Sojak var. gracilis.

Leaves pubesdent with short, appressed, non-curly hairs on abaxial surface. Infloresence pubescent with short, appressed, non-curly hairs.

Slope, thickets, forest; elev. 1100--2300 m. Gansu, Hebei, Henan, Hubei, Liaoning, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shanxi, Sichuan.

3b. Cornus bretschneideri var. crispa Fang & W.K. Hu, J. Sichuan Univ., Nat. Sci. ed. 3: 157, Pl. 3, fig. 1, 1980. Juen Mao Lai Mu

Swida bretschneideri var. crispa (Fang & W.K Hu) Fang & W.K . Hu.

Leaves densely pubesdent with long, curly, soft hairs. Infloresence densely pubescent with long, curly, soft hairs.

In sparse forests at elev. 580-1800 m. Gansu, Hebei, Heilongjiang, Jilin, Liaoning, Nei Mongol, Shaanxi, Shanxi.

4. Cornus coreana Wanger. in Fedde, Pepert Sp. Nov. 6: 99, 1908.
Korean Lai Mu
Swida coreana (Wanger.) Sojak.

Deciduous trees up to 20 m. Barks light brown, rectangular split. Yong branches more or less 4-angled, pubescent with brown short hairs. Old branches brown, glabrous. Leaves opposite, 1.2-2 cm-petiolate, paperous, long elliptic to broad ovate, 6--8 x 1.7-2.5(--4) cm; apex short acuminate; base rounded or broad cuneate, rarely oblique, 4--5-veined, sometimes revolute at margin; abaxial surface light green, with brown and grayish white, appressed short hairs, rough to touch; small veins reticular. Corymbose cymes 3.5-5 cm wide, with white and brown, short hairs. Flowers white, ca 5.2 mm in diam.; pedicels 2-3.2 mm. Calyx lobes lanceolate, unequal, 0.5-1.1 mm, longer than the disk. Stamens ca. 4.2 mm, equal to the petals; anthers oblong, color?. Style clavate, ca 3 mm. Stimga subcapitate, not wider than the style. Fruit globose, ca 5 mm in diam., black at ripening; stones ? (shape?). Fl. May--Jun, fr. Sept--Oct.

In forests. Elv. 0--300 m. Liaoning [Korea]

The specimens cited by Wangerin when he published the species are very similar to C. walteri, except that C. walter has larger flowers and darker color of bark. Its morphology seems overlaps with that of C. walteri. The status of the species may need further evaluation.

5. Cornus hemsleyi Schneid. & Wanger. In Fedde, Repert. Sp. Nov. 7: 229, 1909.
Hong Lian Zhi

Swida hemsleyi (Schneid. & Wanger.) Sojak; Cornus alsophila W. W. Smith, Notes Bot. Gard. Edinb. 10: 19, 1917; C. polyantha Fang & W. K. Hu, J. Sichuan Univ., Nat. Sci. ed. 3: 158, Pl. 4, fig. 2, 1980; Swida alsophila (W. W. Smith) Holub.

Shrubs or small trees 2-5 m. Barks redish brown, brown or dark gray. Young branches red or green first, later red, slightly 4-angled or rounded, glabrous or with appressed soft hairs. Old branches purple red to deep brown, glabrous, with yellowish brown elliptic lenticels. Leaves opposite, paperous, elliptic to ovate elliptic, or broad ovate, 4.5-9.3 (--13) x 1.8-4.8 (--6.2) cm, 6-8-veined, small veins conspicusly reticulate; apex acuminate or short acuminate; base rounded, broad cuneate to nearly cordate, sometimes slightly oblique; abaxial surface more or less rough to touch, densely papillate, pubescent with dense to sparse appressed, short, white hairs; axils of veins sometimes with grayish and/or light brown long hairs. Corymbose cymes flat to convex, 5-9 cm wide, pubescent with light brown short hairs or nearly glabrous after anthesis. Flowers white to yellowish, small, 6-7 (--8.5) mm in diam.; pedicels 1-5.5 mm. Calyx lobes triangular, 0.4-1 mm, equal to or longer than the disk. Stamens longer than or rarely equal to petals; anthers ovate oblong, grayish blue to grayish white, or yellowish. Style cylindric or slightly expanded at apex, but not clavate, 1.8-4 mm. Stigma subcapitate, wider than the style, slightly 0--4-lobed. Fruit globose, 4--5 mm in diam. Purple red to Black; stones globose, vertically compressed, ca 2.3-3.1 x 1.8-2.5 mm, inconspicuously 8-ribbed. Fl. Jun-Jul, fr. Aug-Sept.

In thickets, mixed forests, by streams. Elv. 1000--3980 m. Gansu, Guizhou, Hebei, Henan, Qinghai, Shaanxi, Shanxi, Sichuan, Yunnan, Xizhang.

Oil extracts from seeds are used for soap industry. Leaves and barks are used as source of industrial tannin.

6. Cornus koehneana Wanger. in Fedde, Pepert Sp. Nov. 6: 99, 1908.
Chuan Xia Lai Mu
Swida koehneana (Wanger.) Sojak.

Trees 6-10 m. Barks dark brown. Young branches slender, more or less 4-angled (or ridged), purple-red green, pubescent with grayish appressed hairs. Old branches yellow-brown to gray-brown, glabrous, sparsely scattered with yellowish white long elliptic lenticels. Leave opposite, 1.2-2.2 cm-petiolate, thick paperous, long elliptic to ovate elliptic, 3.3-8 x 2.3-4.5 cm; apex acuminate or extended acuminate; base cuneate or broad cuneate, slightly oblique, 4-veined; abaxial surface grayish green, with grayish white, thin, appressed short hairs; small veins inconspicuously reticular. Corymbose cymes loose and spreading, ca 6 cm wide, with branches slightly arched inward (bent toward the center), pubescent with grayish appressed hairs. Flowers white, ca 7 mm in diam.; pedicels 0.5-3 mm. Calyx lobes triangular, 0.4 mm, longer than the disk. Stamens ca. 4 mm, shortern than the petals; anthers light yellow, oblong ovate. Style clavate, ca 3.8 mm. Stimga capitate, not wider than the style. Fruit ovate globose, ca 5 x 6 mm; stones subovoid, 4 x 5 mm. Fl. May-Jun, fr. Jul-Aug.

In mixed forest in valley or slopes; elev. 1700-2200 m. Gansu, Shaanxi, Shanxi, Sichuan. Specimens of the species are sparse. They appear to be very similar to C. walteri. Examination of more specimens, especially living specimens, of the species are needed to determine its relationship to C. walteri.

7. Cornus macrophylla Wall in Roxb., Fl. Ind., ed. Carey & Wallich, 1: 431. 1820.
Lai Mu

Cornus brachypoda C. A. Meyer, Mem. Acad. Petersb. 7: 223. 1844; C. crispula Hance, J. Bot. xix. 216. 1881; C. corynostylis Koehne in Gartenfl. 45: 286. Pl. 51, fig. 4 A-B. 1896; C. taiwaniensis Kenehira, Formosan Trees, 282. 1917; C. longipedunculata Fang & W. K. Hu, in Fl. Sichuan 1: 344. Pl. 131, fig. 1-3. Addenda: 469. 1981; C. alpina Fang & W. K. Hu, J. Sichuan Univ., Nat. Sci. ed. 3: 160. PL. 6. fig. 2. 1980; Swida macrophylla (Wall.) Sojak; S. macrophylla var. longipedunculata (Fang & W. K Hu) Fang & W. K. Hu; S. alpina (Fang & W.K. Hu) Fang & W. K. Hu.

Trees 2--15 (--25) m, rarely shrubs. Barks gray-brown or gray-black, smooth. Young branches stout, more or less 4-angled, early sparsely pubescent with grayish short hairs, later glabrous. Old branches dark brown, with yellowish white to grayish elliptic to rounded lenticels and semicircular leaf scars. Leaves opposite, 1.5-3 cm-petiolate, paperous, broad elliptic, broad ovate or ovate oblong, rarely elliptic, 9-16(--18) x 3.5-8.8 cm; apex acute or short acuminate; base rounded, broadly cuneate, rarely oblique, 5--8-veined; abaxial surface light green to slightly whitish, papillate, with grayish white, appressed short, later deciduous hairs; small veins conspicuous. Paniculate, sometimes corymbose cymes 8-12 cm wide, sparsely or rarely densely pubescent with elevated hairs; penduncles sometimes redish. Flowers white, fragrant, ca (7--)8--10 mm in diam.; pedicels 0.3-4.5 mm. Calyx lobes triangular or broad triangular, 0.4-0.5 mm or rarely 0.8--1.0 mm, longer (or taller) than the disk. Stamens equal or slightly longer than the petals; anthers onblong or obovate oblong, yellow (?) or rarely blue. Style cylindric, slightly expanded at apex to apparent clavate, 2--4 mm. Stimga subcapitate, wider than the style, slightly lobed. Fruit subglobose, ca 4.5--6 in diam., purple black to blue black; stones compressed globose, (2.6--)3-4 mm in diam., 6 or 8-ribbed. Fl. Jun-Jul (Aug), fr. Aug-Sept (Oct).
Slopes, by streams, or in dense forests. Elev. 5-3600 m. Gansu, Shaanxi, Shandong, Taiwan, Xizang, and provinces of south of the Yangtzi River [Pakistan, Myanmar, India, Bhutan, Sikkim, Nepal, Afghanistan].

