Species

C H A P T E R

2 Species 2.1 Actinidia arguta 2.1a Actinidia arguta var. arguta 2.1b Actinidia arguta var. giraldii 2.2 Actinidia callosa 2.2a Actinidia callosa var. callosa 2.2b Actinidia callosa var. acuminata 2.2c Actinidia callosa var. discolor 2.2d Actinidia callosa var. henryi 2.2e Actinidia callosa var. strigillosa 2.3 Actinidia chengkouensis 2.4 Actinidia chinensis 2.4a Actinidia chinensis var. chinensis 2.4b Actinidia chinensis var. deliciosa 2.4c Actinidia chinensis var. setosa 2.5 Actinidia chrysantha 2.6 Actinidia cylindrica 2.6a Actinidia cylindrica var. cylindrica 2.6b Actinidia cylindrica var. reticulata 2.7 Actinidia eriantha 2.8 Actinidia farinosa 2.9 Actinidia fasciculoides 2.9a Actinidia fasciculoides var. fasciculoides 2.9b Actinidia fasciculoides var. cuneata 2.9c Actinidia fasciculoides var. orbiculata 2.10 Actinidia fortunatii 2.11 Actinidia fulvicoma 2.11a Actinidia fulvicoma var. fulvicoma 2.11b Actinidia fulvicoma var. cinerascens 2.11c Actinidia fulvicoma var. hirsuta 2.11d Actinidia fulvicoma var. pachyphylla 2.12 Actinidia glaucocallosa 2.13 Actinidia grandiflora 2.14 Actinidia hemsleyana 2.15 Actinidia henryi 2.16 Actinidia holotricha 2.17 Actinidia hubeiensis 2.18 Actinidia indochinensis 2.18a Actinidia indochinensis var. indochinensis 2.18b Actinidia indochinensis var. ovatifolia 2.19 Actinidia kolomikta 2.20 Actinidia laevissima 2.21 Actinidia lanceolata 2.22 Actinidia latifolia 2.22a Actinidia latifolia var. latifolia 2.22b Actinidia latifolia var. mollis

46 46 48 49 49 51 52 54 56 57 59 59 61 64 66 68 68 70 71 74 76 76 77 78 79 81 81 83 85 86 87 89 91 92 94 96 97 98 99 100 103 104 106 106 108

2.23 Actinidia liangguangensis 2.24 Actinidia lijiangensis 2.25 Actinidia linguiensis 2.26 Actinidia longicarpa 2.27 Actinidia macrosperma 2.27a Actinidia macrosperma var. macrosperma 2.27b Actinidia macrosperma var. mumoides 2.28 Actinidia melanandra 2.28a Actinidia melanandra var. melanandra 2.28b Actinidia melanandra var. glabrescens 2.29 Actinidia melliana 2.30 Actinidia obovata 2.31 Actinidia persicina 2.32 Actinidia pilosula 2.33 Actinidia polygama 2.34 Actinidia rongshuiensis 2.35 Actinidia rubricaulis 2.35a Actinidia rubricaulis var. rubricaulis 2.35b Actinidia rubricaulis var. coriacea 2.36 Actinidia rubus 2.37 Actinidia rudis 2.37a Actinidia rudis var. rudis 2.37b Actinidia rudis var. glabricaulis 2.38 Actinidia rufa 2.39 Actinidia rufotricha 2.39a Actinidia rufotricha var. rufotricha 2.39b Actinidia rufotricha var. glomerata 2.40 Actinidia sabiifolia 2.41 Actinidia sorbifolia 2.42 Actinidia stellatopilosa 2.43 Actinidia styracifolia 2.44 Actinidia suberifolia 2.45 Actinidia tetramera 2.46 Actinidia trichogyna 2.47 Actinidia ulmifolia 2.48 Actinidia umbelloides 2.48a Actinidia umbelloides var. umbelloides 2.48b Actinidia umbelloides var. flabellifolia 2.49 Actinidia valvata 2.50 Actinidia venosa 2.51 Actinidia vitifolia 2.52 Actinidia zhejiangensis 2.53 Actinidia hypoleuca 2.54 Actinidia strigosa

Kiwifruit: The Genus ACTINIDIA. http://dx.doi.org/10.1016/B978-0-12-803066-0.00002-2 Copyright © 2016 China Science Publishing & Media Ltd. Published by Elsevier Inc. All rights reserved.

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Kiwifruit: The Genus ACTINIDIA

2.1  Actinidia arguta 软枣猕猴桃 ruan zao mi hou tao

Actinidia arguta (Siebold & Zuccarini) Planchon ex Miquel in Ann. Mus. Bot. Ludg.-Bat. 3: 15. 1867. Trochostigma arguta Siebold & Zuccarini in Abh. Akad. Wiss. München. 3: 727. 1843. TYPE: not designated.

2.1a  Actinidia arguta var. arguta 软枣猕猴桃 (原变种) ruan zao mi hou tao (yuan bian zhong)

Actinidia melanandra Franchet var. latifolia E. Pritzel in Bot. Jahrb. Syst. 29: 470. 1900. Actinidia arguta var. cordifolia (Miquel) Bean in Trees & Shrubs Brit. Isl. 1: 162. 1914. Actinidia cordifolia Miquel in Ann. Mus. Bot. Ludg.-Bat. 3:15. 1876. TYPE: Japan. Kiusiu, Pierot s.n.

Figure 2.1  Natural range of Actinidia arguta var. arguta.

Actinidia megalocarpa Nakai ex Nakai & Kitagawa in Rep. 1st. Sci. Exp. Manch. IV. 1: 9, t. 3. 1933 (Pl. Nov. Jehol.). TYPE: Japan. Chang-Shan-Yü, N.H.K. September 13, 1933. Actinidia chartacea Hu in Bull. Fan Mem. Inst. Biol. Bot. 10: 128. 1940. TYPE: China. Sichuan: Mo-Tien-Ling, August 31, 1930, F.T. Wang 22454a (Holotype, PE). Actinidia arguta var. purpurea (Rehder) C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 208. 1984. A. purpurea Rehder in Sarg. Pl. Wils. 2: 378. 1915. TYPE: China. Sichuan: Tachien-lu, E.H. Wilson 1314 (Holotype, A). Actinidia platyphylla A. Gray ex Miquel 1. c. TYPE: Japan. Hokkaido: Cape Sangar, J. Small s.n. (Holotype, GH). Distribution: China: Yunnan, Guangxi, Guizhou, Hunan, Jiangxi, Fujian, Zhejiang, Sichuan, Chongqing, Hubei, Anhui, Gansu, Shaanxi, Henan, Shanxi, Shandong, Hebei, Beijing, Tianjin, Liaoning, Jilin, Heilongjiang, and Taiwan; most parts of Japan and Far East region of Russia (Figure 2.1).

Chapter 2 Species

Morphological characteristics Large deciduous climbing vines. Stems One-year-old shoots are gray, light gray or russeted, glabrous, or sparsely covered with white pubescence; lenticels are distinct, long-shuttle shaped, light color. Twoyear-old stems are grayish brown, glabrous and c. 4 mm in diameter; pith is lamellate, white.

Figure 2.2  Stem and leaf of Actinidia arguta var. arguta.

Figure 2.3  Flower and fruit of Actinidia arguta var. arguta.

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Leaves Chartaceous; ovate, oblong, and occasionally broad obovate, 8-12 cm long and 5-10 cm broad; leaf base is nearly round or broad truncate, leaf apex is acute or mucronate. The upper surface is dark green and glabrous. Margin serrations are dense and appressed. The lower surface is light green, main veins and secondary vein axils covered with grayish-white or yellow fascicled hairs. The petiole is green or light red, 3-7 cm long (Figure 2.2).

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Kiwifruit: The Genus ACTINIDIA

Pistillate flowers Predominantly single, cyme inflorescence is axillary with one to three flowers. Pedicel c. 7-15 cm long. Flowers are 12-20 mm in diameter; five to six sepals of calyx are ovate, c. 6 mm long. Four to six petals, obovate and c. 7-10 mm long, 4-7 mm wide, white to light green. Filaments are white, 4 mm long, c. 44 filaments per flower. Anthers are black or dark purple, sagittate. Styles are white, usually 1822 per flower, 6-7 mm long. Ovary is bottle-shaped and glabrous, 6-7 mm long. Blooming is in May.

Morphologically, A. arguta is closely related to A. melanandra and A. hypoleuca (Li, 1952; Liang, 1984; Li et al., 2007a); this relationship is supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). AFLP analysis showed a Nei’s genetic diversity of 0.0907 and a Shannon’s polymorphic information index of 0.1325 for genotypes of A. arguta in the National Germplasm Repository for Actinidia, Wuhan Botanic Garden (WBG), Chinese Academy of Sciences (CAS), Wuhan, Hubei, P.R. China (Li, 2006).

Staminate flowers  Cyme inflorescence, many flowered; shape of petals is similar to that of pistillate flowers. Usually 44 stamens. Filaments are white, 3-5 mm long. Anthers are dark-brown or dark purple to black, c. 2 mm long. Ovary vestigial.

Chromosomes: 2n = 2x = 58, 4x = 116, 6x = 174, 8x = 232

Fruit  Mostly ovoid or globose, not spotted (immaculate) and glabrous. The color of unripe fruit varies from light green, dark green to yellow-green; nearly ripe fruit from green to light red to purplish-red. Apex of fruits is round or rostrate. Fruit c. 4 cm long and 2.5 cm in diameter. Fruit average 5-7.5 g. Fruit flesh is green or emerald-green and juicy. The fruit are sweet with slightly acid taste and ripen in August. Vitamin C (Vc) content is 81-430 mg per 100 g fresh weight. Soluble solid content is 14-15% and total acidity 0.9-1.3%. The fruit is suitable for fresh consumption and processing (Figure 2.3).

Diagnostic characters and species relationships Lamellate pith, glabrous bottle-shaped ovary, spotless fruits (no lenticels), and leaf glaucous below.

Figure 2.4  Natural range of Actinidia arguta var. giraldii.

2.1b  Actinidia arguta var. giraldii 陕西猕猴桃(变种) shaanxi mi hou tao

Actinidia arguta var. giraldii (Diels) Voroshilov in Byull. Glavn. Bot. Sada (Moscow) 84: 33. 1972. A. giraldii Diels in Bot. Jahrb. Syst. 36 (Beibl. 82): 75. 1905. TYPE: China. Shaanxi, July 1891, giraldi 4065. Actinidia arguta var. nervosa C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 309. 1984. TYPE: China. Zhejiang: Tiantaishan, alt. 900 m, July 20, 1959, Zhejiang Bot. Exped. 28283 (Holotype, PE). Actinidia melanandra Franchet var. kwangsiensis (H.L. Li) C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 210. 1984. A. kwangsiensis H.L. Li in J. Arnold. Arbor. 33: 26. 1952. TYPE: China. Guangxi: Luocheng, alt. 1000 m, June 20, 1926, R.C. Ching 6185 (Holotype, NY; Isotype, PE). Distribution: China: Guangxi, Hunan, Jiangxi, Zhejiang, Sichuan, Chongqing, Hubei Gansu, Shaanxi, and Henan (Figure 2.4).

Chapter 2 Species

Morphological characteristics Stems  One-year-old shoots are gray-greenish brown covered with yellow-green downy hairs; pith is lamellate, white. Two-year-old stems are grayish brown, glabrous. Leaves Chartaceous; broad ovate, ovate, unequal; leaf blade is 6-16 cm long and 6-11 cm broad; the leaf bases are broad cuneate, rounded or subcordate, the apices are mucronate and mostly twisted. The leaf margin is undulate, sharp teeth appressed. The upper surface is green; the apical half is glabrous, but the petiolar half of the leaf is covered by many curly downy hairs; veins densely covered with downy hairs. The lower surface is green, densely covered with curly downy hairs, similar to the upper surface. The petiole is green or light red, sparsely covered with downy hairs, 4.5-7 cm long. Pistillate flowers  Flowers are yellowish white; anthers are purple. Staminate flowers  Cyme inflorescence, 3-14 flowers per inflorescence. Fruit  Oblong-ovoid or cylindric-ellipsoid; skin is thick and amaranth in color. Apex is slightly protruded and base is flat. Calyx shed, not persistent. Fruit c. 3 cm long and 1.7 cm in diameter. The fruit average 2.1 g, amaranth colored pulp with medium-sized core, light sweet, medium juicy and taste without aroma. The fruit ripen in August (Figure 2.5).

Figure 2.5  Stem, leaf, and fruit of A. arguta var. giraldii. Photographs provided by Zhangying.

Diagnostic characters and species relationships A. arguta var. giraldii differs from A. arguta var. arguta by the underneath of the leaf having curly tomentum on midvein and lateral veins or covering the entire leaf surface. The variety is very closely related to A. arguta var. arguta and may also be related to A. melanandra and A. hypoleuca.

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2.2  Actinidia callosa 硬齿猕猴桃 ying chi mi hou tao

Actinidia callosa Lindley in Nat. Syst. ed. 2, 439. 1836. TYPE: Nepal, N. Wallich 6634 (Lectotype: N. Wallich 6634, sheet Herb. K negative 18308).

2.2a  Actinidia callosa var. callosa 硬齿猕猴桃(原变种) ying chi mi hou tao (yuan bian zhong)

Actinidia arisanensis Hayata in Icon. Pl. Formos. 8: 11. 1919. Actinidia rankanensis Hayata, op. cit. 13. Actinidia remoganesis Hayata, op. cit. 13. Actinidia callosa var. formosana Finet & Gagnepain in Bull. Soc. Bot. France 52, Mém. 4: 20. 1905 (Contr. Fl. As. Or.). Actinidia formosana Hayata in Icon. Pl. Formos. 8: 12. 1919. Actinidia callosa var. pubiramula C.Y. Wu in Fl. Yunnan. 1: 67. 1977. TYPE: China. Yunnan: Xichou, May 9, 1959, Q.A. Wu 8016 (Holotype, KUN). Distribution: China: Yunnan, Guangxi, Guangdong, Guizhou, Hunan, Jiangxi, Fujian, Zhejiang, Sichuan, Chongqing, Hubei, Gansu, Shaanxi, and Taiwan (Figure 2.6).