7a. Inflorescence sparsely pubescent with appressed hairs; anthesis June to July; flowers larger 8-10 mm in diam; fruit stones 3-4 mm in diam…………………………...............................................var. macrophylla
7b. Inflorescence densely pubescent with elevated hairs; anthesis July to August; flowers smaller, 7-8 mm in diam; fruit stones smaller, ca 2.6 mm in diam………………………………………………………...var. stracheyi

7a. Cornus macrophylla var. macrophylla
Lai Mu

Trees 3-15 (--25) m. Old branches with grayish white elliptic lenticels. Inflorescence sparsely pubescent with white to yellowish appressed hairs. Flowers 8-10 mm in diam. Calyx lobes broad triangular, 0.4-0.5 mm, slightly longer than the disk. Style thickened at apex or sometimes typical clavate. Fruit stone 3-4 mm in diam, 6-ribbed. Fl. Jun-Jul, fr. Aug.-Sept.

Slopes, by streams, or dense forests. Elev. 5-3600 m. Gansu, Shaanxi, Shandong, Taiwan, Xizhang, and provinces of south of the Yangtzi river [Pakistan, Myanmar, India, Bhutan, Sikkim, Nepal, Afghanistan].

7b. Cornus macrophylla var. stracheyi C B. Clarke in Hook. F. Fl. Brit. Ind. 2: 744. 1879.
Mi Mao Lai Mu
Cornus stracheyi (C. B. Clarke) Hemsl. Kew Bull. 1909: 333. 1909; Swida stracheyi (C. B. Clarke) Sojak in Bovit. Bot. & Del. Sem. Hort. Bot. Univ. Carol. Prag. 1960: 10. 1960.

Trees or shrubs, 3-9 m. Old branches with yellowish white rounded lenticels. Inflorescence densely pubescent with brownish or brownish and white hairs. Flowers small, 7-8 mm in diam. Calyx lobes triangular, unequal, 0.8-1, longer than the disk. Style slightly thickened at apex. Fruit stones smaller, ca. 2.6 mm, 8-ribbed. Fl Jul-Aug, fr. Sept-Oct.
In mixed woods or on the edge of woods. Elev. 1700-3400 m. Yunnan, Xizang [India, Nepel].

8. Cornus oligophlebia Merr. J. Arn. Arb. 23; 187. (Rec. Indochin. Pl. 3). 1942.
Zhang Ye Lai Mu
Swida oligophlebia (Merr.) W. K. Hu, Bull. Bot. Res. 4: 102. 1984.

Trees 6-10 m. Young branches green, more or less pubescent with brown hairs. Old branches dark brown. Leaves opposite, rarely alternate, 1.1--15 cm-petiolate, leathery, broad elliptic to ovate, 8-11 x 4-4.5 cm; apex acuminate; base broadly cuneate or rounded, sometimes oblique; blade slightly revolute at margin, 3(-4)-veined; abaxial surface light green, not papillate, sparsely pubescent with white short hairs or nearly glabrous; small veins conspicuous horizontal. Paniculate cymes short, large, spreading, 8.5-11cm wide, glabrous or with appressed gray hairs. Flowers white, ca 9 mm in diam.; pedicels 1--2 mm. Calyx lobes broad triangular, slightly longer (or taller) than to the disk. Stamens shorter to longer than the petals; anthers narrow obovate. Style cylindric, 3 mm. Stimga small, flat, punciform. Fruit subglobose, ca 3--4 mm in diam., purple black to blue black; stones compressed globose, (2.6--)3-4 mm in diam. Fl. Sept., fr. Jan.

In forests. Elev. 1200-1500 m. SE Yunnan [Vietnan, Myanmar, Thailand, India, Bhutan, northwest Himalaya].
The species appears to be rare in China, with a restricted distribution in Yunnan province and adjacent countries. The species stands out from the rest of the subgenus by a combination of features including leathery leaves, large, short, spreading paniculate inflorescences, punciform stigma, sometimes subopposite leaves, a flowering time in fall and fruiting time in winter. These features suggest affinity to C. oblonga. The species thus may represent an relatively old lineage in subg. Kraniopsis or actually a member of subg. Yinquania. Both are restricted to SW China. Phylogenetic analyses will help to test these hypotheses.

9. Cornus papillosa Fang et W.K Hu in Flora of Sichuan, 1: 330, Pl. 125, figs. 1-4, Addenda: 468, 1981.
Ru Tu Mao Lai
Swida papillosa (Fang & W. K. Hu) Fang & E. K. Hu; Swida polyantha (Fang & W.K. Hu) Fang & W.K.Huv

Small trees ca 5 m. Yong branches light green, with sparse yellowish white soft hairs. Old branches deep brown, glabrous, with rounded or elliptic lenticels. Leaves opposite, paperous, elliptic to oblong elliptic, 8--12 x 4-7 cm, 7-9 veined, small veins conspicuous; apex more or less cuspidate; base curneate or broad cuneate; abaxial surface gray, with dense papillae, blackish streaks, and appressed (or flat) hairs; axils of veins often clustered with light yellow long hairs. Corymbose 5-9 cm wide, sparsely pubescent with light yellowish white, short hairs. Flowers white, ca. 7 mm in diam.; pedicels 1-5.5 mm. Calyx lobes ovate to linear triangular, 0.6-0.8 mm, equal to or longer than the disk. Stamens longer than petals; anthers ovate oblong, yellow. Style cylindric, sligtly expanded at apex, 3.4 mm. Stigma compressed head, wider than the style, slightly 3--4-lobed. Fruit globose, ca 3 mm in diam. black; stones globose, vertically compressed, 2.7 x 1.8 mm. Fl. Jun, fr. unknown.
In forests. Evl. Ca 3000 m. Sichuan, Yunnan.

10. Cornus parviflora Chien in Sinensia 2: 99, 1931.
Xiao Hua Lai Mu
Swida parviflora (Chien) Holub.

Trees or shrubs, 3--8 m. Barks yellow brown. Yong branches slender, 4-angled, with sparse grayish hairs. Old branches grayish brown, with sparse yellow brown lenticels. Leaves opposite, paperous, long elliptic to elliptic lanceolate, 4-6.5 x 1.6-3.3 cm, 3-4- veined; apex acuminate to more or less caudate; base curneate or broad cuneate; abaxial surface light green, non-papillate, with sparse white, appressed short hairs. Corymbose cymes 4--12 cm wide, pubescent with light yellowish white, short hairs, occationally with a lanceolate or ovate lanceotate, leaf-like bract; internodes of inflorescence thin, long. Flowers white, small, ca 4.5 mm in diam.; pedicels 0.3--2 mm. Calyx lobes broad triangular 0.3-0.4 mm, slightly longer than the disk. Stamens shorter than petals; anthers narrow obovate. Style cylindric, ca 2 mm. Stigma small, punciform. Fruit obovate or nearly oblong, ca 5-6 long, 4 mm in diam. Fl. Jul, fr. Aug--Spet.
In dense or sparse forests or open hillside. Evl. 330--2500 m. Guizhou, Guangxi.

11. Cornus quinquenervis Franch., J. Bot. 10: 307. 1896.
Xiao Lai Mu
Cornus paucinervis Hance, J. Bot. n.s. 10: 217. 1881; Swida paucinervis (Hance) Sojak.

Shrubs, 1-3(--4) m. Barks blackish gray, smooth. Yong branches green or purplish red, 4-angled, with grayish short hairs. Old branches brown, glabrous. Leaves opposite, chatarceous, elliptic lanceolate, lanceolate, rarely oblong ovate, 4-9 (--10) cm x 1-2.3(--3.8) cm, (2--) 3(-4)- veined; veins nearly straight and ascending below, arched inward near the leaf margin; apex acuminate to obtuse acummiate; base curneate; abaxial surface light green, with sparse, white, appressed, later deciduous, short hairs. Corymbose cymes 3.5--8 cm wide, pubescent with white, appressed short hairs. Flowers white to yellowish white, small, ca 9--10 mm in diam.; pedicels 2--9 mm. Calyx lobes lanceolate triangular to acute triangular, ca 1 mm, conspicuously longer than the disk. Stamens shorter than petals; anthers oblong ovate, yellowish white. Style conspicuously clavate, ca 3.5 mm. Stigma small, punciform or truncate. Fruit rounded globose, ca 5 in diam., black when ripe; stones subglobose, ca 4 mm in diam., inconspicuously 6-ribbed. Fl. Jun--Jul, fr. Oct--Nov.
Hillside, mountain forests, thickets by streams. Elev. 50-2500 m. Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangsu, Shaanxi, Sichuan, Yunnan.

Cornus paucinervis Hance has long been used as the name of this species. However, this name has two earlier hononames (i.e., this name had been used by two paleobotanists and published before Hance: Heer, in Fl. Tertiaria Helvetiae. 1859; Ettingshausen, in Die Fossile Flora der Alteren Braunkohlenformation der Wetterau. Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Cl., Abt. 1, 57: 807. Pl.3. 1868. See Eyde, Bot. Rev. 54: 268-269. 1988). Thus, the name becomes invalid for the species. The next properly published name is Cornus quinquenervis which becomes the valid name for the species.

12. Cornus schindleri Wanger. In Fedde, Repert, Sp. Nov. 4: 337. 1907.
Kang Ding Lai Mu
Cornus monbeigii Hemsl., Kew Bull. 333. 1909; C. xanthotricha Fang & W. K. Hu, J. Sichuan Univ., Nat. Sci. ed. 3: 159. Pl. 6, fig. 1. 1980; C. monbeigii Hemsl. Subsp. crassa Fang & W. K. Hu, J. Sichuan Univ., Nat. Sci. ed. 3: 158. Pl. 5, fig. 1, 1980; C. monbeigii Hemsl. Subsp. poplifolia Fang & W. K Hu, J. Sichuan Univ., Nat. Sci. ed. 3: 158. Pl. 5, fig. 2. 1980; C. lixinensis Fang & W. K. Hu, Fl. Sichuan 1: 339. Pl. 129, fig. 1-3. Addenda: 468. 1981; C. poliophylla Schneid. & Wanger. In Fedde, Pepert. Sp. Nov. 7: 228. 1909; C. malifolia Fang & W. K. Hu, in Fl. Sichuan 1: 335. Pl 128, fig. 6-8. Addenda: 468. 1981; Cornus poliophylla var. praelonga Fang & W. K. Hu, in Fl. Sichuan 1: 341. Pl. 130. fig. 7-8. Addenda: 469. 1981; C. daijinensis Fang & W. K. Hu, in Fl. Sichuan 1: 341. Pl. 129. fig. 4-6. Addenda: 469. 1981; C. scabrida Franch. In Nouv. Arch. Mus. Paris 3(7): 250. 1885; Swida monbeigii (Hemsl.)Sojak; S. monbeigii (Hemsl.) Sojak var. xanthotricha (Fang et W. K Hu) Fang & W. K. Hu; S. monbeigii (Hemsl.) Sojak var. crassa (Fang & W. K. Hu) Fang & W. K. Hu; S. monbeigii (Hemsl.) Sojak var. poppulifolia (Fang & W. K. Hu) Fang & W. K. Hu; S. schindleric (Wanger.) Sojak; S. schindleri (Wanger.) Sojak vr. Lixianensis (Fang & W. K. Hu); S. poliophylla (Schneid. & Wanger.) Sojak; S. poliophylla (Schneid. & Wanger.) Sojark var. malifolia (Fang et W. K. Hu) Fang & W. K. Hu; S. poliophylla (Schneid. & Wanger.) Sojak var. praelonga (Fang & W. K. Hu) Fang & W. K. Hu; S. daijinensis (Fang & W. K. Hu) Fang & W. K. Hu; S. scabrida (Franch.) Holub.

Trees or shrubs, 2--8 (--10) m. Barks brown. Yong branches 4-angled, densely pubescent with brown or grayish hairs, rarely glabrous. Old branches redish to grayish purple-brown, glabrous, with sparse, whitish, rounded or elliptic lenticels. Leaves opposite, paperous to thick paperous, elliptic, ovate elliptic to broad ovate, rarely long elliptic or rounded; elliptic lanceolate, 4-11 (--15) x 2.5-6.5 (--8) cm, 6-8 (--9)-veined; apex acuminate or short accuminate; base cuneate to rounded, cordate, rarely slightly oblique; abaxial surface light green to grayish green, often papillate and conspicuously pubescent with whitish to brownish curly, spreading hairs on veins or on the entire surface. Corymbose cymes (5--)6-10 cm wide, densely pubescent with yellowish to brown (rarely rusty-red), later often deciduous curly hairs. Flowers white, ca (6--)7-8 mm in diam.; pedicels 1-5(--6) mm. Calyx lobes triangular, rarely lanceolate, (0.2) 0.3-0.5 mm, longer, rarely shorter, than the disk. Stamens equal or longer than petals; anthers oblong to narrow oblong, light blue to gray, yellow. Style cylindric, ca 2--3(--4) mm. Stigma capitate to disciform, wider than the style, often slightly lobed. Fruit subglobose, purple redish black, black when ripe, ca 4--6 mm in diam; stones compressed globose or ovate, 3--4 mm x 2.8--3.5 mm, conspicuously or inconspicuously 8-ribbed, rarely 10-ribbed in ripe fruits. Fl. May--Jul, fr. Aug.--Oct.
In dense or sparse forests or open hillside. Evl. 1100--3200 m. Gansu, Henan, Hubei, Shaanxi, Sichuan, Yunnan, Xizang.
The species shows somewhat continuous morphological variation across its distributional range. In particular, the ratio of filament length to petal length show a east-north-west-south pattern, with a higher ratio in the east and lower ratio in the west. Due to this association with geography, two subspecies are recognized (Xiang in Bull. Bot. Resear. 9: 125. 1989).

1a. Midrib of leaves with scattered, appressed brown hairs beneath; mature flower buds subrotund; ratio of length of filaments to length of petals is equal or greater than 1.4; hairs on surface of ovary usu. appressed ………......................subsp. poliophylla.

1b. Midrib of leaves with curly (crisped) yellowish to yellow hairs beneath; mature flower buds conical; ratio of length of filaments to length of petals is less than 1.4; hairs on the surface of ovary usu. patent……...........................................subsp. schindleri.

12a. Cornus schindleri subsp. schindleri
Kang Ding Lai Mu
Midveins of leaves on abaxial surface pubescent with yellowish to yellow crisped hairs. Flower buds conical when mature. Filaments are 1.4 times shorter than the petals. Ovary pubescent with patent (spreading) hairs.

S & W Sichuan, NE & NW Yunnan, SE Xizang.

12b. Cornus schindleri subsp. poliophylla (Schneid. & Wanger.) Xiang
Hui Ye Lai Mu

Midveins of leaves on abaxial surface pubescent with brown, appressed hairs. Flower buds nearly rounded when mature. Filaments are 1.4 times as long as or longer than the petals. Ovary pubescent with appressed hairs.

SE Gansu, Henan, W Hubei, S Shaanxi, NE Sichuan,

13. Cornus ulotricha Schneider. & Wanger. In Fedde, Repert., Sp. Nov. 7: 228. 1909.
Juen Mao Lai Mu

Swida ulotricha (Schneid. & Wanger.) Sojak var. leptophylla W. K Hu, Bull. Bot. Res. 4(3): 106. 1984; S. ulotricha (Schneid. & Wanger.) Sojak.

Trees, rarely shrubs, (1.5--)7-15(--20) m. Barks rough, gary, rectangulary split. Yong branches redish brown, sparsely pubescent with short hairs. Old branches yellowish brown, glabrous. Leaves opposite, chataceous, broad ovate to broad elliptic, 1.5-2.8 cm petiolate, 9--15 x 3-8.5 cm, 6--7-veined; apex short accuminate; base rounded, sometimes slightly oblique; abaxial surface grayish, non-papillate or only inconspicuously papillate, pubescent with both whitish appressed and brown elevated (V-shaped) or curly hairs; hairs often deciduous; curly hairs on veins dense. Corymbose cymes, broad, 8-12 cm wide, pubescent with a mixture of grayish appressed short hairs and slightly curly hairs; lateral branches arched inward; smaller branches nearly all oneside, toward the center of the inflorescence. Flowers white, ca 6-8 mm in diam.; pedicels, thick, 0.2--2 mm. Calyx lobes broad triangular, 0.3-0.4 mm, shorter than or equal to the disk. Stamens shortern than or equal to petals; anthers oblong ovate, yellow. Style cylindric, ca 3 mm. Stigma semiglobose to disciform, wider than the style. Fruit subglobose, bluish black, ca 4.3-4.5 mm in diam; stones subglobose, 3--4 mm in diam. Fl. May--Jun, fr. Jul.-Aug.
In mixed forests, by streams. Evl. 850-2650 m. Gansu, Guizhou, Henan, Hubei, Shaanxi, Sichuan, Yunnan, Xizang.

14. Cornus walteri Wanger. in Fedde, Pepert Sp. Nov. 6: 99, 1908.
Mao Lai
Cornus yunnanensis L, J. Arn. Arb. 25: 312, 1944. C. walteri Wanger. Var. insignis Fang & W. K.Hu, Fl. Sichuan 1: 349, Addenda: 469, 1981; C. walteri var. confertiflora Fang & W.K Hu, in Fl. Sichuan 1: 349, Pl 132, fig. 5-7, Addenda, 470, 1981; Swida walteri (Wanger.) Sojak; S. wlateri var. insignis (Fang & W. K. Hu) Fang & W.K. Hu; S. walteri var. confertiflora (Fang & W.K.Hu) Fang & W.K. Hu.

Trees 6-15 m. Barks dark gray, rectagularly split. Yong branches more or less 4-angled, green, densely pubescent with grayish white short hairs. Old branches yellow-green, glabrous. Leave opposite, (0.8--) 3.5 cm-petiolate, paperous, long elliptic to broad ovate, 4-12 (--15) x 1.7-5.5(--8) cm; apex short to long acuminate; base cuneate, rarely rounded or cordate, often oblique, 4 (--5)-veined; abaxial surface light green, with grayish white, appressed short hairs, rough to touch; small veins inconspicuously reticular. Corymbose cymes dense 7--9 cm wide, with white, short hairs. Flowers white, fragrant, ca 9.5 mm in diam.; pedicels 0.8-2.7 mm. Calyx lobes triangular, 0.4 mm, equal to the disk. Stamens ca. 4.8-5 mm, equal or longer than the petals; anthers light yellow, rarely rosy red, oblong ovate. Style clavate, ca 3.5 mm. Stimga capitate, not wider than the style. Fruit globose, ca 6-7(--8) in diam., black; stones compressed globose, 5 x 4 mm, inconspicuously ribbed. Fl. May--Jun, fr. Aug--Oct.