Morphological characteristics Large, deciduous climbing vines. Stems  One-year-old shoots are brown, glabrous and smooth; lenticels conspicuous, relatively large, dense, oblong, yellow-white, internodes 1-2 cm long. Two-yearold stems are dark-brown and glabrous, lenticels inconspicuous, round or oblong, dark yellow-brown. Older stems are light brown and glabrous, lenticels are sparse, oblong or linear shape, yellow-brown; pith is lamellate, brown. Leaves  Thick, chartaceous to subcoriaceous, oblong or nearly elliptic, leaf blade is 5-10 cm long and 2.5-6 cm broad, almost bilateral symmetry, base auriculate-cordate, broad rounded or truncate, with a mucronate upper part. The upper surface is dark green, lustrous and glabrous. Almost entire margin at base of leaf, leaf margin is densely serrated, mucronulates are brown and appressed, main vein and secondary veins are green and glabrous. The lower surface is light green and glabrous, main vein and secondary veins are whitish-green, sparsely covered with white pubescence. The petiole is purple-red, glabrous, c. 1.8-2.7 cm long and slender (Figure 2.7). Pistillate flowers Single. Pedicel is c. 1.0 cm long, whitishgreen and glabrous. Five sepals of calyx are short elliptical to ovate, c. 4 mm long and 2 mm broad, light reddish-brown, with four distinct dark-brown longitudinal stripes, the inner surface of sepals is glabrous and the outer surface is partially covered with sparse, light purplish pubescence on

50

Kiwifruit: The Genus ACTINIDIA

Figure 2.6  Natural range of Actinidia callosa var. callosa.

milky-white, c. 24 per flower, 4 mm long with slightly swollen stigmata. Ovary is round, c. 3 mm long and 3 mm broad, covered with dense, slight, red pubescence. Staminate flowers  Cyme inflorescence, one to three flowers per inflorescence. Pedicel is c. 7 mm long, light purplish-red, green and sparsely covered with purplish-red, short pubescence. Peduncle is c. 4 mm ­ long, light purplish-red and sparsely covered with light, purplish-red pubescence. Calyx of five sepals, oblong, c. 6 mm long and 3 mm broad, green, sparsely covered with distinct red stripes in middle area of inner surface, with purplish-red pubescence on margin but densely covered with purplish-red pubescence on outer surface. Corolla is smaller than that of pistillate flowers, c. 1.9 cm in diameter, five petals, obovate, about 9 mm long and 4 mm broad, white, with numerous slender, light red, radiating longitudinal stripes extending to the apices of the petals, also branching at the end of the stripes. Filaments are white, c. 18 per flower, 6 mm long. Anthers are yellow, sagittate. Ovary is vestigial, tiny, densely covered with purplish pubescence.

Figure 2.7  Stem and leaf of Actinidia callosa var. callosa.

margins but otherwise nearly glabrous. Flowers are c. 2.2 cm in diameter, five petals, elliptical, c. 9 mm long and 4 mm broad, white, with light red radiating stripes ­extending to the apices of petals, distinct. Filaments are white, c. 27 per flower, 3 mm long. Anthers are yellow, sagittate. Styles are

Fruits  Nearly globose or short cylindrical. Fruit skin is green, glabrous; many lenticels elliptical or round, small, yellow-brown. Fruit apex is narrowly pointed, flatly truncated, emarginate in middle, styles persistent or shed, fruit shoulders are wide, round, nearly truncate, sepals persistent or shed, glabrous. Fruit stalk is light brown, glabrous and caducous, c. 1.9 cm long. Fruits are c. 2.3 cm long and 2.1 cm in diameter. The fruit weigh 8.7 g, fruit flesh is jade-green. The seed are small, elliptical, and

Chapter 2 Species

r­eddish-brown. The fruit ripen in mid-­S eptember, optimum sweet-acid flavor without astringent or peppery taste, juicy and crisp texture. Vc content is 16.6 mg per 100 g fresh weight. Soluble solid content is 7.8% and titratable acidity 1.16% (Figure 2.8).

Diagnostic characters and species relationships Readily recognized by conspicuous lenticels on 1-yearold stems, often lamellate pith, one- to three-flowered inflorescence and often globose and spotted fruits. Based on morphological characters, A. callosa is considered to be closely related to A. fortunatii, A. lanceolata, A. pilosula, A. trichogyna, A. venosa, and A. sabiifolia (Li, 1952; Liang, 1984; Li et al., 2007a,d). Molecular genetic data support a close relationship between A. callosa and A. ­lanceolata

Figure 2.8  Fruit of Actinidia callosa var. callosa.

Figure 2.9  Natural range of Actinidia callosa var. acuminata.

51

and A. pilosula but not with A. sabiifolia (Li et al., 2002a; Huang et al., 2002a; Chat et al., 2004; Li, 2006). Chromosomes: 2n = 2x = 58, 4x = 116

2.2b  Actinidia callosa var. acuminata 尖叶猕猴桃 jian ye mi hou tao

Actinidia callosa Lindley var. acuminata C.F. Liang in Fl.  Reipubl. Popularis Sin. 49(2): 314. 1984. TYPE: China. Hunan: Rucheng, July 16, 1960, Hunan Exped. 8569 (Holotype, HNNU). Distribution: China: Yunan and Hunan (Figure 2.9).

Morphological characteristics Stems The upper part of young shoots is white-green, lenticels are yellow-green, the basal part of current shoots is covered with long and yellow hispid hairs, with white or light brown, long, hispid hairs on the middle part, occasionally covered with pubescence, shed late in season. Old stems are gray-white to dark-brown, glabrous or covered with pubescence, lenticels are yellow-white; pith is lamellate, ­ white-green or brown-green. Leaves Chartaceous, oblong or broad lanceolate, leaf blade is c. 10.2-14.0 cm long and 4.8-7.2 cm broad, base flat to shallow cordate, nearly cordate, apex acuminate. The upper surface is light green to dark-brown. Leaf margin is serrated with wavy teeth pointed inward, red-brown and dense. Main vein and secondary veins sparsely covered with white hairs.

52

Kiwifruit: The Genus ACTINIDIA

Flowers Inflorescence is covered with long brown hairs.

As a variety of A. callosa, its species relationships are probably as for A. callosa var. callosa.

Fruit  Nearly long cylindric. Fruit skin is dark green, densely covered with alternately large and small white spots. Fruit shoulder is wide with persistent calyx, curling inward. The stalk is about 1.5-2.2 cm long, sparsely covered with yellow pubescence. The fruit average 1-1.5 g, mostly single. The fruit ripen in October to November. Vc content is 4.4 mg per 100 g fresh weight. Soluble solids content is 5.6% and titratable acidity 0.63%.

2.2c  Actinidia callosa var. discolor

Diagnostic characters and species relationships A. callosa var. acuminata differs from A. callosa var. callosa by its leaf having a long acuminate apex, and the outer surface of the sepals being covered by yellow-brown tomentum.

异色猕猴桃 yi se mi hou tao

Actinidia callosa Lindley var. discolor C.F. Liang in Fl.  Reipubl. Popularis Sin. 49(2): 315. 1984. TYPE: China. Guangxi: Hexian, in sparse forest, alt. 500 m, October 11, 1959, Z.T. Li 603855 (Holotype, IBK). Actinidia fanjingshanensis S.D. Shi & Q.B. Wang in Acta Bot. Yunnan. 16(4): 345. 1994. TYPE: China. Guizhou: Fanjingshan, alt. 580 m, August 15, 1991, Q.B. Wang et al. 91001 (Holotype, GBG; Isotype, KUN!). Distribution: China: Yunnan, Guangxi, Guangdong, Guizhou, Hunan, Jiangxi, Fujian, Zhejiang, Hubei, Anhui, and Taiwan (Figure 2.10).

Figure 2.10  Natural range of Actinidia callosa var. discolor.

Morphological characteristics Stems One-year-old shoots are brown and glabrous, lenticels are distinct, small, densely, elliptical, white, internodes c. 1.2-0.7 cm long. Two-year-old stems are red-brown, glabrous and slender, lenticels are distinct, small, sparsely, elliptical, light yellow; pith is lamellate, brown. Leaves Chartaceous and elliptical; leaf blade is c. 6.0-12.0 cm long and 3.5-6.0 cm broad, basal part cuneate, mostly bilateral symmetry; apex is relatively wide, mucronate or acuminate. The upper surface is dark green and glabrous. Leaf margin is serrated, more distinct and larger teeth in upper part, almost entire margin in basal part, tooth tips appressed or spreading. Main vein and secondary veins are

yellow-green, glabrous. The lower surface is green and glabrous, main vein and secondary veins are white-green and glabrous. The petiole is light green, glabrous, and 2.5-8.5 cm long (Figure 2.11). Pistillate flowers Cyme inflorescence mostly with three flowers. Peduncle is c. 1.4-1.6 cm long, green, sparsely covered with light purple-red pubescence, with white ­protuberances; pedicel is medium stout, light purple-red green, sparsely ­covered with light purple-red and white pubescence, the central pedicel is c. 10 mm long while lateral pedicels are c. 7 mm long. Calyx has five sepals, green, elliptical, c. 5 mm long and 2 mm broad, sparsely covered with light purple-red and white pubescence on

Chapter 2 Species

53

Figure 2.11  Stem and leaf of Actinidia callosa var. discolor.

both surfaces. The flowers are c. 2 cm in diameter, white, five petals are nearly oblong, c. 9 mm long and 5 mm broad, the petal base is narrow, slightly red, with dense radiating stripes up to apex, branching at tip. Filaments are white, 3 mm long, c. 27 per flower. Anthers are y­ ellow,

Figure 2.12  Flower and fruit of Actinidia callosa var. discolor.

relatively large, elliptical with deep, longitudinal dehiscing grooves. Styles are white, usually 20 per flower, 4 mm long, stout, with slightly swollen stigmata. Ovary is nearly globose, densely covered with light purple-red pubescence, 3 mm in diameter.

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Kiwifruit: The Genus ACTINIDIA

Staminate flowers Predominately single, pedicel is c. 1.6 cm long, green-white to light red, glabrous. Calyx of five sepals, oblong, c. 6 mm long and 2 mm broad, sparsely covered with light purple-red pubescence on inner surface, light brown pubescence on outer surface and relatively more densely covered with pubescence on the basal part of both surfaces. Flowers are c. 2.2 cm in diameter, five petals, white, with slight light purplish-red on the basal part, oblong, c. 1.4 cm long and 0.6 cm broad, with short and sparse longitudinal stripes only at the basal part. Filaments are white to light purple, c. 4 mm long, 27 per flower. Anthers are yellow, elliptical, with distinct longitudinal dehiscing grooves; ovary is vestigial small cone-shape, densely covered white and light purple pubescence. Fruit  Nearly ellipsoid, green fruit skin covered with short tomentum, with abundant fruit spots, yellow-brown, large, round or elliptical. Fruit apex is round and wide, with small and short rostrate; fruit shoulder is narrow, almost glabrous with persistent calyx. The stalk is c. 2.2 cm long, ­yellow-green and glabrous. Fruit are c. 2.4 cm long and 0.9 cm in diameter. The fruit is average 1.8 g; jade-green pulp with relatively small core. Numerous seed are relatively small, elliptical, purple-red with smooth seed coats. The fruit ripen in early October, acid flavor, very astringent and tart, little juice and soft pulp. Vc content is 12.54 mg per 100 g fresh weight. Soluble solid content is 11.6%, sugar content 2.14%, and acid content 1.04% (Figure 2.12).

Diagnostic characters and species relationships The variety is distinguished from A. callosa var. callosa by the different colors of the upper and lower leaf surfaces, especially in dried specimens when the lower surface

Figure 2.13  Natural range of Actinidia callosa var. henryi.

­ ecomes grayish yellow, and by the fruit being subglobose b to ovoid. As a variety of A. callosa, its relationships with other species are probably the same as those of A. callosa var. callosa. The close relationships between A. callosa var. discolor and A. lanceolata and A. pilosula are supported by the molecular genetic data (Li, 2006). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. callosa var. discolor in the National Germplasm Repository for Actinidia at WBG, were 0.1223 and 0.1758, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58, 4x = 116

2.2d  Actinidia callosa var. henryi 京梨猕猴桃 jing li mi hou tao

Actinidia callosa var. henryi Maximowicz in Trudy Imp. S.-Peterburgsk. Bot. Sada 11: 36. 1890. SYNTYPES: China. Hubei: Yichang, A. Henry 3471, 3494, 3955, 4377A; Badong, A. Henry724. Actinidia curvidens Dunn in Kew Bull. 1906: 1. 1906. TYPE: China. Hubei: Yichang, A. Henry 3564, 4377, 3494, 3955 (GH). Actinidia callosa var. ephippioidea C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 315. 1984. TYPE: China. Yunnan: Fengqing, alt. 2400 m, July 9, 1938, T.T. Yü 16621 (Holotype, PE). Distribution: China: Yunnan, Guangxi, Guangdong, Guizhou, Hunan, Jiangxi, Fujian, Zhejiang, Sichuan, Chongqing, Hubei, Anhui, Gansu, and Shaanxi (Figure 2.13).