In mixed sparse or dense forests. Elv. 300-2500(--3000) m. Hebei, Liaoning, Shaanxi, all provinces of eastern China, Central China, southern China, and southwestern China.

Furits are source of oil; wood is hard and used for tools. May be used for street ornamental.

15. Cornus wilsoniana Wanger. In Fedde, Repert., Sp. Nov. 6: 97. 1908.

Guang Pi Lai Mu
Cornus fordii Hemsl. In Kew Bull. 334. 1909. C. kweichouensis Li, J. Arn. Arb. 25: 311, 1944; Swida wilsoniana (Wanger.) Sojak.

Trees 5-18 m, rarely up to 40 m. Barks gray to greenish gray, rectagularly split. Yong branches grayish green, more or less 4-angled, pubescent with grayish, appresshed short hairs. Old branches brown, glabrous, with brown, oblong elliptic lenticels. Leave opposite, 0.8--2 cm-petiolate, paperous, elliptic or ovate elliptic, 6-12 x 2-5.5 cm; apex shortly acuminate to accuminate; base cuneate to broadly cuneate, slightly revolute at margin, 3--4-veined; abaxial surface grayish green, densely pubecent with white, appressed short hairs, and papillae, rough to touch. Paniculate to corymbose cymes 6--10 cm wide, with white, short hairs. Flowers white, ca 7 mm in diam.; Calyx lobes triangular, 0.4 -0.5 mm, longer than the disk. Stamens ca.6-6.8 mm, equal to the petals; anthers yellow, narrow oblong. Style cylindric, sometimes slightly expaned near the apex, 3.5--4 mm. Stimga small, disciform, not wider than the style. Fruit globose, ca 6--7 in diam., purple black to black; stones globose, 4-4.5 mm in diam, ribs inconspicuous. Fl. May, fr. Sept--Nov.

In forests. Elv. 130-1130 m. Fujian, Gansu, Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangxi, Shaanxi, Sichuan, Zhejiang.

Furits are source of oil (up to 30% oil); leaves were used for livestock feeding. Wood is dense and used for farming tools and furniture. Crown is nicely shaped and good candidates for street ornamental.

4. Subg. Cornus L.

Shan Zhu Yu Ya Shu

Cornus sect. Macrocarpium Spach. Hist. Nat. Veg. 8:101. 1939; Cornus subg. Sinocornus Q. Y. Xiang, Acta Phytotax. Sin. 25:128. 1987; C. sect. Cornotypus Dumortier. Florula Belgica 83: 1827; C. subg. Eukrania Raf. Alsograph. Am. 59. 1838; Cornus b. Tanycrania Endlicher. Gen.Pl. 798. March. 1839; C. subg. Tanycrania Endlicher ex Reichenbach. Repert. Herb. 143. 1841; C. subg. Macrocarpium (Spach) Schneider; Macrocarpium (Spach) Nakai; C. sect. Tanycrania (Endlicher ex Reichenbach) Ledebour.

Deciduous small trees or shrubs. Flower buds terminal or axillary, ovate globose; pubescent with gray or brown, appressed hairs. Leaves paperous, opposite, ovate, elliptic, or ovate lanceolate, petiolate; adaxial surface glabrous or nearly glabrous; abaxial surface pubescent with appressed hairs or sometimes tomentosa. Inflorescences umbellate cymes, terminal or axillary, blooming before leaves expanded; subtended by 4 decussate, scale-like bracts. Bracts deciduous after anthesis; outer pair of bracts larger than the inner pair. Calyx tube cup-like, 4-dentate. Petals yellow, sublanceolate. Filaments awn-shape. Anthers oblong, 2-locular. Ovary 1 or 2-locular. Style short, cylindric. Stigma truncate. Fruits long elliptic to oblong, red to black; stones long elliptic, 1 or 2--seeded.

4 species in Europe, eastern Asia, and western North America. 2 species in China.

1a. Branches monopodial; inflorescence axillary; peduncles short, 2-3 mm; fruits 1.2-1.8 cm; lower surface of leaf with clusters of light brown hairs in axils of lateral veins..........................................................1. C. officinalis
1b. Branches sympodial; inflorescence terminal; peduncles longer, 5-12 mm; fruits smaller, 6-8(-10) mm; lower surface of leaf with clusters of grayish hairs in axils of lateral veins................................................2. C. chinensis

1. Cornus officinalis Sieb. et Zucc., Fl. Jap. 1:100. t. 50. 1835.
Shan Zhu Yu

Macrocarpium officinale (Sieb. et Zucc.),

Trees or shrubs 4--10 m; axis sympodial. Bark gray-brown. Leaf winter buds terminal or axillary, alone or associated with flower buds. Flower buds terminal, pubescent with yellow brown short hairs. Leaves ovate lanceolate or ovate elliptic, 5.5-10 x 2.5-4.5 cm; 6-7-veined; baxial surface light green, sparsely pubescent with short appressed hairs; axils of lateral veins with dense, light brown, long soft hairs. Umbellate inforescences terminal; bracts ovate, chataceous to leathery, ca 5--8 mm long, pubescent; peduncles short and thick, ca 2 mm, pubescent. Pedicles slender, 0.5-1.0 cm, densely pubescent with soft hairs. Calyx teeth borad trangular, ~0.6 mm. Petals ligular-lanceolate, 3.3 mm, reflexed. Stamens 1.8 mm; anthers elliptic. Ovary obovate, 1 mm, densely pubescent. Style ca 1.5 mm. Fruits long elliptic, 1.2-1.8 x 0.5--0.7 cm., red to purple red; stones narrow elliptic, 1.2 cm with a few unequal ribs. Fl. Mar-Apr; fr. Sept-Oct.

On mountains, slopes, edge of forests or in forests at 400--1500 (--2100) m. Anhui, Gansu, Henan, Hunan, Jiangsu, Jiangxi, Shaanxi, Shandong, Shanxi, Zhejiang [Korea, Japan].

Fruits are called "Zhu Yu" or "Zao Pi" in Chinese medicine, prescribed as an astringent tonic in impotence, spermatorrhea, lumbago, vertigo, night sweats.

2. Cornus chinensis Wanger. in Fedde, Repert. Sp. Nov. 6:100. 1908 et in Engl., Pflanzenreich, 41 (IV. 220):80. 1910.
Hua Zhu Yu
Macrocarpium chinense forma longipedunculatum Fang et W. K. Hu, J. Sichuan Univ., Nat. Sci. ed. 3:160. Pl. 7.1. 1980; M. chinense forma microcarpum W. K. Hu., in Fl. Sichuan 1:354, Pl. 136. Addenda: 470. 1981; M. chinense forma jinyangense W. K. Hu, in Fl. Sichuan 1: 354. Pl. 136. Addenda: 470. 1981; Cornus chinensis Wanger. forma microcarpa (W.K. Hu) W. K. Hu; C. chinensis forma jinyangensis (W. K. Hu) W. K. Hu; C. chinensis forma longipedunculata (Fang et W.K.Hu) Fang et W.K. Hu; Macrocarpium chinense (Wanger.) Hutch.

Trees 4-8 m; axis monopodial; bark dark brown. Leaf winter buds terminal or axillary, small, narrow cornicle. Flower buds in pairs, lateral, separated by a terminal leaf bud in the center, subglobose, acute at apex, pubescent with yellow brown hairs. Leaves ovate lanceolate to long elliptic, 6--11 x 2.8--5.5 cm, 5-6 veined; abaxual surface sparsely pubescent with grayish white, appressed hairs and a cluster of conspicuous gray long hairs in axils of veins, or sometimes toomentosa. Umbellate inflorescences lateral; bracts paperous to leathery, broad ovate to elliptic, 6.5-7 mm in length, pubescent with appressed hairs on both sides; peduncles purplish brown, 5--12 mm., more or less pubescent. Calyx triangular lanceolate, 0.7 mm. Petals lanceolate, 4 mm in length. Stamens 1.6 mm; anthers subglobose. Ovary bell-shape, ca 1 mm, pubescent with gray short hairs. Style 1--1.4 mm, glabrous. Pedicels slender, 8--9 mm, pubescent with long, yellow hiars. Fruits long elliptic, 6--8(--10) x 3.4--4 mm, purple brown to black; stones long elliptic, ca 7.5 mm with a few ribs. Fl. Apr; fr. Sept.

Slope, edge of mixed forests or in dense forests, Elev. 750--2500 (--3500) m. Gansu, Guangdong, Guizhou, Henan, Hubei, Shaanxi, Sichuan, Yunnan, Xizhang, Zhejiang.

Fruits have similar medicinal uses as C. officinalis.

5. Subg. Syncarpea (Nakai) Xiang, Act. Phytotax. Sin. 25:128. 1987.

Si Zhao Hua Ya Shu

Benthamia Lindl., Bot. Reg. 19: tab. 1579. 1833; Cornus sect. Cephalocrania Hance, Journ. Linn. Soc. 13:105. 1873, nom. tent.; Cornus sect. Benthamia C.B. Clarke in Hook. f., Fl. Brit. India 2: 745, 1879; Benthamia subg. Syncarpea Nakai, Bot. Mag.Tokyo 23: 41, 1909. March; Dendrobenthamia Hutch., Ann. Bot. N. S. VI(21): 92, 1942; Benthamidia Spach. Hist. Nat. Veg. 8: 92: 1839; Cornus subg. Benthamidia (Spach) K. Koch; Benthamidia sect. Cephalocrania (Hance) Hara; Cornus subg. Benthamia (C. B. Clarke) Schneider.

Evergreen or deciduous small trees or shrubs. Flower buds always terminal, globose to ovate, exposed or covered by scales. Leaf buds axillary or terminal, covered by a scale or exposed. Leaves opposite, petiolate, leathery or subleathery to paperous; blade ovate, elliptic or oblong lanceolate. Inflorescences capitulate cymes, terminal; bracts 4, white to light yellowish. Calyx tubular, slightly to conspicuously 4-lobed. Petals white to yellowish. Filaments slender; anthers elliptic, rarely ovate elliptic, yellow, brown, blue black. Ovary 2-locular. Style cylindric, often ridged and pubescent. Stigma truncate to capitate. Fruits fused into a compound, multiple stoned berry, red orange to red, globose or semi-globose; stones asymmatric, 1-seeded.

The species delimitation in the subgenus is controversial. Over 15 new taxa have been published since 1950s. Xiang (1987, Bull. Bot. Res. 7(2): 33-52) recognized 13 subspecies among four species, C. capitata, C. honghongensis, C. kousa, and C. multinervosa. We follow this treatment. However, preliminary allozyme investigation by Dudley and Santamour (1994,
Phytologia, 77: 425-430) suggested that C. capitata subsp. capitata is quite divergent from C. capitata subsp. angustata. Thus, in the present treatment, we recognize C. angustata (Chun) T.R. Dudley as a species separate from C. capitata. Further studies are needed to test the various species delimitations in the group.
5 species from Himalaya to eastern Asia. All in China. Gangsu, Henan , Nei Mongol, Shaanxi, Shanxi, and all provinces of the Yangtzi River [Nepal, Bhutan, northeastern India, Mong Jia La Guo, Korea, Japan, Loas and Veitnam].

Wood is hard, used for tools; mature fruits edeble with a sweet taste, used for winary or sold as wild fruits in free market of villages; excellent garden ornamentals due to their showy bracts.

  1a. Evergreen small trees; leaves subleathery, leathery, or thick leathery; 
        inflorescence buds exposed in winter.
        2a. Mature leaves grayish below; abaxial surface typically pubescent with short, 
              light gray or white hairs and asperous, rarely smooth or tomentosa; hairs appressed, 
              persistent, rarely deciduous.
              3a. Peduncles short and thick, 2-3(-5) cm; fruitresences compressed globose....... 1. C. capitata
              3b. Peduncles slender, 5-8(-10) cm; fruitrescences globose..................................2. C. angustata 
        2b. Leaves yellowish or pale green below; abaxial surface glabrous or more or 
              less pubescent with brown hairs, smooth to touch; hairs fine, appressed, deciduous, 
              leaving brown residuous dots on leaf surface after falling.......................................3. C. hongkongensis
  1b. Deciduous small trees; leaves paperous; inflorescence buds completely covered 
        by two pairs of decussate, pubescent scales; buds mixed.
              3a. Veins 3-4(5), curved inward, extending upward; lower ones not reaching 
                    the apex; base of peduncles conspicuously ring-like thickened; anthers light 
                    yellow to dark blue?; fruit stones light yellow, without red spots......................4. C. kousa
              3b. Veins 5-6 (7), curved inward, all extending almost to the apex; base of 
                    peduncles not conspicuously thickened; anthers dark blue; fruit stones scattered 
                    with red spots..............................................................................................5. C. multivervosa
  
1. Cornus angustata (Chun) T.R. Dudley
Jien Ye Si Zao Hua
Cornus kousa Buerg var. angustata Chun In Sunyatsenia 1: 285, 1934;Cynoxylon elliptica Pojark. In Notul. Syst. Inst. Bot. Nom. Kom. Acad. Sci. URSS. 12: 188, 1950; Dendrobenthamia hupeiensis Fang, Acta Phytotax. Sin. 2(2): 95, P. 11, 1953; D. wuyishanensis Fang & Hsieh, J. Sichuan Univ. Nat. Sci. 3: 161, Pl. 8, fig.1, 1980; Cornus capitata Wall. subsp. angustata (Chun) Xiang; Benthamidia japonica var. angustata (Chun) Hara;Dendrobenthamia angustata (Chun) Fant; D. angustata var. wuyishanensis (Fang & Hsieh) Fang & W. K. Hu.

Evergreen trees or shrubs, 4--12 m; Barks gray or gray-brown; young branches grayish green, typically pubescent with white, appressed hairs, rarely tomentose. Old branches gray brown, nearly glaberous. Flower buds globose, exposed and subtended by four small, green, young bracts. Leaf buds subtending flower buds, covered by small, triangular to lanceolate scales. Leaves thin leathery to thick leathery, grayish green, oblong eliptic or obovate elliptic or lanceolate, or elliptic, 7--9(--12) x 2--4(--5) cm, 3--4- veined; apex acuminate caudate; base cunnate to broad cuneate; abaxial surface gray-green, densely pubescent with white, appressed hairs, rough to touch, or rarely tomentose; axil of veins sometimes aggregated with white elevated soft hairs. Capitulate cymes globose, 55--80 (--95)-flowered, 0.8--1.2 cm in diameter; bracts yellowish turning white, long ovate to obovate, 2.5--5.0 x 0.9--2.2 cm. Calyx tube short, ca 0.7--1 mm long, truncate to slightly 4-lobed. Petals ovate 2.8 mm. Style cylindric, ca 1.5 mm, densely pubescent with white hairs. Fruitirescence globose, ca 1.5--2.5 cm in diameter, red when mature, pubescent with white, appressed hairs. Penduncles slender, 5.5--10 cm. Fl. June--July, fr. Oct.--Nov.

1a. Yong branches and abaxial surface of leaves pebesent with white appressed hairs……………..var. angustata
1b. Yong branches and abaxial surface of leaves tomentose……………...................................….var. mollis

1a. Cornus angustata var. angustata
Jien Ye Si Zao Hua
Yong branches, abaxial surface of leaves and bracts pubescent with white appressed fine hairs.
Open slope, by streams, or in dense forests; elev. 300--2200 m. Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi?, Sichuan,.
Ripe fruits are sweet-taste and edible.

1b. Cornus angustata var. mollis (Rehd.) Xiang
Rong Mao Jian Ye Si Zao Hua
Cornus capitata Wall. var. mollis Rehd. In Sarg., Pl. Wils. 2: 579. 1916; C. capitata var. hypoleuca Levl., Cat. Pl. Yunnan, p. 59. 1916; Dendrobenthamia angustata var. mollis (Rehd.) Fang; Benthamidia capitata (Wall.) Hara var. mollis (Rehd.) Hara.

Yong branches, abaxial surface of leaves and bracts all tomentose.

Slope, forests; elev. 750--2100 m. W. Hubei, E. Sichuan, NE. Yunnan.

2. Cornus capitata Wall., in Roxb. Fl. Ind. Ed. Carey et Wall., 1:434. 1820
Tou Zhuang Si Zao Hua
Dendrobenthamia emeiensis Fang & Hiseh, Fl. Sichuan, 1: 368. Pl. 141, fig. 1-6, Addenda, 471, 1981; Dendrobenthamia brevipedunculata Fang & Hiseh, J. Sichuan Univ., Nat. Sci. 3: 161, Pl. 2, 1980; Benthamia fragifera Lindl., Bot. Regist. 19: t1579; Cynoxylon yunnanense Pojark. Notul. Syst. Inst Bot. Nom. Kom. Acad. Sci. URSS. 12:185.1950; C. glabriuscula Pojark, l.c. 12: 185. 1950; C. capitata (Wall.) Nakai; Benthamidia capitata (Wall.) Hara; Dendronbenthamia capitata (Wall.) Hutch.; Cornus capitata Wall. subsp. emeiensis (Fang & Hiseh) Xiang; C. capitata Wall. subsp. brevippedunculata (Fang & Hiseh) Xiang; Dendronbenthamia capitata (Wall.) Hutch. var. emeiensis (Fang & Hiseh) Fang & W. K. Hu; D. tonkinensis Fang var. brevipendunculata (Fang & Hsieh).