Chapter 2 Species

Morphological characteristics Stems One-year-old shoots are green and glabrous; lenticels are distinct, small, sparsely, round or elliptical, yellow-white, internodes c. 2.0-6.0 cm long. Two-year-old stems are brown, glabrous and smooth, very slender, lenticels are distinct, dense and small, round or linear, white. Older stems are brown and glabrous, lenticels are not distinct, sparsely covered, elliptical and gray-brown; pith is lamellate, white. Leaves Chartaceous to thick chartaceous, broad lanceolate, leaf blade is c. 8.0-10.5 cm long and 4.0-5.5 cm broad, mostly

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bilateral symmetry, basal cuneate, apex mucronate. The upper surface is green, glabrous and lustrous. Leaf margin is serrated with dense, small teeth spreading or appressed, tip purplish color, almost entire in the basal part. Main vein and secondary veins are green, sparsely covered with light ­purplish-red short pubescence. The lower surface is light green, sparsely covered with white pubescence. Main vein and secondary veins are white-green and sparsely covered with long white pubescence. The petiole is medium stout, c. 2.4-4.9  cm long and sparsely covered with light ­purplish-red, short pubescence (Figure 2.14).

Figure 2.14  Stem and leaf of Actinidia callosa var. henryi.

Pistillate flowers  Cyme inflorescence, predominantly three flowers, occasionally single. Peduncle is short, slender, light purple-red and densely covered with light purple-red pubescence, while pedicel is slender, c. 6 mm long, light purple-red and densely covered with light purple-red pubescence. Calyx of five sepals, brown-green, elliptical, c. 5 mm long, 3 mm broad, and densely covered with purple-red pubescence on both surfaces. Flowers are c. 2.0 cm in diameter, five petals, occasionally six, white, nearly elliptical, c. 19 mm long and 8 mm broad, narrow at basal part with red color extending halfway up petal. Filaments are white, with light purple-red color at base, c. 4.5 mm long and 30 filaments per flower. Anthers are yellow, kidney-shaped, c. 2 mm long and 0.5 mm broad, with deep longitudinal dehiscing grooves. Styles are milky-white, c. 24 per flower, 3 mm long and becoming more stout from base to apex with slightly swollen stigmata, white, with some red. Ovary is elliptical, densely covered with ­purple-red pubescence, c. 4 mm long and 3 mm broad. Fruit Long cylindrical, with some irregularly shaped fruits and some are elliptic or ovate; fruit skin green to dark green, covered with irregular, relatively large, yellow-brown spots; both ends of the fruits are retuse with persistent calyxes. Fruit are c. 2.0-4.5 cm long and 1 cm in diameter. Fruit average 5.1-5.7 g, the largest in the species A. callosa, with green or dark green pulp and large core. Numerous seed are flat,

elliptical, yellow-brown. The fruit ripen in October; acid and astringent flavor, rich in pectin and less juicy. Vc content is 15.5 mg per 100 g fresh weight. Soluble solids content is 11%, sugar content 7.4%, and acid content 1.02% (Figure 2.15).

Diagnostic characters and species relationships It is distinguished from A. callosa var. callosa by its mammilliform, cylindrical fruit. As a variety of A. callosa, its species relationships may be similar to those of A. callosa var. callosa. In additional, A. callosa var. henryi appears to be closely related to A. obovata and A. hubeiensis (Liang, 1984; Sun and Huang, 1994). A close relationship to A. hubeiensis and A. pilosula is indicated by molecular genetic data (Chat et al., 2004; Li, 2006). The genetic diversity index of A. callosa var. henryi natural populations was 0.870 revealed by SSR markers (Liu et al., 2007), while the average genetic diversity (h) and haplotype genetic diversity (H) based on cpSSR markers was 0.087 and 0.546, respectively (Zhang et al., 2007). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. callosa var. henryi in the National Germplasm Repository for Actinidia at the WBG were 0.0118 and 0.0173 respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58, 4x = 116

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Kiwifruit: The Genus ACTINIDIA

Figure 2.15  Flower and fruit of Actinidia callosa var. henryi.

2.2e  Actinidia callosa var. strigillosa 毛叶硬齿猕猴桃 mao ye ying chi mi hou tao

Actinidia callosa var. strigillosa C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 315. 1984. TYPE: China. Guizhou: Kaili, alt. 750 m, May 8, 1959, South Guizhou Exped. 1398 (Holotype, PE).

are light green, glabrous, slightly concave from the surface. The lower surface is light green and mainly glabrous except sparsely strigillose, main vein and secondary veins are green-white, purplish-red on the petiolar end of the raised main vein. The petiole is purplish, sparsely covered with purplish-red pubescence, and c. 2.5-5.0 cm long (Figure 2.17).

Stems  One-year-old shoots are yellow-brown and glabrous; lenticels are protruded, elliptical or linear, gray-yellow or ­yellow-brown; internodes are 1.0-6.5 cm long. Two-year-old stems are dark-brown and glabrous; lenticels are protruded, elliptical, yellow-brown; pith is amellate, green.

Staminate flowers Predominately single, rarely cyme inflorescence with two flowers; pedicel is c. 0.5 cm long, green, glabrous, peduncle c. 0.8 cm long, green, glabrous. Calyx of six sepals, white-green, elliptical, c. 6 mm long and 5 mm broad, glabrous. Flowers are white. Corolla is c. 3.5 cm in diameter, six petals overlapping, ovate, c. 1.7 cm long and 1.4 cm broad, with longitudinal stripes. The filaments are white, 34-36 per flower and 9 mm long. Anthers are yellow, sagittate. Ovary is vestigial, tiny, cone-shaped, covered with white pubescence.

Leaves  Coriaceous, long ovate or nearly elliptical; leaf blade is c. 10-12.5 cm long and 6.5-8.5 cm broad, base obtuse, round and predominately bilateral symmetry, apex acuminate. The upper surface is green and glabrous, except in lateral vein axils. Main vein and secondary veins

Fruit Nearly globose. Fruit skin is green and glabrous with many large, round, light brown-yellow fruit spots. Fruit apex is flat and glabrous, with persistent styles; fruit shoulder is round, persistent sepals are relatively large, dark-brown, elliptical, glabrous. Fruit stalk is c. 2.4 cm long, reddish-brown,

Distribution: China: Yunan, Guangxi, Guizhou, and Hunan (Figure 2.16).

Morphological characteristics

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57

Figure 2.16  Natural range of Actinidia callosa var. strigillosa.

Figure 2.17  Stem and leaf of Actinidia callosa var. strigillosa.

glabrous. Fruit are c. 1.8 cm long and 1.9 cm in diameter. Fruit average 4.8 g, green pulp with medium-sized, elliptical, whitish core. Numerous seed are small, elliptical or round, purple-red. Fruit are slightly acid, astringent and tart taste, less juicy with slightly crispy texture (Figure 2.18).

Diagnostic characters and species relationships A. callosa var. strigillosa is distinguished from A. callosa var. callosa by its leaves being adaxially sparsely strigillose. As a variety of A. callosa, its specific relationships with other species are probably the same as those of A. callosa var. callosa. However, molecular genetic data (Chat et al., 2004; Li, 2006) showed that A. callosa var. strigillosa may

have originated by natural hybridization and that its genetic origin is in the A. chinensis species complex (Li, 2006). Chromosomes: 2n = 4x = 116

2.3  Actinidia chengkouensis 城口猕猴桃 cheng kou mi hou tao

Actinidia chengkouensis C.Y. Chang in Acta Sci. Nat. Univ. Szechuan. 1976: 74. 1976. TYPE: China. Chongqing: Chengkou, July 15, 1958, T.L. Dai 101228 (Holotype, SZ; Isotype, PE). Distribution: China: Sichuan, Chongqing, and Hubei (Figure 2.19).

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Kiwifruit: The Genus ACTINIDIA

Figure 2.18  Flower and fruit of Actinidia callosa var. strigillosa.

Morphological characteristics Stems One-year-old shoots are densely covered with brown setose. Two-year-old shoots are glabrous; pith is lamellate, brown. Leaves Chartaceous, broadly obovate, suborbicular, 6-12 cm long, 7-12 cm broad, short hispid on upper surface, conspicuously white-downy on lower surface. The veins are densely covered with brown, long hispid. Petioles are densely covered with brown hispid.

Figure 2.19  Natural range of Actinidia chengkouensis.

Pistillate flowers  Cyme inflorescence, one to three flowers. Calyx has four sepals. Flowers are white becoming yellow after, numerous stamens. Ovary is subglobose, densely pubescent. Fruit Fruit globose or ovoid, densely covered with yellowish hispid. Mature fruit unknown.

Diagnostic characters and species relationships The species is characterized by leaf underneath having hispid on midvein and lateral veins, and sparsely stellate tomentose.

Chapter 2 Species

A. chengkouensis is morphologically close related to A. chinensis var. deliciosa (Liang, 1984). It may be derived from a hybrid origin from A. chinensis var. deliciosa with another species.

2.4  Actinidia chinensis 中华猕猴桃 zhong hua mi hou tao

Actinidia chinensis Planchon in London J. Bot. 6: 303. 1847. TYPE: China. R. Fortune 39 (Holotype, K).

2.4a  Actinidia chinensis var. chinensis 中华猕猴桃(原变种) zhong hua mi hou tao (yuan bian zhong)

Actinidia chinensis var. rufopulpa (C.F. Liang & R.H. Huang) C.F. Liang & A.R. Ferguson in Guihaia 5(2): 72. 1985. A. chinensis var. chinensis f. rufopulpa C.F. Liang & R.H. Huang in Acta Phytotax. Sin. 20(1): 101. 1982. TYPE: China.

Figure 2.20  Natural range of Actinidia chinensis var. chinensis.

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Hubei: Wufeng, Chengguan Shawan, alt. 1150 m, August 31, 1980, R.H. Huang 4016 (Holotype, IBK). Actinidia chinensis var. jinggangshanensis (C.F. Liang) C.F. Liang & A.R. Ferguson in Guihaia 5(2): 72. 1985. A. chinensis var. chinensis f. jinggangshanensis C.F. Liang in Act. Phytotax. Sin. 20(1): 101. 1982. TYPE: China. Jiangxi: Yongxin, June 17, 1959, S.K. Lai 928 (Holotype, LBG). Actinidia multipetaloides H.Z. Jiang in J. Southwest Agric. Univ. 17(2): 93. 1995. TYPE: China. Yunnan: Suijiang, June 10, 1989, S.S. Sun & H. Zhang 89-3023 (Holotype, Herbarium, Institute of Horticulture, Yunnan Academy of Agricultural Sciences). Distribution: Occurs widely occurs in Central, Southeast China with greatest abundance in eastern Guizhou, Guangxi, Hunan, Jiangxi, Fujian, southwest Zhejiang, Hubei, southern and western Anhui, Henan, and Jiangsu (Figure 2.20).

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Kiwifruit: The Genus ACTINIDIA

Morphological characteristics Large deciduous climbing vines. Stems One-year-old shoots are gray-greenish brown and glabrous, or sparsely covered with white-downy hairs, readily shed; lenticels are relatively large, few, round or oblong, light yellowish-brown; internodes 3-6 cm long. ­Two-year-old stems are dark-brown, glabrous and c. 6 mm in diameter; lenticels are round or oblong, yellow-brown; pith is lamellate and very pale or creamy white. The over-wintering bud is exposed, covered only by bud scales with short, yellow-brown, downy hairs.

Leaves  Thick and papery; broadly ovate, nearly round and occasionally broadly obovate, leaf blade is 10-11 cm long and 11-14 cm broad; bilaterally symmetrical; basal cordate; apex round, truncate, minutely obtuse or retuse. The upper leaf surface is dark green and glabrous. Entire margin at leaf base, but on the upper parts the margin has small brown teeth terminating the veinlets. Main vein and secondary veins are glabrous and inconspicuous. The lower surface is grayish-green, densely covered with white stellate hairs; main and secondary veins are whitish-green, densely covered with white, very short pubescence. The petiole is light pinkish green, pubescence readily shed, and 8-9 cm long and c. 3 mm in diameter (Figure 2.21).

Figure 2.21  Stem and leaf of Actinidia chinensis var. chinensis.

Pistillate flowers Predominantly single, sometimes in cyme inflorescences with two to three flowers. Pedicel is c. 3-4 cm long, green color, covered by brown tomentum. Calyx has five to six elliptic or obovate sepals, c. 10 mm long and 7 mm broad, densely covered with yellow-brown tomentum. The flowers are c. 4 cm in diameter, white, becoming yellow to gold 1 day after opening; five to seven petals, nearly round at apex and c. 20 mm long, radiating stripes on the surface. About 155 stamens per flower. Filaments are white to light green; 8 mm long. Anthers are yellow, sagittate. Styles are white, usually 30 per flower, 6 mm long with slightly swollen stigma. Ovary is broadly globose, covered with white pubescence, 8 mm in diameter. The pistillate flowers produce a sweet fragrance. Staminate flowers Cyme inflorescence, two to three flowers, pedicel is c. 3.6 cm long. Calyx has four to six sepals, predominantly five to six, yellowish green and long obovate, overlapping. Opening flowers are white, becoming yellow to gold from 1 day after opening; flowers are smaller than pistillate flowers, c. 2.5 cm in diameter, four to six petals, predominantly six petals, broadly obovate, obtuse-round or weakly retuse at apex; wave-like wrinkles on the edge of the petal. The filaments vary in length, 40-47 per flower. Anthers are yellow. Ovary is vestigial, covered with white to pale brown pubescence.

Fruit Mostly ellipsoid or ovoid sometimes with blunt protrusion at stylar end, green or dark-yellow to brown skin covered with brown, fine tomentum, usually readily shed when the fruit mature. Fruit shoulder is round with persistent calyx. The stalk is about 3-7 cm long, green or greenish-brown, sparsely covered with light yellow tomentum. Fruit are c. 4 cm long and 3 cm in diameter. The fruit average 22 g, but those of selections from the wild can be much larger, >100 g. Yellow or green pericarp with small, round, white core, occasionally with red pigments around the core. Numerous seed in fruit, dark purple to almost black, elliptical with concave furrows in the seed coat. Fruit ripen in September, sweetly acid flavor, juicy and fine texture. Vc content is 50-420 mg per 100 g fresh weight. Soluble solid content is 7-19% and titratable acidity 0.9-2.2%. The fruit are suitable for fresh consumption and processing (Figure 2.22).

Diagnostic characters and species relationships Distinguished by broadly obovate leaves, leaf apex truncate or emarginated and fruits finely tomentose, but hairs easily shed. Over-wintering buds exposed, not buried in bark. Actinidia chinensis is a morphologically variable and is best treated as a species complex containing several closely related varieties. A. chinensis var. chinensis is very closely

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61

Figure 2.22  Flowers (pistillate, staminate) and fruit of Actinidia chinensis var. chinensis.

related to A. chinensis var. deliciosa and A. chinensis var. setosa. In addition, what appears to be a close relationship with A. obovata, A. grandiflora, A. stellatopilosa, A. hubeiensis, A. lijiangensis, and A. sorbifolia (Liang, 1984; Li et al., 2007a,d) is supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The natural population genetic diversity (PIC) of A. chinensis var. chinensis was >0.757 revealed by SSR markers (Li et al., 2004; Liu et al., 2007), while the average genetic diversity (h) and haplotype genetic diversity (H) based on cpSSR markers were 0.068 and 0.421 at the population level, 0.151 and 0.869 at the variety level, respectively (Zhang et al., 2007). The genetic diversity (PIC) of cultivars of A. chinensis var. chinensis was 0.89 based on SSR markers (Zhen et al., 2004). The Nei’s genetic diversity and the Shannon’s polymorphic information index of the live collection of

A. chinensis var. chinensis in the National Germplasm Repository for Actinidia at WBG, were 0.1450 and 0.2099, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58, 4x = 116

2.4b  Actinidia chinensis var. deliciosa 美味猕猴桃 mei wei mi hou tao

Actinidia chinensis var. deliciosa (A. Chevalier) A. Chevalier in Rev. Bot. Appl. Agric. Trop. 21: 241, 1941. A. latifolia (Gardner & Champion) Merrill var. deliciosa A. Chevalier in Rev. Bot. Appl. Agric. Trop. 20: 12, 1940. TYPE: “Fructification du même,” Planche II in Chevalier, 1940 (Lectotype, Pl. 2 in Chevalier, 1940: 15).

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Kiwifruit: The Genus ACTINIDIA

Actinidia chinensis var. hispida C.F. Liang in Acta Phytotax. Sin. 13(4): 33. 1975. TYPE: China. Guizhou: Fanjingshan, September 11, 1931, Q.Y. Jiao et al. 438 (Holotype, PE). Actinidia deliciosa (A. Chev.) C.F. Liang & A.R. Ferguson in Guihaia 4(3): 181. 1984. Actinidia deliciosa var. chlorocarpa (C.F. Liang) C.F. Liang & A.R. Ferguson in Guihaia 4(3): 182. 1984. A. chinensis var. hispida f. chlorocarpa C.F. Liang in Guihaia 2(1): 4. 1982. TYPE: China. Yunnan: Huize, September 1981, Z.Y. Yang & F.C. Tang1033 (Holotype, IBK). Actinidia deliciosa var. longipila (C.F. Liang et R.Z. Wang) C.F. Liang & A.R. Ferguson in Guihaia 4(3): 182. 1984.—A. chinensis var. hispida f. longipila C.F. Liang et R.Z. Wang in Act. Phytotax. Sin. 20(1): 102. 1982. TYPE: China. Shaanxi: Meixian, X.M. Zhang 707 (Holotype, IBK).

Figure 2.23  Natural range of Actinidia chinensis var. deliciosa.

Actinidia deliciosa var. coloris T.H. Lin & X.Y. Xiong in Guihaia 11(2): 117. 1991. TYPE: China. Hunan: Shimen, in shrubs, alt. 1200 m, October 3, 1990, X.Y. Xiong 001 (Holotype, IBK). Distribution: China: Yunnan, Guangxi, Guizhou, Hunan, Jiangxi, Sichuan, Chongqing, Hubei, Gansu, Shaanxi, and Henan (Figure 2.23).

Morphological characteristics Large, very vigorous deciduous vine. Stems  One-year-old shoots are green and covered with short, coarse, gray-brown hairs. Two-year-old stems are ­reddish-brown, glabrous; lenticels are white, round dot or

Chapter 2 Species

ovate shaped; internodes 1.5-3 cm long; pith is lamellate in older stems and pale brown. Leaves Papery to thick papery; nearly round and occasionally oblong, leaf blade is 8-12 cm long and 5.5-12.5 cm broad; bilaterally symmetrical; base lightly cordate or near cuneate; apex round, minute obtuse or retuse. The upper leaf surface is dark green and glabrous or puberulent. Leaf margin is

63

nearly entire and green with small spine-like awns spreading out. Main vein and secondary veins are yellowish green and main veins are sparsely covered by short yellow-brown tomentum. The lower leaf surface is light green, densely covered with pale yellow stellate hairs; main and secondary veins are yellow-green, covered with pale yellow tomentum. The petiole is sparsely covered with brown tomentum, 4.08.5 cm long (Figure 2.24).

Figure 2.24  Stem, emerging shoot, and leaves of Actinidia chinensis var. deliciosa.

Pistillate flowers  Predominantly single; pedicel is c. 4 cm long, green, covered with pale brown tomentum. Calyx has five to six elliptical or obovate sepals, c. 8 mm long and 7 mm broad, densely covered with light brown tomentum. The flowers are c. 5.5 cm in diameter, white, changing to yellow to apricot-yellow 1 day after opening; six to seven petals, predominantly six, obovate, c. 25 mm long and 20 mm broad, with longitudinal stripes on the surface. Filaments are white, 8 mm long, c. 150 filaments per flower. Anthers are yellow, sagittate. Styles are white, usually 37 per flower, 5 mm long with slightly swollen stigma. Ovary is short cylindrical, 6 mm in diameter, covered with white to light brown pubescence. The pistillate flowers produce a sweet fragrance. Staminate flowers Cyme inflorescence, two to three flowers; green pedicel, c. 2.5 cm long, covered by pale brown tomentum. Calyx has four to six sepals, predominantly six, light brown, covered by pale brown tomentum. Flowers are white when first opening, changing to yellow after 1 day; flowers are smaller than pistillate flowers, c. 4 cm in ­diameter, four to six petals, predominantly six, obovate, c. 20 mm long and 15 mm broad, with longitudinal stripes on the surface. The filaments vary in length, 200 per flower. Anthers are yellow, sagittate. Ovary is vestigial, tiny cone, covered by light brown tomentum. Fruit Mostly ellipsoid or cylindrical or ovoid. The fruit skin is green to brown, densely covered with long yellow-brown

hispid hairs, not readily shed when the fruit mature. Fruit apex is protruded, nearly round, often narrower than the middle part of the fruit. Calyces are persistent. The stalk is dark green, about 5-7 cm long. Fruit are c. 6.3 cm long and 3.2 cm wide, cross section often oval. The fruit average 34 g, but individual vines can have much larger fruit, some commercial cultivars up to 200 g. Fruit pericarp bright green with small, round, creamy white core. Occasionally, inner pericarp around core has red pigments. The numerous seeds in fruit are dark purple to black, elliptical with concave furrows in the seed coat. Fruit ripen in October to November, sweetly acid flavor, juicy and fine texture. Vc content is ­50-240 mg per 100 g fresh weight. Soluble solid content is 8-25% and total acidity 1.6%. The fruit are suitable for fresh consumption and processing (Figure 2.25).

Diagnostic characters and species relationships The variety is characterized by flowering branchlets and petioles being covered by yellow-brown strigose hairs, leaves broadly ovate or obovate and fruits densely hispid. Overwintering buds deeply buried in the bark. As a variety of A. chinensis, it is very closely related to A. chinensis var. chinensis and A. chinensis var. setosa. Its relationships with other species may be as the same as those of A. chinensis var. chinensis (Liang, 1984; Li et al., 2007a,d), as supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006).

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Kiwifruit: The Genus ACTINIDIA

Figure 2.25  Flowers (pistillate, staminate) and fruit of Actinidia chinensis var. deliciosa.

The natural PIC of A. chinensis var. deliciosa was >0.781 revealed by SSR markers (Li et al., 2004; Liu et al., 2007), while the average genetic diversity (h) and haplotype genetic diversity (H) based on cpSSR markers was 0.014 and 0.114 at population level, 0.041 and 0.516 at the taxon level, respectively (Zhang et al., 2007). The genetic diversity (PIC) of the cultivars group of A. chinensis var. deliciosa was 0.91 based on SSR markers (Zhen et al., 2004). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. chinensis var. deliciosa in the National Germplasm Repository for Actinidia at WBG, were 0.1450 and 0.2099, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 4x = 116, 6x = 174, 8x = 232, occasionally 12x = 358

2.4c  Actinidia chinensis var. setosa 刺毛猕猴桃 ci mao mi hou tao

Actinidia chinensis var. setosa H.L. Li in J. Arnold. Arbor. 33: 56. 1952. TYPE: China. Taiwan: Mt Arisan, E.H. Wilson 10802 (Holotype, US). Actinidia setosa (H.L. Li) C.F. Liang & A.R. Ferguson in Guihaia 5(1): 72. 1985.

Distribution: China: Mt Arisan of Taiwan (endemic) (Figure 2.26).

Morphological characteristics Strong deciduous climber. Stems One-year-old shoots are covered with brownish red stiff hairs. Two-year-old stems are brownish red with oblong lenticels; pith is lamellate, white or pale yellow. Leaves Papery: nearly round; leaf blade is 12-17 cm long and 10-15 cm broad; retuse to mucronate at apex and round or cordate at base. Margin is serrated as long and thin teeth. The upper leaf surface is covered with scabrid-hispid pubescence. The lower surface is silvery-gray, densely covered with white stellate hairs; main and secondary veins are ­whitish-green, densely covered with white stellate hairs. The petiole is covered with ferruginous hispid-setose pubescence, and c. 3.5-7.5 cm long (Figure 2.27). Flowers  Cyme inflorescence with few flowers. Pedicel is c. 1-2 cm long, with small bracts on which there are stripes on the surface. Calyx has five sepals, c. 14-15 mm long and 10-12 mm broad; broadly obovate, round at apex with short

Chapter 2 Species

65

Figure 2.26  Natural range of Actinidia chinensis var. setosa.

Figure 2.27  Stem, young breaking shoot, and leaves of Actinidia chinensis var. setosa.

fingernail-like tips. The flowers are c. 0.8-1.0 cm in diameter, opening white but turning yellow-orange; five petals, sometimes three to four; long ovate, c. 8-10 mm long. Styles are c. 5-6 mm long with slightly swollen stigma. Ovary is nearly globose, 6-7 mm in diameter, densely covered with white pubescence.

core. Numerous seeds in fruit, long, elliptical. The fruit ripen in early July, light sweetly acid flavor and juicy. VC content is 79.5 mg per 100 g fresh weight. Soluble solids content is 10.5% and total acidity 1.3% (Figure 2.28).

Fruit  Mostly nearly globose to ellipsoid, c. 2-3 cm across. The skin is densely covered with long, stiff, coarse brown hairs. The fruit average 28 g; pericarp light green with large

The variety is characterized by flowering branchlets and petioles being covered by stiff, reddish-brown hairs and the fruit being densely hispid.

Diagnostic characters and species relationships

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Kiwifruit: The Genus ACTINIDIA

Figure 2.28  Flowers and fruit of Actinidia chinensis var. setosa.

As a variety of A. chinensis, it is also very closely related to A. chinensis var. chinensis and A. chinensis var. deliciosa. Its relationships with other species may be as the same as A. chinensis var. chinensis and others (Liang, 1984; Li et al., 2007a,d), as supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The Nei’s genetic diversity and the Shannon’s polymorphic information index of A. chinensis var. setosa plants growing in the National Germplasm Repository for Actinidia at WBG were 0.0158 and 0.0230, respectively, as revealed by AFLP markers (Li, 2006).

2.5  Actinidia chrysantha

Chromosomes: 2n = 2x = 58

Large deciduous climber.

Figure 2.29  Natural range of Actinidia chrysantha.

金花猕猴桃 jin hua mi hou tao

Actinidia chrysantha C.F. Liang in Fl. Reipubl. Pop. Sin. 49: 318. 1984. TYPE: China. Guangxi: Lingui, in shrubs, June 23, 1952, C.F. Liang 30398 (Holotype, IBK). Distribution: China: Guangxi, Guangdong, Jiangxi, Hunan, and Fujian (Figure 2.29).

Morphological characteristics

Chapter 2 Species

Stems  One-year-old shoots are gray-brown, covered with sparse, tea-brown downy hairs. Two-year-old stems are ­reddish-brown, lenticels are distinct; pith is lamellate, light brown.

Figure 2.30  Stem, breaking shoot, and leaves of Actinidia chrysantha.

Figure 2.31  Pistillate flowers and fruit of Actinidia chrysantha.

67

Leaves Thin, papery, round or ovate; lamina is 7-14 cm long and 4.5-6.5 cm broad Upper leaf surface is grassy green or green, glabrous. Lower surface is glaucous and slightly rusty, and ­glandular-tomentose on midvein and lateral veins when young.