Evergreen trees or shrubs, 3--15(--20) m; Barks brown to blackish gray; young branches grayish green, pubescent with white, appressed hairs. Old branches gray brown, nearly glaberous. Flower buds globose, exposed and subtended by four small, green, linear lancelate bracts. Leave buds exposed. Leaves thin leathery to coraceous, grayish green, long eliptic or oblong lanceolate, 5--12 x 2--3.5(--4) cm, 3--4-veined, acuminate to shortly caudate at apex, cunnate to broad cuneate at base; abaxial surface gray-green, densely pubescent with white, thick, appressed hairs, rough to touch; axil of veins often pitted or rarely with a cluster of hairs. Capitulate cymes globose, 50-100-flowered, ca 1.2 cm in diameter; bracts white, obovate or broad obovate, rarely rouned, 3.5--6.2 x 1.5--5 cm. Calyx tube short, ca 1.2 mm long, hardly lobed to consipucuously 4-lobed; lobes rounded. Petals oblong, 3--4.0 mm long. Style cylindric, ~1.5 mm, densely pubescent with white hairs. Fruitirescence compressed (or semi-) globose, 1.5--2.5 cm in diameter, purple red when mature, pubescent with small, white hairs. Penduncles short and stout, (1.5) 4--5 (--8) cm. Fl. May--July, fr. Sept.--Nov.

In evergreen or mixed forests; elev. 1000--3200 m. Southwestern China: Guizhou, Sichuan, Yunnan, Xizang [India, Nepal, Bhudan, Myanmar].

Ripe fruits are sweet-taste and edible. Bark is used medicinally; branches and leaves are used for tannin.

In western Guizhou, the eastern edge of its distributional range, the species comes into contact with Cornus angustata. In this area, the distinction of the two species is somewhat obscured. Intermediate specimens, such as specimens with leaves like C. angustata, but fruitrescnece like C. capitata or vice versa, are found. In addition, there are also specimens sparsely pubescent with fine, white hairs and smooth to touch on the abaxial surface (unlike either C. capitata or C. angustata, both of which are densely pubescent with coarse hairs and rough to touch) and compressed globose fruitrescence (like C. capitata) supported by slender peduncles (like C. angustata). This may suggest that either the two species hybridize in the region of their contact or C. capitata and C. angustata represent two subspecies currently undergoing differentiation. The two taxa are preliminarily distinguished by only the peduncle and shape of fruitirescence (stout vs. slender, compressed globose vs. globose) and whether the axil of veins pitted or non-pitted. However, as discussed above, a comparison of allozymes between the two taxa (represented by few specimens) showed quite different profiles between the two taxa. Additional molecular analyses should be helpful for understanding the origin of these variation.

3. Cornus hongkongensis Hemsl., J. Linn. Soc. Bot. 25: 345. 1888.
Xiang Gang Si Zao Hua

Evergreen trees or shrubs, 3--15 (--25) m. Barks gray, dark gray or backish brown, smooth; young branches green o purplish green, sparsely pubescent with brown, appressed hairs or rarely densely pubescent with brown hairs or glabrous. Old branches light gray, gray green, gray brown, with or without lenticels. Winter flower buds globose to cornical, exposed, subtended by four green, later expanded and petaloid, bracts. Leaf buds subtending flower buds, covered by small, triangular to lanceolate scales. Leaves thin subleathery to thick coriaceous, elliptic, oblong elliptic, or obovate oblong, 6.2--13 (--16) x 2.5--6.3 (--7.5) cm, short acuminate to caudate at apex, cuneate, broad cunnate to rounded at base; abaxial surface light green or powder-green, glabrous or sparsely pubescent with white or brown and white hairs when young, often glabrous when old except sometimes in axils of veins, 3--4 (--5)-veined. Capitate cymes globose, ca 40--70-flowered, 0.7 -1.3(--2.0) cm in diameter; bracts yellowish to white, broad elliptic, broad ovate, rounded, obovate oblong elliptic, or obovate, 1.6--4 x 1.3--2 (--4.2) cm, sparsely pubescent or glabrous. Calyx tube short, 0.7--1.3 mm long, shallowly 4-lobed, rarely 5-lobed; lobes truncate to rounded teeth. Petals elliptic, oblong elliptic, ovate-elliptic, ovate-lanceolate to ovate, 1.5--4.2 x 0.8--1.1 mm, sometimes slightly united at base. Style cylindric, 0.5--1.5 mm, sparsely pubescent with white hairs or glabrous. Compound fruit globose, 1.5--2.5 cm in diameter, red, yellow-red when mature, nearly glabrous or slightly pubescent with fne white hairs; peduncles 4--8(--10) cm. Fl. Apr.-Jun, fr. Oct.--Dec.

In thickets, borad-leaved evergreen forest; roadside, by streams, valleys, slopes, and forests. Elev. 650-2500 m. Six subspecies in Guangdong, Fujian, Guangxi, Guizhou, Hunnan, Hong Kong, Jiangxi, Sichuan, Yunnan, Zhejiang.

The species is highly variable in vegetative morphology and was divided into several species by other authors based on minor differences, such as pubescence, shape, and size of various parts. The variation overlaps, but is also more or less associated with geographic distribution. To recognize this pattern, Xiang (Bull. Bot. Res. 7: 33. 1987) recognized six subspecies within the species. This treatment is followed here and need to be evaluated by both field and laboratory data.


  1a. Leaves obovate, large, 8.5--16 cm x 3.8--7.5 cm, thick leathery; inflorescence 
        15--20 mm in diam. (excl. bracts); bracts large, broad obovate or rounded, 4--5 
        x 3-4.2 cm .&............................................................................................................3e. subsp. gigantea
  1b. Leaves ovate to long elliptic, smaller, 5.5-10 (--13.5) x 2.7-5.8 (--6.3) 
        cm; inflorescence smaller, 7--13 mm in diam; bracts smaller, various shaped.
        2a. Outer surface of petals glabrous or nearly so; style glabrous..................................3b. subsp. tokinensis.
        2b. Outer surface of petals pubescent with short appressed fine hairs; style 
              pubescent with white hairs.
              3a. Leaf lateral veins, particularly the secondary lateral veins, inconspicuous..........3d. subsp. elegans
              3b. Leaf lateral veins conspicuous; secondary lateral veins mosstly horizontal.
                    4a. Leaf abaxial surface and veins densely pubescent with red-brown, coarse 
                          hairs.........................................................................................................3f. subsp. ferruginea
                    4b. Leaf abaxial surface sparsely pubescent with thin hairs; veins hairless 
                          or nearly so.
                          5a. Leaves subcoriaceous; axils of veins on lower surface often covered by a 
                                cluster of "Y"-shaped soft hairs; glabrous or rarely pubescent with 
                                brown hairs........................................................................................3c. subsp. melanotricha
                          5b. Leaves leathery, thick leathery; no cluster of "Y" shaped hairs 
                                in axils of veins, sparsely pubescent with brown and white, appressed hairs 
                                on lower surface when young; often glabrous with conspicuous brown hair scars 
                                 (brown spots) when old.....................................................................3a. subsp. hongkongensis
  
3a. Cornus hongkongensis Hemsl. subsp. hongkongensis
Xiang Gang Si Zao Hua
Benthamia japonica Sieb. & Zucc. var. sinensis Benth. In Hook. Kew J. Bot. 4:165. 1852; Dendrobenthamia latibracteata Fang & Hsieh, J.. Sichuan Univ., Nat. Sci. 3: 163, pl. 10, 1, 1980; Dendrobenthamia hongkongensis (Hemsl.) Hutch; Benthamia hongkongensis (Hemsl.) Nakai; Cynoxylon hongkongensis (Hemsl.) Nakai; Benthamidia hongkongensis (Hemsl.) Hara.

Evergreen trees or shrubs, 5-15 (25) m; bark deep gray or deep brown; old branches gray or brown, glabrous, with conspicuous lenticels. Leaves coriaceous elliptic to long elliptic, rarely obovate elliptic, 6.0--13 x 2.8--6.3 cm, pubescent with white and brown, soft, appressed hairs on both sides when young, gradually become glabrous and scattered with brown spots (the hair scars) when old; 2-3(-4)-veined. Inflorescent 50--70-flowered, ca 1.0 cm in diam.; bracts white, broad elliptic to obovate-broad elliptic, 2.8--4.1 x 1.7-3.5 cm, nearly glabrous on both sides. Peduncles 3.5--10 cm, pubescent with brown appressed hairs. Flowers small, fragrant; calyx tubular, 0.7--0.9 mm, with brown hairs at the base, pubescent; petals oblong elliptic, 2.2--2.4 x 1--1.2 mm, light yellow; filaments 1.9--2.1 mm, slightly pubescent; anthers elliptic, dark brown; style ca 1.0 mm, pubesent with white hairs. Compound fruit ca. 2.5 cm in diam., with white thin hairs, yellow or red when mature with a 3.5--10 cm long peduncle. Fl. May-Jun; Fr. Nov.-Dec.

In evergreen broad-leaved forests; elev. 600-1800 m. Southern China: Guangdong, Guangxi, Guizhou, Hunan, Hong Kong [Veitnam].

Wood is used as building material; fruits edible, used for wine brewing.

3b. Cornus hongkongensis Hemsl. subsp. tonkinensis (Fang) Xiang, Bull. Bot. Reserch. 7(2): 42. 1987.
Dendrobenthamia tonkinensis Fang, Acta Phytotax. Sin. 2: 103. Pl. 15. 1953; Cornus tonkinensis (Fang ) Tard.-Bl.