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Kiwifruit: The Genus ACTINIDIA

The petiole is light red and glabrous. Leaf margin is serrated with small teeth when young but crenate when old (Figure 2.30). Pistillate flowers  Cyme inflorescence, one to three flowers per inflorescence. The flowers have five petals, c. 1.5 cm long and 1.8 cm broad, bright yellow. About 35 stamens per flower. Anthers are yellow. Fruit Short cylindrical, ovoid or globose. Fruit apex protruded or slightly protruded, persistent calyx. Fruit skin is maroon brown or brownish-green, glabrous. Obvious and abundant fruit spots are brown or yellowish-brown, round or square, medium size. Fruit average 10-30 g, 2.4-4.0 cm long and 2.2-3.1 cm diameter. Green or light green flesh, fine texture, medium-sized core, juicy, sweetly acid taste with aroma. 100-250 seed per fruit, seed are c. 2 mm long, 1.3-1.7 g per 1000 seeds. Vc content is 34-57 mg/100 g fresh weight, total sugar content is 8.3%, and acid content is 1.3%. Fruit ripen in November in Guangxi (Figure 2.31).

Diagnostic characters and species relationships

this is not supported by molecular genetic data (Li, 2006). The natural PIC of A. chrysantha was from 0.653 to 0.830 revealed by SSR markers (Liu et al., 2007), while the average genetic diversity (h) and haplotype genetic diversity (H) based on cpSSR markers was 0.134 and 0.604, respectively (Zhang et al., 2007). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living A. chrysantha resources in the National Germplasm Repository for Actinidia at WBG were 0.0828 and 0.1209, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 4x = 116

2.6  Actinidia cylindrica 柱果猕猴桃 zhu guo mi hou tao

Actinidia cylindrica C.F. Liang in Fl. Reipubl. Popularis. Sin. 49(2): 315. 1984. TYPE: China. Guangxi: Rongshui, near mountain road, alt. 600 m, June 6, 1957, D.Z. Chen 476 (Holotype, IBK).

The species is distinguished by the underneath of the leaves being glaucous with glandular-fine hairs on midvein and lateral veins, leaf margin crenate and flowers yellow.

2.6a  Actinidia cylindrica var. cylindrica

Actinidia chrysantha was considered, morphologically, to be closely related to A. indochinensis (Liang, 1984), but

Distribution: China: Rongshui, Guangxi (endemic) (Figure 2.32).

柱果猕猴桃(原变种) zhu guo mi hou tao (yuan bian zhong)

Figure 2.32  Natural range of Actinidia cylindrica.

Morphological characteristics Small to medium-sized, semi-evergreen climber. Stems One-year-old shoots are relatively slender, light green, light red-purple on upper surface of the shoots; internodes

are c. 1.3-3.0 cm long. The staminate plants are covered with sparse tomentum. Older stems are dark to yellow-brown or gray-brown. Stems of pistillate plants are glabrous, relatively rough, lenticels distinct, sparse, oblong or linear shape, and light pale gray; pith is lamellate and white.

Chapter 2 Species

Figure 2.33  Stem and leaf of Actinidia cylindrica.

Leaves Leaves of pistillate plants are wider than those of staminate plants; thick, becoming leathery, lanceolate, leaf blade is c. 5-13 long and 2.5-5.5 cm wide; apex acute or acuminate and basal subrounded. The upper surface is green and lustrous. Leaf margin is finely serrulate (very small teeth) with tiny red spines, shallowly crenate-serrate on 2-year leaves. The lower surface is glabrous, the petiole and veins on lower surface of staminate plants are covered with sparse tomentum. The petiole is dark light purple, tomentose, c. 1.31.5 cm long and 2-2.8 mm in diameter (Figure 2.33). Pistillate flowers  Cyme inflorescence, mostly with one to three flowers. Pedicel is c. 8-10 mm long and 1 mm in diameter. Calyx has five, sometimes six, sepals are short, elliptical, c. 6 mm long and 4 mm broad. The flowers are c. 1.4 cm in diameter, red, five to six petals, sometimes two to four, rounded, acute rounded at basal, invaginated, with white longitudinal stripes, glabrous. Stamens 34-37 per flower, filaments are c. 1.0-2.1 mm long. Anthers are yellow, long, ­deltoid-shaped, obtuse ended and c. 1.5 mm long. Styles are c. 1-2 cm long, 16-22 per flower, with slightly swollen stigma, nearly round. Superior ovary is long, pubescent, cylindrical. Staminate flowers  Predominantly single, rarely two flowers per inflorescence, pedicel is c. 9 mm long, glabrous, light green-white. Calyx has predominantly five, occasionally four, sepals c. 4-5 mm long, elliptic and glabrous, with retuse longitudinal stripes on the surface, ridged between stripes. Corolla is of similar size to that of pistillate flowers, pink-red, four to five petals, elliptical, the middle part is relatively narrow and invaginated, ladle-like, c. 7 mm long and 6 mm broad. Stamens, 24 per flower, filaments are white, c. 2-5 mm long. Anthers are yellow, long shape, mucronate at apex and c. 1.6 mm long. Rudimentary ovary, milky-white and overshadowed by stamens. Fruits Long cylindrical, yellow-copper green skin is smooth and glabrous, fruit spots are small and dense, round or elliptic raised shapes, brown. Fruit apex is slight protruded, with calyx persistent, very small, deltoid, dark-brown and glabrous. The stalk is readily shed. Fruit are about 1.9-2.2 cm

Figure 2.34  Pistillate flowers and fruit of Actinidia cylindrica.

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Kiwifruit: The Genus ACTINIDIA

long and 8-9 mm in diameter. The fruit average 0.5-1 g, light emerald-green pulp with small, green core. Small seeds are black and flat oblong shape. The fruit ripen in mid-October, sweetly acid flavor and juicy. Vc content is 28.4 mg per 100 g fresh weight. Soluble solid content is 12-13.2%, sugar content 6.1% and acid content 1.28% (Figure 2.34).

Diagnostic characters and species relationships It is characterized by inconspicuous lenticels on branchlets, small lamellate pith, and consistently cylindrical fruits with unreflexed sepals. Morphologically, Actinidia cylindrica is considered to be related to A. fortunatii and A. henryi (Liang, 1984; Li et al., 2007a,d), and this is supported by molecular genetic

data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). Chromosomes: 2n =2x = 58

2.6b  Actinidia cylindrica var. reticulata 网脉猕猴桃 wang mai mi hou tao

Actinidia cylindrica var. reticulata C.F. Liang in Guihaia 2(1): 3. 1982. TYPE: China. Guangxi: Rongshui, alt. 670 m, October 2, 1981, Y. Li 3003 (Holotype, IBK). Distribution: China: limited distribution in Guangxi and Guizhou (Figure 2.35).

Figure 2.35  Natural range of Actinidia cylindrica var. reticulata.

Morphological characteristics Stems Current shoots are light green, smooth and glabrous, upper part being slender. One-year-old shoots and old stems are dark-brown with slight luster. Lenticels are small, relatively numerous, round or short linear shapes. One-year-old shoots of male plants are c. 3.5 mm in diameter; internodes c. 2.5-8.5 cm long while internodes of female plants are c. 5.2-15.5 cm long.

Leaves Thick and becoming leathery; oblong, lanceolate or broad lanceolate; leaf blade is c. 5.5-18.7 cm long and 7.58.3 cm broad; basal obtuse-rounded; retuse near the petiole, apex acuminate or mucronate. The upper surface is darkbrown or green, glabrous. Relatively large serrations on leaf margin are irregular and ciliate. Main vein, secondary veins and small veins are invaginated, making vein-between areas

Chapter 2 Species

elevated from the surface. The lower surface is light green, veins are distinctly protruded from the surface, vein meshes also distinct. In male plants, the lower leaf surface is densely covered with stellate hairs which have a tiny protuberance at

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the basal part of the stellate hair clusters, the hairs spreading; the stellate hairs of female plants are inconspicuous. The petiole is light green with brown rusty spots on the surface toward to sun, and c. 3-4 cm long (Figure 2.36).

Figure 2.36  Stem and leaves of Actinidia cylindrica var. reticulata.

Pistillate flowers  Dichasium inflorescence, two to seven flowers per inflorescence, the apical flower opens first, and the corolla opens entirely. Pedicel is c. 1.0-2.3 cm long, light green, glabrous, uppermost pedicel is c. 1.3 cm long while lateral pedicels are c. 7 mm long. Calyx has five sepals which are elliptic or mucronate at apex with wrinkled strips, pale white. The flowers are pink to red, petals are nearly round with basal part relatively narrow. Stamens c. 40-50 per flower, filaments are white, c. 2.3 mm long. Anthers are yellow, long shape with slightly acute apex, medium dehiscing grooves, c. 1.3 mm long and 0.5 mm broad. Styles are white, 27-34 per flower, 2.5-3.0 mm long with slightly swollen stigma. Ovary is elliptical, white and glabrous, c. 4 mm long and 3 mm in diameter.

truncate, pistils persistent and very distinct. Fruit stalk is c. 7-10 mm long, green brown color, with slender middle part. Fruit average 2.1 g, dark green pulp with small and green core. Seeds are very small, elliptic, slightly flat, blackbrown in color. The fruit ripen in mid-October, sweetly acid flavor and juicy. Vc content is 146 mg per 100 g fresh weight. Soluble solid content is 16.9%, sugar content 4.36%, and acid content 1.0% (Figure 2.37).

Staminate flowers  Dichasium inflorescence, occasionally cyme inflorescence, two to seven flowers, peduncle is c. 2.2 cm long and 1.2 mm in diameter, apical pedicel is c. 1.1 cm long, while lateral pedicels are c. 0.4 cm long, light green. Calyx has predominately five sepals, occasionally four or six, nearly white, oblong, obtuse at apex, concave and glabrous. Opening flowers are pink; corolla is c. 1.3-2.0 cm in diameter, petals are elliptic, with basal part relatively narrow, invaginated. Stamens 40-50 per flower, filaments are c. 4 mm long, slender, light pink near to petals. Pistil is rudimentary, overshadowed by stamens. Styles are white, glabrous, five per flower, and c. 0.5 mm long with slightly swollen stigma. Ovary is c. 1.2 mm long and 1.0 mm in diameter.

As a variety of A. cylindrica, the relationships with other species are probably the same as those of A. ­cylindrica var. cylindrica. In addition, it appears also to be closely related to A. fortunatii and A. henryi, and this is supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. cylindrica var. reticulate in the National Germplasm Repository for Actinidia at WBG were 0.0670 and 0.0978, respectively, as revealed by AFLP markers (Li, 2006).

Fruits Short cylindrical or nearly round, dark yellow-­copper green fruit skin, fruit spots are very small, round or slightly long round, light green brown color. The middle part of the fruits is relatively narrow; calyx is relatively large, usually covering fruit shoulders; fruit apex is slight protruded, nearly

Diagnostic characters and species relationships A. cylindrica var. reticulata differs from A. cylindrica var. cylindrica by the lower leaf surface having very ­distinct midvein and lateral veins, and reticulate veinlets conspicuous.

2.7  Actinidia eriantha 毛花猕猴桃 mao hua mi hou tao

Actinidia eriantha Bentham in J. Proc. Linn. Soc., Bot., 5: 55. 1861. TYPE: China. Fujian, Lindley 1836 (Holotype, K). Actinidia davidii Franchet in Nouv. Arch. Mus. Paris Sér. 2, 5: 57. 1884.

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Figure 2.37  Pistillate flowers and fruit of Actinidia cylindrica var. reticulata.

Actinidia eriantha f. alba C.F. Gan in Guihaia 3(1): 18. 1983. TYPE: China. Zhejiang: Qingyuan, May 22, 1982, C.F. Gan 101 (Holotype, IBK). Actinidia eriantha var. calvescens C.F. Liang in Guihaia 11(2): 118. 1991. TYPE: China. Guangxi: Guilin, May 9, 1987, C.F. Liang 34511 (Holotype, IBK).

Figure 2.38  Natural range of Actinidia eriantha.

Actinidia eriantha var. brunnea C.F. Liang in Guihaia 11(2): 118. 1991. TYPE: China. Guangxi: Guilin, May 15, 1987, C.F. Liang 34514 (Holotype, IBK). Distribution: China: Yunnan, Guangxi, Guizhou, Hunan, Jiangxi, Fujian, and Zhejiang (Figure 2.38).

Chapter 2 Species

Morphological characteristics Large, very vigorous, deciduous climber. Stems One-year-old shoots are yellowish-brown and densely covered by a thick layer of milky-white to dirty yellow tomentum; lenticels are indistinct; internodes 2.9-4 cm long. Two-year-old stems are brown, sparsely covered with white ash-like downy hairs, indistinct lenticels, and c. 8 mm in diameter; pith is white, lamellate. Leaves  Thick and chartaceous, somewhat brittle and easily damaged, ovate, occasionally cone-shaped; leaf blade is

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8-16 cm long and 6-11 cm broad; bilaterally slightly asymmetric; acuminate or shortly obtuse at apex. The upper leaf surface is dark green and glabrous, shiny; main and secondary veins are green and glabrous. The marginal serrations are not distinct, only with small, light green, spine-like awns spreading outward. The lower leaf surface is light ­grayish-green, densely covered by white stellate tomentum and pubescence; main and secondary veins are whitish-green, densely covered with long white tomentum. The petiole is white to yellowish-brown, c. 1.5-3.1 cm long (Figure 2.39).

Figure 2.39  Stem, leaves, and flower buds of Actinidia eriantha.