Evergreen trees or shrubs, 4-15 m; bark dark gray; old branches gray with lenticels. Leaves leathery, oblong ovate or long elliptic, 4.5--11 (--13) x 1.7--5.3 (--6) cm, nearly glabrous, 3(--4)-veined. Inflorescences 40--50-flowered, ca 0.8 cm in diam; bracts white, broad elliptic to broad ovate, 1.6--1.8 x 1.3--1.5 cm, slightly pubescent on both surfaces. Calyx nearly hairless. Petals 2.2 x 0.8--1.0 mm, glabrous; filatments short, glabrous; anthers broad elliptic, 0.7 mm. Style short and thick, ca 0.5 mm, hairless. Compound fruit 1.5--2.0 cm in diam, red when mature with a 4--7.5 cm peduncle. Fl. June; fr. Dec.

In evergreen broad-leaved forest; elve. 1120--2500 m. South to southwestern China: S. Yunnan and SW Guangxi [N. Veitnam];.

3c. Cornus hongkongensis Hemsl. subsp. melanotricha (Pojark.) Xiang, Bull. Bot. Reserch. 7 (2): 42. 1987
Guang Ye Xiang Gang Si Zao Hua
Cynoxylon melanotricha Pojark. In Notul. Syst. Inst. Bot. Nom. Kom. Acad. Sci. URSS. 12: 191. 1950; Dendrobenthamia jinyunensis Fang & W. K. Hu in Fl. Sichuan 1: 362. Pl. 137, fig. 1-6, addenda 471, 1981; Dendrobenthamia gigantea (Hand.-Mazz.) Fang var. caudate Fang & W,K, Hu in Fl. Sichuan 1: 360. Pl. 138, addenda 471, 1981; Dendrobenthamia melanotricha (Pojark.) Fang; D. ferruginea (Wu) Fang var. jinyunensis (Fang & W. K. Hu) Fang & W. K. Hu.

Evergreen trees or shrubs, 3--12 m; bark dark gray or dark brown; old branches gray brown, glabrous, often without lenticels. Leaves subleathery or leathery, elliptic to long elliptic, 6--10 x 2.7--5 cm, 3(--4)-veind; veins conspicuous; abaxial surface sparsely pubescent with white or brown, appressed short hairs when young, often glabrous when old; axils of veins with clusters of dark brown, "Y" shaped long hairs. Inflorescences ca 40-flowered, greenish yellow, 1.0 cm in diam; bracts broad elliptic or broad obovate, 2--4 x 1--3.5 cm, yellowish green at first, creamy white later, glabrous. Calyx tube 0.9 mm, with brown hairs at base; petals long elliptic or long ovate, 2.8--3 x 1.0 mm, sparsely pubescent on outer surface. Filaments 2.2 mm, glabrous; anthers ovate elliptic, brown, 1.0 mm long. Style ca 1.3 mm pubescent with white hairs. Compound fruit 2--2.5 cm in diam., red when mature with a 3.7--9.5 cm peduncle. Fl. May-Jun; fr. Oct.-Nov.

In evergreen broad-leaved forests; elev. 450-1800 m. Southwestern China: Guizhou, Hunnan, Sichuan, Yunnan.

The hard wood is used for making farming tools.

3d. Cornus hongkongensis Hemsl. subsp. elegans (Fang & Hsieh) Xiang, Bull. Bot. Reserch. 7 (2): 42. 1987
Xiu Li Xiang Gang Si Zao Hua
Dendrobenthamia elegans Fang & Hsieh, J. Sichuan Univ., Nat. Sci. 3: 162. Pl. 9, fig. 1. 1980; D. rotundifolia Fang & Hsieh, J. Sichuan Univ., Nat. Sci. 3: 162/ Pl. 8, fig. 2. 1980; D. elegans Fang & Hsieh var. rotundifolia (Fang & Hsieh) Fang & W. K. Hu.

Evergreen small trees or shrubs, 3-8 (--15) m; bark grayish or gray-brown; old branches gray or gray brown, glabrous. Leaves leathery, elliptic to oblong elliptic, 5.5-8.2 x 2.5-3.5 cm, 3--4-veind; secondary lateral veins inconspicuous; abaxial surface glabrous; sometimes pitted in vein axils. Inflorescences ca 45--55-flowered, 8 mm in diam; bracts obovate elliptic, 3.5--4 x 1.8-2.5 cm, sparsely pubescent with brownish fine, apprssed hairs. Calyx tube 0.7--0.9 mm, pubescent on outer surface, with a whorl of red-brown long hairs at the base; petals ovate elliptic, 2.0-2.5 x 0.8--1.0 mm, sparsely pubescent on outer surface. Filaments 1.8-2.0 mm; anthers elliptic, ca. 0.6 mm long. Style 0.7-0.9 mm, pubescent with white short hairs. Compound fruit 1.5-2.0 cm in diam., red when mature with a 4.5-9.0 cm peduncle. Fl. May-Jun; fr. Nov.

By streams, in forests; elev. 250-1200 m. Southeastern China: Fujian, Jiangxi, Zhejiang.

3e. Cornus hongkongensis Hemsl. subsp. gigantea (Hand.-Mazz.) Xiang, Bull. Bot. Reserch. 7 (2): 42. 1987.
Dai Ye Xiang Gang Si Zao Hua
Cornus hongkongensis var. gigantean Hand.-Mazz. Symb. In. 7: 690. 1933; Dendrobenthamia pachyphylla Fang & W. K. Hu, J. Sichuan Univ., Nat. Sci. 3: 163. Pl. 9, fig. 2, 1980; D. gigantean (Hand.-Mazz.) Fang var. caudate Fang & W.K. Hu, in Fl. Sichuan 1: 360. Pl. 138, fig. 7, addenda: 471. 1981; Benthamidia hongkongensis var. gigantea (Hand.-Mazz.) Hara; Dendrobenthamia gigantea (Hand.-Mazz.) Fang; Cornus gigantea (Hand.-Mazz.) Tard.-Bl.;

Evergreen small trees or shrubs, 2--6 m; bark dark brown, smooth; old branches gray or gray green. Leaves leathery to thick leathery, obovate, rarely broad elliptic, 8.5--16 x 3.8-7.5 cm, caudate-acumminate at apex, often 4-veind; veins conspicuous; abaxial surface sparsely pubescent with appressed short hairs when young, glabrous when old; axils of veins without or rarely with clusters of dark brown hairs. Inflorescences ca 60-flowered, 1.4-2.0 cm in diam; bracts yellowish to white, broad obovate or nearly rounded, ca 4 x 3-4.2 cm, nearly glabrous. Calyx tube 1.3 mm, often shallowly 4-lobed, rarely 5-lobed, pubescent on outer surface. Petals ovate lanceolate, 4.2 x 1.1 mm, sparsely pubescent on outer surface. Filaments ca 4 mm; anthers ovate elliptic, ca. 1.2 mm long. Style ca 1.5 mm, pubescent with white hairs. Compound fruit 2.5 cm in diam., yellow-red when mature; penduncle 8-9.5 cm. Fl. Apr--May; fr. unknown.

In valleys, sparse forests, evergreen broad-leaved forests; elev. 700--1700 m. Southwestern China: W. Guizhou, S. Sichuan, NE. Yunnan.

3f. Cornus hongkongensis Hemsl. subsp. ferruginea (Wu) Xiang, Bull. Bot. Reserch. 7 (2): 42. 1987.
He Mao Xiang Gang Si Zao Hua
Cornus ferruginea Wu, Bot. Jahrb. 71: 199. 1940; Dendrobenthamia ferruginea (Wu) Fang var. jiangxiensis Fang & Hsieh, J. Sichuan Univ., Nat. Sci. 3: 164. 1980; Benthamidia ferruginea (Wu) Hara; Cynoxylon ferruginea (Wu) Pojark; Dendrobenthamia ferruginea (Wu) Fang.

Evergreen small trees or shrubs, 5-12(--20) m; young branches densely pubescent with brown hairs; old branches dark gray green. Leaves subleathery to paperous, long elliptic to broad elliptic, 8-14 x 3.5-6.0 cm, 4 (--5)-veind; veins conspicuous; abaxial surface powder-green (or pale green), pubescent with brown, thick hairs, later gradually glabrous; veins with long brown hairs. Inflorescences 60--70-flowered, ca. 1.1 cm in diam; bracts yellowish white, broad obovate elliptic, ca 4-4.5 x 2.5--3 cm, sparsely pubescent with thin appressed hairs. Calyx tube ca 1.2 mm, often shallowly 4-lobed, pubescent with white and brown hairs on outer surface. Petals long elliptic, ca. 2.5 mm long, sparsely pubescent on outer surface. Filaments ca 1.7 mm; anthers elliptic, yellow, ca. 0.8 mm long. Style ca 1.5 mm, pubescent with white hairs. Compound fruit .3-1.8 cm in diam., red when mature; penduncle 8-9.5 cm. Fl. Jun; fr. Oct.-Dec.

Slope, valleys, roadside, sparse forest, or dense forest; elev. 200--1100 m. Southern China: Guangdong, Guangxi, Guizhou(?).

Fruits are sweet and edible.