Pistillate flowers Cyme inflorescence with usually three flowers; peduncle 5-10 mm long, pedicels c. 3-5 mm long, white grayish-green, densely covered with white tomentum. Calyx has two, usually three sepals, broadly ovate, c. 7-9 mm long, densely covered with milky-white to pale brown tomentum. Flowers are c. 4 cm in diameter, usually rose pink, sometimes white, often deep red; five to six petals, nearly obovate and c. 15 mm long, 10 mm broad. Stamens 115-130 per flower; filaments are pink, 5-7 mm long; anthers are yellow, sagittate. Styles, usually 37-39 per flower, 3-4 mm in length, are white, with slightly swollen stigma. Ovary is nearly globose or elliptical, densely covered with white tomentum, 7 mm in diameter. Staminate flowers  Cyme inflorescence with usually three flowers. Peduncle is c. 8-10 mm and the terminal pedicel is c. 13 mm and secondary pedicels are 0.5-1 mm long. Calyx has three triangular sepals, c. 11 mm long, densely covered by white tomentum. Flowers when first opening are usually pink aging to orange pink, c. 4 cm in diameter, six to seven petals, long elliptical, c. 17 mm long and 11 mm broad. Stamens, c. 160 per flower, filaments are pink, anthers are yellow, oblong, blunt at apices. Ovary is rudimentary and densely covered with white tomentum.

Fruit Mostly long cylindrical or ovoid, densely covered by milky-white hairs, sometimes pale brown, usually persistent but sometimes molting as fruit mature. Fruit skin is green, densely covered with small golden-yellow spots; sepals persistent. Fruit apex is nearly flat shaped with retuse center. Fruit shoulder is nearly flat cut or rounded shape; calyx is persistent. The stalk, c. 1.9 cm long, is densely ­covered with white tomentum. Fruit are c. 4 cm long and 2.2 cm in diameter. Fruit average 30-50 g but some selections have considerably larger fruit; fruit pulp very dark green with small and round core. Numerous seed, brown, flat-elliptical in shape. Fruit ripen mid-late September, acid flavor, juicy and fine texture. VC content is exceptionally high, 561-1379 mg per 100 g fresh weight. Soluble solid content is 5-16% and total acidity 1.3-2.9%. The fruit is suitable for fresh consumption and for processing and has considerable commercial potential (Figure 2.40).

Diagnostic characters and species relationships The species is distinct in that branchlets, buds, abaxial leaf surface, petioles, inflorescences, sepals, and fruit always have a white indumentum; the flowers are usually pink to red. Morphologically, A. eriantha appears to be related to A. fulvicoma, A. chinensis and A. zhejiangensis (Li, 1952;

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Kiwifruit: The Genus ACTINIDIA

Figure 2.40  Flowers and fruit of Actinidia eriantha.

Liang, 1982b, 1984; Li et al., 2007a,d), but molecular genetic data does not support a close relationship between A. eriantha and A. chinensis or A. fulvicoma whereas it does support a close relationship between A. eriantha and A. zhejiangensis in maternal lineage (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The natural PIC of A. eriantha ranges from 0.705 to 0.827 revealed by SSR markers (Liu et al., 2007, 2010), while the average genetic diversity (h) and haplotype genetic diversity (H) based on cpSSR markers was 0.063 and 0.333, respectively (Zhang et al., 2007). The Nei’s genetic diversity and the Shannon’s ­polymorphic information index of living A. eriantha resources in the National Germplasm Repository for Actinidia at WBG were 0.0986 and 0.1425, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58

2.8  Actinidia farinosa 粉毛猕猴桃 fen mao mi hou tao

Actinidia farinosa C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 319. 1984. TYPE: China. Guangxi: Tianlin, Naxiang Town, in grass near road, June 14, 1955, Z.T. Li 600772 (Holotype, IBK).

Distribution: China: limited distribution in two counties, Tianlin and Loucheng, Guangxi (endemism) (Figure 2.41).

Morphological characteristics Medium vigorous, semi-evergreen climber. Stems  New flowering shoots are densely covered by ­yellowish-brown woolly hair, readily shed, lenticels not obvious. Two-year-old stems are sparsely covered with ­residual coarse hairs and small, almost invisible lenticels. Pith is lamellate and dirty white. Leaves Broadly ovate or ovate, apex acute or short acuminate, leaf base slightly cordate, not symmetrical bilaterally. Petiole, 3.5-4 cm, has dense brownish woolly tomentum, readily shed; leaf lamina broadly ovate to ­orbicular-ovate, 9-15  × 7-9.5 cm; papery; upper leaf surface pale green, shiny, and the conspicuous midrib, secondary veins and veinlets have stiff, appressed bristly hairs; lower leaf surface appears glaucous and has thick brownish cottony-tomentum, readily shed, and slender, bristly hairs on midvein and lateral veins. Leaf margin, often pinkish, has indistinct teeth, small wavy teeth toward apex.

Chapter 2 Species

Figure 2.41  Natural range of Actinidia farinosa.

Figure 2.42  Stem, leaves, staminate flowers, and fruit of Actinidia farinosa.

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Kiwifruit: The Genus ACTINIDIA

Pistillate flowers  Usually single, sometimes up to three in inflorescence; calyx has five ovate sepals, 4-5 × 2.5 mm, densely tomentose on outer surface, glabrous on inner surface; corolla pink or white, five petals obovate to oblong obovate, 4-7 × 4-5 mm; filaments are c. 4 mm in length and anthers 1.5 mm; ovary is cylindrical, c. 2 mm, covered in dense brownish tomentum; styles c. 0.5 mm.

diversity and the Shannon’s polymorphic information index of living collection A. farinosa in the National Germplasm Repository for Actinidia at WBG were 0.0513 and 0.0749, respectively, as revealed by AFLP markers (Li, 2006).

Staminate flowers  Inflorescences cymose, one to three flowers, densely tomentose; peduncles very short 2-4 mm but pedicels 0.5-1.5 cm; single tapering bract, c. 4.5 mm, diamond-shaped, with dense long tomentum, readily shed. Calyx has five ovate sepals, 4-5 × 2.5 mm, densely tomentose on outer surface, glabrous on inner surface; corolla pink or white, five petals obovate to oblong obovate, 4-7 × 4-5 mm. About 40 stamens, filaments are c. 4 mm long, anthers brown, c. 1.5 mm.

2.9  Actinidia fasciculoides

Chromosomes: 2n = 2x = 58

簇花猕猴桃 cu hua mi hou tao

Actinidia fasciculoides C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 312. 1984. TYPE: China. Yunnan: Xichou, September 11, 1947, K.M. Feng 11643 (Holotype, PE).

2.9a  Actinidia fasciculoides var. fasciculoides 簇花猕猴桃(原变种) cu hua mi hou tao (yuan bian zhong)

Fruit  Small fruit, ovoid-cylindrical, hairs eventually shed to become glabrous, olive-green skin, small pale lenticels (Figure 2.42).

Distribution: China: Guangxi and Yunnan (Figure 2.43).

Diagnostic characters and species relationships

Morphological characteristics

The species is characterized by broadly ovate leaves, the lower surface of which has thick brownish cottony-­ tomentum, the hairs readily shed, and pink flowers. Morphologically, Actinidia farinosa is considered to be related to A. fulvicoma var. hirsuta (Liang, 1984; Li et  al., 2007a,d), and this is supported by molecular genetic data in the paternal lineage (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The Nei’s genetic

Figure 2.43  Natural range of Actinidia fasciculoides var. fasciculoides.

Large vigorous deciduous climber. Stems  One-year-old shoots are green or light purple, covered with light purple pubescence; lenticels are distinct, white, elliptical or linear in shape. Two-year-old stems are blackish brown, glabrous; lenticels are distinct, ­grayish-white, oblong or linear; pith is solid, brown. Older stems are r­eddish-brown; lenticels are large, white, linear, or round.

Chapter 2 Species

Leaves Petiole is purplish-red, sparsely covered with short, purplish pubescence, about 2.5-3.5 cm long. Leaf lamina is chartaceous, oblong or obovate; c. 6.5-9.5 cm long, 4.56.5 cm broad; basal part is round, usually bilaterally symmetrical; leaf apex is acute; upper surface of the leaf is dark green and shiny; midvein is pale green; up to seven secondary veins on each side of the primary vein, concave from the surface. Lower leaf surface is green, covered with white hispid hairs; purplish-red veins, the primary vein is covered with purplish-red pubescence and secondary veins with stiff white hairs. Leaf margin is serrated with small but conspicuous spine-like, purplish-red teeth, Young developing leaves are purplish-red (Figure 2.44). Pistillate flowers  Cyme inflorescences of one to six flowers, pedicle glabrous. Calyx has five sepals, broadly ovate, acute at apex, 4-5 mm long, sparsely covered with red tomentum. The flowers are light pink, 15-18 mm in diameter, five petals, broadly obovate, 8 mm long, round at apex. Ovary is cylindrical or ovoid, 2-3 mm long, style 3-4 mm long. Staminate flowers  Cyme inflorescences of up to six flowers, clustered. Pedicel is c. 6-14 mm long, sparsely covered with red pubescence. Calyx has five sepals, broadly ovate, red coloration toward apex, sparsely covered with

Figure 2.44  Stem, emerging shoots, and leaves of Actinidia fasciculoides.

Figure 2.45  Flower of Actinidia fasciculoides.

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red ­pubescence. The flowers are white, c. 14-16 mm in ­diameter; five petals, ladle shape, obtuse at apex. Anthers yellow (Figure 2.45). Fruit Long, cylindrical, 1.5-2 cm long, emarginated at fruit apex. Fruit skin is very dark green, with obvious brown lenticels nearly round or irregularly shaped, glabrous. Dark green pulp with numerous small seed. The fruits ripen September to October, Vc content is 7.3 mg/100g fresh weight and 0.33% acid.

Diagnostic characters and species relationships The species is characterized by the brownish solid pith and two- to six-fruited infructescence with very short peduncle. Actinidia fasciculoides was considered to be related to A. umbelloides (Liang, 1984).

2.9b  Actinidia fasciculoides var. cuneata 楔叶猕猴桃 xie ye mi hou tao

Actinidia fasciculoides var. cuneata C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 313. 1984. TYPE:China. Guangxi: Tianlin, alt. 800 m, November 24, 1959, Q.B. Huang 41099 (Holotype, IBK).

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Kiwifruit: The Genus ACTINIDIA

Figure 2.46  Natural range of Actinidia fasciculoides var. cuneata.

Distribution: China: Yunnan and Guangxi (Figure 2.46).

Morphological characteristics

As a variety of A. fasciculoides, it is closely related to A. umbelloides.

Stems  One-year-old stems are green, sparsely covered with purplish-red and white pubescence; lenticels are white, elliptical. Two-year-old stems are reddish-brown, glabrous; sparse lenticels are white, elliptical; internodes are about 9-22 cm long. Old stems are grayish brown and glabrous, lenticels are not distinct; pith is solid, brown.

2.9c  Actinidia fasciculoides var. orbiculata

Leaves  Thin, chartaceous and obovate-elliptical shape, about 7.0-9.0 cm long, 4.0-5.0 broad; basal part is cuneate or broadly cuneate, usually bilaterally symmetrical; leaf apex is acute, but acuminate on new young leaves. Upper surface of the leaf is green; new leaves on apical parts of shoots ­ purplish-red; glabrous; primary and secondary veins are light green. Leaf margin is almost entire, but toward apex, sparse, short, spine-like bristles appressed, green in color. Lower surface of the leaf is ash-white green, glabrous. Primary vein and secondary veins are green, covered with short white or purple-red pubescence toward base. Secondary veins, six to seven, on each side of the primary vein. Petioles, c. 2.0-2.5 cm long, are green, sparsely covered with short ­purplish-red pubescence.

Distribution: China: Guangxi (endemism) (Figure 2.47).

Diagnostic characters and species relationships A. fasciculoides var. cuneata differs from A. fasciculoides var. fasciculoides by the leaf lamina being obovate to elliptical and the leaf base being cuneate (wedge-shaped or narrowly triangular).

圆叶猕猴桃 yuan ye mi hou tao

Actinidia fasciculoides var. orbiculata C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 313. 1984. TYPE: China. Guangxi: Longzhou, alt. 400 m, October 10, 1957, S.Q. Chen 13557 (Holotype, IBK).

Morphological characteristics Leaves  Relatively small, c. 7 cm long, 4.5 cm broad; leaf margin is slightly serrated, basal part is entire, five secondary veins on each side of the midvein. The petiole is c. 3-8 cm long. Fruit The stalk is about 2 mm long.

Diagnostic characters and species relationships A. fasciculoides var. orbiculata differs from A. ­fasciculoides var. fasciculoides by its usually smaller, rounder leaves with finer serrations and often longer petioles. As a variety of A. fasciculoides, it is, like A. fasciculoides var. fasciculoides, thought to be closely related to A. umbelloides.

Chapter 2 Species

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Figure 2.47  Natural range of Actinidia fasciculoides var. orbiculata.

2.10  Actinidia fortunatii 条叶猕猴桃 tiao ye mi hou tao

Actinidia fortunatii Finet & Gagnepain in Bull. Soc. Bot. France 53: 574. 1906. TYPE: China. Guizhou: Pinyfa, June 8, 1905, Cavalerie & Fortunat 2350 (Holotype, P). Actinidia glaucophylla F. Chun in Sunyatsenia 7: 14. 1948. TYPE: China. Guangdong: Ruyuan, June 9, 1933, S.P. Ko 52835 (Holotype: IBSC). Actinidia longicauda F. Chun in Sunyatsenia 7: 14. 1948. TYPE: China. Guangxi: Quanzhou, June 12, 1937, C.S. Chun 81944 (Holotype, IBSC). Actinidia glaucophylla var. asymmetrica (F. Chun) C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 234. 1984.— Actinidia asymmetrica F. Chun in Sunyatsenia 7: 13. 1948. TYPE: China. Guangxi: Wuming, Damingshan, May 14, 1924, Liu 566 (Holotype, IBSC). Actinidia gracilis C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 316. 1984. TYPE: China. Guangxi: Du’an, October 22, 1957, Y.K. Li P0-1827 (Holotype, IBK). Actinidia glaucophylla var. rotunda C.F. Liang in Guihaia 2(1): 4. 1982. TYPE: China. Guangxi: Daxin, April 11, 1981, H.N. Qin 59 (Holotype, IBK).