4. Courns kousa Hance, J. Linn. Soc. Bot. 13: 1.5. 1873.
Dong Yin Si Zao Hua
Cornus? Japonica DC., Prodr. 4: 273. 1830; Benthamia japonica Sieb. & Zucc. Fl. Jap. 1: 38, t. 16. 1836; Cornus kousa Buerg. Ex Miq. In Ann. Mus. Bot. Lugd.-Bat. 2: 159, 1865, pro syn.; Benthamia kousa Nakai, Bot. Mag. Tokyo 23: 41. 1909; Benthamia japonica a. typical, b. minor, c. exsucca, Nakia, Bot. Mag. Tokyo 28: 314. 1914; Benthamia viridis Nakai, Chosen-Shokubutsu, 1: 426, fig. 532, March, 1914; Cynoxylon kousa Nakai, C. japonica (Sieb. & Zucc.) Nakai; Dendrobenthamia japonica (Sieb & Zucc.) Hutch; Benthamidia japonica (Sieb. & Zucc.) Hara; Dendroenthamia japonica (A. P. DC.) Fang.

Deciduous small trees 2--10 m. Barks gray; young branches light green or redish green, pubescent with white, appressed hairs or rarely densely pubescent with white long soft hairs. Old branches dark brown, with or without lenticels. Winter buds mixed, globose, completely covered by two pairs of scales. Leaves thin paperous to thick coraceous, ovate to ovate elliptic, 5.5--12 x 3.5-7.0 cm; apex acuminate; base broad cuneate or rounded; margin entire or rarely finely dentated; abaxial surface light green or powder-green, pubescent with white appressed hairs, papillate or non-papillate; axil of veins with yellow or brown, long, soft hairs, 4--5-veined. Capitate cymes globose, ca 40--50-flowered, ca 0.8-1.0 cm in diam; bracts white, rarely yellow or pinkish, ovate or ovate-lanceolate, nearly glabrous. Calyx tube 4-lobed; lobes broad triangular to rouned, pubescent with white appessed hairs on inner side. Petals greenish to yellowish, oblong elliptic to ovate, 1.5--4.2 x 0.8--1.1 mm, sometimes slightly united at base. Anthers yellowish or dark blue to blackish. Style cylindric, densely pubescent with white hairs. Disk of flower glabrous or hirsute. Compound fruit globose, 1.5--2.0 cm in diameter, red when mature, slightly pubescent with fine white hairs; peduncles 5.5-6.5 cm, with or without a ring of conspicuous thickening at base. Fl. May--Jun, fr. Sept--Oct.

Two subspecies. China, Korea, and Japan. I subspecies in China.

4a. Cornus kousa Hance subsp. chinensis (Osborn) Q. Y. Xiang, Bull. Bot. Res. (Harbin) 7(2):46.
Si Zao Hua

Cornus kousa Hance var. chinensis Osborn, Gard. Chron. Ser. 3, 72: 310. 1922. Benthamia chinensis Hort. Ex Lavallee, Arb. Segrez. 129. 1877, pro syn.; Cynoxylon sinensis Nakai, J. Jap. Bot. 15: 741. 1939; Cynoxylon pseudokousa Pojark, Notul Syst. Inst. Bot. Nom. Kom. Acad. Sci. URSS. 12: 193. 1950; Dendrobenthamia japonica (A.P. DC.) Fang var. huxiensis Fang & W. K. Hu in F. Sichuan 1: 371, addenda 472, 1981; D. japonica (DC.) Fang var. leucotricha Fang et Hsieh in Fl. Sichuan 1: 371. Pl. 142, fig. 7-11, addenda: 472. 1981; Benthamidia sinensis (Nakai) Yamazaki, J. Japanese Bot. 75: 373. 2000; Cornus kousa Buerg. var. yaeyamensis Hatus., Fl. Ryukyus: 457. 1971, nom. Semi nud.; Cornus yaeyamensis (Hatus.) Hatus; Benthamidia japonica var. chinensis (Osborn) Hara; Dendrobenthamia japnica (A. P. DC.) Fang var. chinensis (Osborn) Fang.

Deciduous small trees. Young branches red brown with rounded lenticels. Leaves paperous to thick chartaceus; abaxial surface powder-green, often densely papillate and pubescent with appressed hairs, rarely with white long soft hairs; hairs gradually deciduous; axil of veins often with a cluster of white to brown long soft hairs. Petals greenish to yellowish. Calyx with a ring of brown short hairs. Anthers yellow, sometimes dark blue or blackish. Penduncles often with a ring of conspicuous thickening at the base.

Valley, shaded slopes, by streams, roadside, mixed, sparse or dense woods. Elev. 450-2200 m. Anhui, Gansu, Guizhou? Henan, Hubei, Hunan, Jiangsu, Jiangxi, Nei Mongol, Shaanxi, Shanxi, Sichuan, Taiwan? Yunnan? Zhejiang [Japan]. ---Check local Floras!!
Fruits are sweet and edible; often used for wine brewing.

5. Courns multinervosa (Pojark.) Xiang Bull. Bot. Res. 7(2): 33. 1987.
Duo mai Si Zao Hua
Cynoxylon multinervosa Pojark. In Nutul. Syst. Inst. Bot. Nom. Kom. Acad. Sci. URSS. 12: 194. 1950; Dendrobenthamia multinervosa (Pojar.) Fang.

Deciduous trees 4-8 (--15) m. Barks blackish brown; young branches green or purplish green, sparsely pubescent with white, appressed hairs. Old branches grayish purple or gray brown, glabrous, with white, elliptic lenticels. Winter buds mixed, globose, completely covered by two pairs of scales. Leaves paperous, long elliptic or ovate elliptic, 6--13 x 3--6 cm; apex acuminate; base cuneate, occationally oblique; margin entire or rarely inconspicuously undulated; abaxial surface light green, pubescent with white appressed hairs, rough to touch; hairs later often deciduous; veins (5--)6(--7)-veined, all nearly extending to the apex. Capitate cymes globose, ca 30--45-flowered, ca 1.0 cm in diam; bracts white or yellow, ovate or elliptic. Calyx tube ca. 0.8 mm, 4-lobed; lobes tooth-like or rounded, pubescent with white or brown appessed hairs on innder side. Petals long elliptic, ca 2.5 x 1.0 mm. Anthers elliptic, blackish brown. Style cylindric, lower part densely pubescent with white hairs. Compound fruit globose, 1.2-1.6 cm in diameter, red when mature; peduncles 7.0-10.0 cm, usually without a thickened ring at base. Fl. May--Jun, fr. Oct--Nov.
In mixed woods; elev. 900-2700 m. Sichuan, Yunnan.

6. Subg. Actocrania Endl. Ex Reichenbach, Repert. Herb. 143. 1841.

Cao Zhu Yu Ya Shu
Cornus sect. Arctocrania Endl. Gen. Pl. 798. 1839; Cornus sect. Cornion Spach Hist. Beg. Phan. 7: 103. 1839; Cornella Rydg., Bull. Torr. Bot. Club. 33: 147. 1906; Arctocrania Nakai, Jap. Bot. Mag. Tokyo 23: 39. 1909; Chamaepericlymenum Hill, Brit. Herb. 331. 1756; Chamaepericlymenum Graebn. in Aschers & Graebn., Fl. Nordost, Flachl. 225. 1898.

Small perennial herbs with vertical stem and creeing rhizomes. Leaves a whorl of six at summit (or opposite at all nodes), entire, palmately or pinnately veined, sessile or shortly petiolated. Inflorescence a terminal minute compound cyme subtended by 4 white or rarely pinkish petaloid bracts. Flowers small; calyx tubular, 4-detated; petals oblong ovate to ovate, white or dark purple or purple above and white below; one or two petals often with a soft awn-like appendage near the apex. Stemens 4; filament short; anthers oblong or oblong ovate. Ovary 2-locular, one pendulous ovule per locule; style cylindric; stigma small, capitate. Fruit globose, red; stones elliptic ovate to subglobose.

3 species circumboreal, from Europe eastward to northeastern Asia to northern North America, also in high mountains of Myanmar. 1 in northeastern China.

1. Cornus canadensis L. Sp. Pl. (ed. 1.) 117. 1753. Chamaepericlymenum canadense (L.) Aschers & Graebn.; Cornella canadensis (L.) Rydb.

Perennial Rhizomatous herb, ca 10-20 cm. Rhizomes slender, creeping. Vertical stems slender, unbranches, Leaves opposite, often appearing to be a whorl of six green leaves at terminal node due to reduction of branches, two larger and four smaller; smaller ones developng from the axil buds of the larger leaves; leaves at lower nodes rudimentary, non-chlorophyllous. Green leaves characeous, obovate to more or less diamond shape, 3.5--4.8 x 1.5-2.5 cm; entire, short accuminate at apex, cuneate at base, 2-3 mm-shortly petiolate, 2-3-veined. Inflorescences minute compound cymes, terminal; bracts white, broad ovate, ca 0.8-1.2 x 0.5-1.1 cm, with 7 parallele veins. Flowers white, ca 2 mm in diam. Calyx tube obovate, ca 1 mm, densel pubescent with grayish white appressed hairs; teeth higher than the disk. Petals creamy white, ovate lanceolate, 1.5-2.0 mm long, reflexed. Stamens ca 1 mm; anthers narrow ovate, yellowish white. Style ca 1 mm, glabrous. Fruit ca. 5 mm in diam; stones elliptic ovate. Fl. Jul-Aug; fr. globose, red when mature.
In sparse mountain coniferous forests; in sunny and moist habitats; edge of woods, mossy beds, old tree stump. Elev. Ca 1200m. Changbei Mountian of Jilin [Korea, Japan, far east of Russia, Canada, North America].