Actinidia glaucophylla var. robusta C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 317. 1984. TYPE: China. Guangxi: Longsheng, alt. 980 m, June 7, 1955, Guangfu Exped. 700486 (Holotype, IBK). Distribution: China: Guizhou (endemism) (Figure 2.48).

Morphological characteristics Relatively small, semi-evergreen or deciduous climber. Stems  Lenticels on 1-year-old shoots are small and indistinct. Two-year-old stems are glabrous, c. 1.5-2 mm in diameter; lenticels not visible, pith lamellate, white. Leaves Hard chartaceous. Young leaves often bright pink, mature leaves very variable depending on plant gender and position on plant, firm, almost fleshy. Petioles are 1-2 cm long, glabrous or sparse woolly hairs, shed as leaves age. Lamina is linear or linear-lanceolate, 7-7 cm long and 1.85.0 cm broad, auriculate or obtuse-rounded at base, acuminate to long acuminate at apex; marginal teeth are sparse and inconspicuous, stiff, appressed, ciliate. The upper leaf

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Kiwifruit: The Genus ACTINIDIA

Figure 2.48  Natural range of Actinidia fortunatii.

Figure 2.49  Stem and leaves of Actinidia fortunatii.

surface is green, lustrous and usually glabrous. Midvein is conspicuous, secondary veins are slender, not straight ascending, anastomosing at the margin, veinlets are reticulate. The lower surface is glaucous, with very sparse long downy hairs, usually shed as leaves age, or glabrous (Figure 2.49). Pistillate flowers  Peduncles are very short, covered with reddish-brown downy hairs; pedicles are c. 9 mm long.

Small bracts are diamond-shaped, 2.5 mm long. Calyx has five sepals, glabrous on both surfaces but with ciliate hairs along the margins. Flowers are white to pink, 8 mm in diameter, cup-like, not opening widely, five petals which are obovate, 5.5 cm long. Filaments are 1.5 mm long, anthers yellow, length similar to filaments or shorter. Ovary cylindrical to globose, c. 2.5 mm in greater diameter, and 2 mm in shorter diameter, covered with dense, yellowish-brown, soft hairs.

Chapter 2 Species

Staminate flowers Male plants are very floriferous; generally six flowers per inflorescence. Flowers are white or pink to red, 6 mm in diameter, cup shaped, not opening widely, length of filaments is twice that of anthers. Fruit  Long-cylindrical, dark olive-green, with dense, ­yellowish-brown, elongated or round lenticels; the fruits are flat at apex, with marginate depressed. The stalk is 0.7 cm long, yellowish-brown, covered with yellowish-brown, short downy hairs. Calyx is persistent, reflexed, also styles persistent. Fruit are 1.5 cm long, 0.7 cm in diameter, average weight is 1.0 g. Fruit pulp is green, with yellowish green core. Numerous seeds are small, black-brown. Fruit tastes acidic (Figure 2.50).

81

Diagnostic characters and species relationships The species is characterized by inconspicuous lenticels on 1-year-old stems, lanceolate to round leaves, lower surface glaucous, white to pink to red flowers and thin cylindrical fruits. The species as currently circumscribed is very variable, especially in leaf characters. Morphologically, Actinidia fortunatii appears to be closely related to A. cylindrica and A. henryi (Li, 1952; Liang, 1982b, 1984; Li et al., 2007a,d), and this is supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006) Chromosomes: 2n = 2x = 58, 4x = 116

Figure 2.50  Pistillate flowers and fruit of Actinidia fortunatii.

2.11  Actinidia fulvicoma 黄毛猕猴桃 huang mao mi hou tao

Actinidia fulvicoma Hance in J. Bot. 23: 321. 1885. TYPE: China. Guangdong: Luofushan, May 1883, B.C. Henry Herb. Propr. 22293 (Holotype, BM).

2.11a  Actinidia fulvicoma var. fulvicoma 黄毛猕猴桃(原变种) huang mao mi hou tao (yuan bian zhong)

Actinidia fulvicoma f. arachnoidea C.F. Liang. TYPE: China. Guangxi: Longzhou, alt. 550-870 m, S.Q. Chen 11830 (Holotype, IBK).

Distribution: China: Yunnan, Guangxi, Guangdong, Guizhou, Hunan, Jiangxi, and Fujian (Figure 2.51).

Morphological characteristics Small to medium-sized semi-evergreen climber. Stems  One-year-old shoots are yellow, covered with yellowish to red pubescence toward the tip, and with grayish

yellow pubescence toward the base. Two-year-old stems are black-brown, covered with short grayish yellow pubescence; lenticels are small, elliptical or linear, grayish-white; pith is solid, light green. Leaves  Thick chartaceous, broad ovate, occasionally elliptical. The leaves are c. 7-9 cm long and 4.5-6.3 cm broad. Broad cuneate or occasionally round at base, bilaterally symmetrical, and mucronate to acuminate at apex. The upper leaf surface is green and glossy. Midvein and secondary veins are light green and conspicuous, sparsely covered with pubescence; parallel reticulate veinlets. The leaf margin is nearly entire, with very sparse, small serrations which have tiny or no awns. The lower leaf surface is grayish-white; midvein and secondary veins protrude and are conspicuous, densely covered with yellow pubescence. Petioles are grayish yellow (Figure 2.52). Pistillate flowers  Cyme inflorescences are axillary from base to tip of flowering shoots, 6-10 flowers per inflorescence; peduncle is 3.1 cm long, and the pedicel is 1.7-2.9 cm long and covered with brownish yellow pubescence. Calyx has four to six sepals, predominantly five, mostly nearly lanceolate, occasionally triangular, grayish yellow, covered

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Kiwifruit: The Genus ACTINIDIA

Figure 2.51  Natural range of Actinidia fulvicoma var. fulvicoma.

Figure 2.52  Stem with emerging shoot and leaves of Actinidia fulvicoma var. fulvicoma.

with long white to grayish yellow pubescence. Flowers are white or pink, c. 17 mm diameter, sweetly fragrant. Corolla has 6-10 petals, in 2 concentric circles, petals long obovate, the wavy margins are reflexed, beak-like at base, with stripe in the center, and 8 mm long and 5 mm broad. Stamens, 20-26 per flower, filaments white, anthers are yellow, nearly round, 1 mm long. Styles, 20-26, are light green and 3 mm long, with swollen stigma. Ovary is globose to short elliptical and pale greenish-white, densely covered with white pubescence, and c. 5 mm long and 4 mm in diameter. Staminate flowers  Cyme inflorescence, up to seven flowers; general peduncle is 10 mm long, yellowish-brown, covered with brownish yellow pubescence; pedicles are 9 mm

long. Calyx has four sepals, elliptical, brownish yellow, outer surface densely covered with brownish yellow pubescence. Flowers are 19 mm in diameter and white aging to yellow. Corolla has five petals, obovate, 9 mm long and 4 mm broad. About 50 stamens per flower, filaments are slender, c. 6 mm long, anthers are yellow and oblong, 1 mm long. Rudimentary ovary. The staminate flowers produce sweet scent, similar to that of A. chinensis. Fruit Nearly cylindrical, dark green, average weight c. 3-4 g per fruit. Styles are persistent at apex which is slightly concave. Fruit shoulder is flat and slightly concave. Fruit is covered with downy yellow hairs; irregular yellow fruit spots can be seen after hairs shed. The stalk densely covered with long downy yellow hairs (Figure 2.53).

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Figure 2.53  Flowers (pistillate, staminate) and fruit of Actinidia fulvicoma var. fulvicoma.

Diagnostic characters and species relationships The species is characterized by leaves with brownish dense stellate and yellowish hairs on lower surface, and long tomentum on 1-year-old shoots. Morphologically, Actinidia fulvicoma is related to A. eriantha, A. farinosa, and A. liangguangensis (Li, 1952; Liang, 1982b, 1984; Li et al., 2007a,d). A fulvicoma is distinguished from A. eriantha by the narrower, longer leaves which are pubescent on their upper surfaces, the hairier stems and petioles, the yellowish or brownish indumentum and the relatively hairless fruit (Li, 1952). A close relationship between A. fulvicoma and A. farinosa or A. liangguangensis is supported by molecular genetic data, but a relationship between A. fulvicoma and A. eriantha was not supported (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. fulvicoma var. fulvicoma in the National Germplasm Repository for Actinidia at WBG were 0.1460 and 0.2131, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58

2.11b  Actinidia fulvicoma var. cinerascens 灰毛猕猴桃 hui mao mi hou tao

Actinidia fulvicoma var. cinerascens (C.F. Liang) J.Q. Li & D.D. Soejarto in Novon 16(3): 362. 2006. Actinidia cinerascens C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 320. 1984. TYPE: China. Guangdong: Luofushan, July 27, 1930, N.Q. Chen 41208 (Holotype, IBK). Actinidia cinerascens var. tenuifolia C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 321. 1984. TYPE: China. Guangdong: Jiaoling, alt. 500-600 m, May 12, 1957, L. Deng 4770 (Holotype, IBK). Actinidia cinerascens var. longipetiolata C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 321. 1984. TYPE: China. Hunan: Daoxian, alt. 950 m, June 26, 1959, P.C. Tam 61840 (Holotype, IBK). Distribution: China: Guangdong, Hubei, and Hunan (Figure 2.54).

Morphological characteristics Small to medium-sized, semi-evergreen climber.

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Kiwifruit: The Genus ACTINIDIA

Figure 2.54  Natural range of Actinidia fulvicoma var. cinerascens.

Stems Flowering shoots and young brachlets are densely covered with brown downy pubescence or velvety tomentum; lenticels are not distinct, nearly linear. Two-year-old stems, c. 2.5 mm in diameter, are glabrous or with short tomentum, lenticels are not distinct; pith is lamellate, white.

long. The flowers are c. 1.5 cm in diameter, white. Calyx has five sepals, rectangular elliptical, c. 4 mm long, covered with hairs on the outer surface and glabrous on the inner surface. Petals, five, are obovate, c. 6-7 mm long. Anthers are yellow, ovate to sagittate, c. 1 mm long; ovary is globose, covered with pubescence.

Leaves  Chartaceous, long obovate, obovate-lanceolate or rectangular lanceolate; lamina is 6-10  cm long and 2.5-6  cm broad; obtuse-rounded to subcordate at base and ­mucronate-acuminate at apex; leaf margin has small, ­eyelash-like spines. The upper leaf surface is green and almost glabrous or partly covered with short coarse hairs. The lower surface is light green, densely covered with white stellate hairs. The petiole is 1-5 cm long, densely covered with short tea-brown pubescence.

Fruit Cylindrical to ovoid, c. 15-20 mm long, spotted on the fruit surface and persistent reflexed calyces. Seed are small, c. 1 mm long.

Flowers  Cyme inflorescence, one to seven flowers, densely covered with dark-brown pubescence. Peduncle is about 1.5-7 mm long; bracts are diamond-shaped, c. 3-4.5 mm

As a variety of A. fulvicoma, its specific relationships with other species are probably the same as those of A. ­fulvicoma var. fulvicoma.

Diagnostic characters and species relationships A. fulvicoma var. cinerascens differs from A. fulvicoma var. fulvicoma in the upper surface of the leaves having thin grayish tomentum and young branchlets and petioles having short rusty-brown velvety tomentum.

Chapter 2 Species

2.11c  Actinidia fulvicoma var. hirsuta 糙毛猕猴桃 cao mao mi hou tao

Actinidia fulvicoma var. hirsuta Finet & Gagnepain in Bull. Soc. Bot. France 52 (Mém. 4): 18. 1906. TYPE: China. Guizhou, July 20, 1898, R.P. Bodinier 2427 (Isotype, K).

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Actinidia fulvicoma f. hirsuta (Finet & Gagnepain C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 252. 1984. Distribution: China: Yunnan, Guangxi, Guangdong, Hunan, and Guizhou (Figure 2.55).

Figure 2.55  Natural range of Actinidia fulvicoma var. hirsuta.

Morphological characteristics Small to medium-sized, semi-evergreen climber. Stems  One-year-old shoots are green, with light brown to rusty-colored hispid hairs, but basal parts of shoot have sparse mealy white powder, readily shed; lenticels are indistinct, brown, oval; internodes are c. 1.0-4.0 cm, pith is solid,

Figure 2.56  Stem and leaves of Actinidia fulvicoma var. hirsuta.

green. Two-year-old stems are reddish-brown, sparsely covered with short brown hispid hairs; lenticels are dense, white to yellowish-brown, round or elliptical; internodes are c. 4.5-6 cm, pith is lamellate, brown. Leaves  Chartaceous, predominantly round, occasionally nearly round; base of leaves, subcordate; leaves ­bilaterally

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Kiwifruit: The Genus ACTINIDIA

symmetrical; leaf apex is usually retuse, occasionally mucronate (with a small spike). Leaves are c. 6.0-8.0 cm long and 6.5-9.0 cm broad. Leaf upper surface is green, glabrous; midvein and secondary veins (5-7 each side of midvein) are yellowish green, densely covered with white and pale brown lanate hairs. Leaf margin is densely serrated with small, green, spine-like awns, distinct, some awns brown. The lower leaf surface is whitish-green, sparsely covered with white stellate hairs. Petiole, c. 4-6 cm long, is red on the upper surface and green on the lower surface, covered with short yellow and white lanate hairs (Figure 2.56).

Flowers Calyx has five sepals are usually outward reflexed, oblong, brown, covered with brown pubescence on outer surface and yellowish-brown pubescence on inner surface, c. 8 mm long and 6 mm broad. Fruit Ovoid to slightly tapering cylindrical, relatively small, densely covered with long, yellowish-brown p ­ubescence. Fruit apex is nearly round with persistent styles, sepals are persistent, reflexed. Fruit flesh green; numerous seed. The stalk is yellowish green, covered with ­yellowish-brown pubescence, 1.5-2.0 cm. The fruit are acid, astringent, with slightly numbing, irritating constituents (Figure 2.57).

Figure 2.57  Staminate flowers and fruit of Actinidia fulvicoma var. hirsuta. Fruit photo from Li Jiewei.

Diagnostic characters and species relationships A. fulvicoma var. hirsuta differs from A. fulvicoma var. fulvicoma in its petioles and young branches being densely long hirsute, and the lower leaf surface having brown and white lanate hairs. As a variety of A. fulvicoma, its specific relationships with other species are probably similar to those of A. fulvicoma var. fulvicoma. In addition, A. fulvicoma var. hirsuta appears to be closely related to A. farinosa and A. liangguangensis, as supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The natural PIC of A. fulvicoma var. hirsuta was 0.783 revealed by SSR markers (Liu et  al., 2007), while the average genetic diversity (h) and haplotype genetic diversity (H) based on cpSSR markers was 0.086 and 0.611, respectively (Zhang et al., 2007). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. fulvicoma var. hirsuta in the National Germplasm Repository for Actinidia at WBG were 0.0513 and 0.0749, respectively, as revealed by AFLP markers (Li, 2006).

2.11d  Actinidia fulvicoma var. pachyphylla 厚叶猕猴桃 hou ye mi hou tao

Actinidia fulvicoma var. pachyphylla (Dunn) H.L. Li in J. Arnold. Arbor. 33: 57. 1952. Actinidia pachyphylla Dunn in J. Linn. Soc., Bot. 39: 409. 1911. TYPE: China. Guangdong: Fenghuangshan, Hongkong Herb s.n. Distribution: China: Guangdong (endemism) (Figure 2.58).

Morphological characteristics Leaves  Almost leathery, long ovate or ovate-lanceolate, c. 9-18 cm long and 4.5-5.0 cm broad. Mid and secondary veins are densely covered with coarse hairs, other parts of upper surface glabrous, lower surface has dense tomentum.

Diagnostic characters and species relationships A. fulvicoma var. pachyphylla differs from A. fulvicoma var. fulvicoma in its coriaceous leaves and young branchlets having a dense brownish tomentum.

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Figure 2.58  Natural range of Actinidia fulvicoma var. pachyphylla.

As a variety of A. fulvicoma, its specific relationships with other species are probably the same as A. fulvicoma var. fulvicoma.

2.12  Actinidia glaucocallosa 粉叶猕猴桃 fen ye mi hou tao

Actinidia glaucocallosa C.Y. Wu in Fl. Yunnan. 1: 68. 1977. TYPE: China. Yunnan: Jingdong, alt. 2800 m, May 24, 1963, Q.A. Wu 9320 (Holotype, KUN). Distribution: China: eastern Yunnan (endemism) (Figure 2.59).

Morphological characteristics Large deciduous climber. Stems  Very young shoots may have fine, downy, brown tomentum. One-year-old stems are green, glabrous and smooth; lenticels are inconspicuous, light yellow, elliptical; internodes 1-2.2 cm long. Two-year-old stems are brown, shiny, relatively stout in diameter and glabrous; lenticels are pronouncedly protruded from the surface, relatively sparse but large, yellowish-brown, round or elliptical; pith is relatively large, lamellate, light brown. Leaves  Thick chartaceous, oblong or broad lanceolate, uniform, c. 7.2-11.5 cm long and 2.9-3.2 cm broad; broadly

c­ uneate to near round at base, mostly bilaterally symmetrical; acuminate at apex. Upper leaf surface is dark green, lustrous, glabrous and smooth. Midvein and lateral veins are green without pubescence. Leaf margin slightly undulating, apical part has fine sparse teeth, short, light purple, spreading; the basal part of the leaf margin without teeth. Lower leaf surface is markedly glaucous, glabrous; midvein protruding, glabrous, seven to nine lateral veins on each side of the m ­ idvein, protruding, glabrous, anastomosing at the leaf margin. The petiole is medium stout, pale green to light purple, 1.7-2.5 cm long. Pistillate flowers  Cyme has several lateral axes, two or three flowers per inflorescence; pedicel is c. 4-8 mm long, light purple, sparsely covered with light purple pubescence. peduncle is c. 6 mm long, light purple-red, also sparsely covered with light purple pubescence. The flowers are c. 1.6 cm in diameter. Calyx has five sepals, green, elliptical, c. 6 mm long, 3 mm broad, inner surface is glabrous with sparse longitudinal stripes, outer surface is sparsely covered by light purple pubescence. Corolla has five petals, broadly elliptical, c. 8 mm long, 5 mm broad, wider in basal part with red longitudinal stripes spreading to the middle of the petal; the basal part of the petals pink, the apical part and the margins are light yellow-­white. About 25 stamens per flower, f­ ilaments are white and 2.5-8 mm long, anthers are yellow, sagittate with indistinct longitudinal grooves. About 14 styles per flower, milky-white, c. 4 mm long, with slightly swollen stigma. Ovary is nearly globose, c. 4 mm in diameter, densely covered with light purple pubescence.

Figure 2.59  Natural range of Actinidia glaucocallosa.

Figure 2.60  Stems, leaves, and fruits of Actinidia glaucocallosa.

Chapter 2 Species

Staminate flowers Cyme has several lateral axes, five to eight flowers per inflorescence. Pedicel is c. 8 mm long, slender and light purple green, densely covered with light purple pubescence; peduncle is c. 13 mm long, slender, also sparsely covered with light purple pubescence. The flowers are c. 1.0 cm in diameter. Calyx has five sepals, nearly deltoid in shape, c. 2 mm long and 1 mm broad, brown-green, glabrous in the middle part of the inner surface with sparse red stripes, sparsely covered with light purple pubescence on the margins; densely covered with light purple pubescence on outer surface. Petals, five, are pink, elliptical, c. 5 mm long and 2 mm broad, with radiating red longitudinal stripes spreading to the margin of the petals. About 15 stamens per flower, filaments are white, 3 mm long, anthers are yellow, relatively small, sagittate. Vestigial tiny ovary, densely covered with light purple pubescence Fruit Flat globose; 2.5 cm long, 2-2.8 cm across, averaging 11-16 g; fruit skin is green or reddish-brown, and glabrous; persistent calyx, persistent styles. The stalk is 1.6-2.0 cm long, stalk end is slightly retuse. Fruit spots are ­yellow-brown, round or linear, protruding and relatively dense. Surface of fruit has longitudinal grooves

Figure 2.61  Natural range of Actinidia grandiflora.

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occasionally, also covered with white mealy powder (Figure 2.60).

Diagnostic characters and species relationships The species is characterized by having very conspicuous lenticels on branchlets, leaves which are glaucous underneath but not glandular-tomentose, fruits globose with prominent lenticels. Actinidia glaucocallosa is considered to be closely related to A. trichogyna and A. laevissima (Liang, 1984).

2.13  Actinidia grandiflora 大花猕猴桃 da hua mi hou tao

Actinidia grandiflora C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 323. 1984. TYPE: China. Sichuan: Tianquan, alt. 1800 m, June 7, 1953, X.L Jiang 34327 (Holotype, IBK). Distribution: China: Sichuan, Yunnan, and Guizhou (Figure 2.61).

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Kiwifruit: The Genus ACTINIDIA

Morphological characteristics Large deciduous climber. Stems  One-year-old shoots are yellow-brown, covered with yellowish-brown tomentum; lenticels are inconspicuous, light yellow, round to elliptical; internodes are 1.5-4.0 cm long. Two-year-old stems are brown with very sparse, coarse brown hairs; lenticels are light yellow, round or elliptical; pith is lamellate, light yellow (Figure 2.62). Leaves Chartaceous, ovate-oblong to elliptical; leaf lamina is 9-14 cm long and 6.5-10.5 cm broad; bilaterally symmetrical; leaf base; auriculate-cordate, shallowly retuse; leaf apex acute. The upper leaf surface is dark green and g­ labrous. Leaf margin is serrate, the teeth appressed or spreading, green; midvein and secondary veins are inconspicuous, yellowish green color and covered with brown ­tomentum. The lower leaf surface is light green, covered with white tomentum and imperfect stellate hairs; ­conspicuous midvein and secondary veins are whitish-green, densely covered with white tomentum. The petiole, c. 4-7 cm long, is purplish-red, sparsely covered with fine yellow-brown tomentum.

­ etals, 10-13 mm long, obovate, white maturing to dirty yelp low. The filaments are slender, c. 4-5 mm long; anthers are yellow, sagittate. Rudimentary ovary is densely tomentose, 3 mm in diameter. Fruit  Fruit are c. 4.9 cm long and 3.9 cm in diameter, averaging 40 g; ellipsoid or cylindrical; fruit skin is brown and covered with fine, grayish-white or yellowishbrown hairs, persistent. The fruit apex is rounded, with beak, persistent styles. Fruit shoulder is narrower, round with persistent calyx. The stalk is brown and glabrous, c. 3 cm long. Green fruit flesh with large, elliptical white core. Seed, purplish-red, elliptical. Fruit ripen in September, acid

Pistillate flowers  Cyme or solitary inflorescence with two to three flowers. Pedicel is about 0.8 cm long, green, covered with white or light brown tomentum. Calyx has five to seven, predominantly five, sepals, c. 10 mm long and 8 mm broad, elliptical or ovate, densely covered with brown tomentum. Flowers large, c. 4.5 cm in diameter, greenish-white, but turning to yellow later; five to six petals, predominantly five, nearly round c. 20 mm long, radiating stripes on the surface. About 135 stamens per flower. Filaments are white to light green, 8 mm long. Anthers are yellow, sagittate. Styles are white, usually 28 per flower, 6 mm long with slightly swollen stigma. Ovary is short cylinder, 6 mm in diameter, covered with dense, white or light brown tomentum. Staminate flowers Cyme inflorescence, two to three flowers. Calyx of six sepals, c. 5-6.5 mm long, ovate to long obovate, short tomentum on outer surface, but inner surface glabrous. Opening flowers are c. 2.0 cm in diameter, six

Figure 2.62  Stem of Actinidia grandiflora.

Figure 2.63  Pistillate flowers and fruit of Actinidia grandiflora.

Chapter 2 Species

flavor but not astringent, juicy and fine texture. Vc content is 50-200 mg per 100 g fresh weight. Soluble solids content is 13.5% and total acidity is 1.05% (Figure 2.63).

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2.14  Actinidia hemsleyana 长叶猕猴桃 chang ye mi hou tao

Morphologically, Actinidia grandiflora appears to be closely related to A. chinensis var. deliciosa and A. obovata (Liang, 1984), and this is supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006).

Actinidia hemsleyana Dunn in J. Linn. Soc., Bot. 38: 355. 1908. TYPE: China. Fujian: Nanping, Hongkong Herb. 2400 (Holotype, HK). Actinidia hemsleyana var. kengiana (F.P. Metcalf ) C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 249. 1984.— A. kengiana F.P. Metcalf in Lingn. Sci. Journ. 11: 16. 1932. TYPE: China. Zhejiang: Jingning, forest understory, August 16, 1926, Y L. Keng 394 (Holotype, A; Isotype, PE). Actinidia subglaucifolia F.P. Metcalf in Lingn. Sci. Journ. 11: 15. 1932. TYPE: China. Fujian: Shouning, August 12, 1926, Y.L. Keng 339 (Holotype, A; Isotype, PE).

Chromosomes: 2n = 2x = 58, 4x = 116

Distribution: China: Zhejiang, Jiangxi, and Fujian (Figure 2.64).

Diagnostic characters and species relationships The species is characterized by having obovate leaves with very sparse, imperfect stellate hairs on the lower surface, and flowers with six petals.

Figure 2.64  Natural range of Actinidia hemsleyana.

Morphological characteristics Vigorous deciduous climber. Stems  Shoots strongly dimorphic. Some 1-year-old shoots grow vigorously, up to 3 m, long and stout, red, densely ­covered with rusty red, bristle-like hairs that are 4 mm long,

bearing narrow leaves; others are much shorter, dark-brown or yellowish-brown, covered with yellowish-brown bristles. Two-year-old stems are brown, dark-brown or grayish brown, densely pubescent with long brown to black ­bristle-like hairs, but becoming glabrous with age; pith is light green, solid becoming brown, lamellate in older shoots.

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Kiwifruit: The Genus ACTINIDIA

Leaves  Chartaceous, lanceolate, obovate-lanceolate or oblong, very variable in size and shape, 7-19.2 cm long, 3-7 cm broad, wider and broader than those of A. lanceolata. Leaves are usually wider above the midpoint, nearly round or cuneate at base. Young leaves can be reddish-brown (Figure 2.65). The upper leaf surface of older leaves is dark green, slightly glossy,

g­ labrous. There are marginal teeth with very short awns mainly on the apical part of the leaf. The lower leaf surface is glaucous, grayish-green, sometimes sparsely covered with grayish-white pubescence. Light green veins protrude conspicuously from the lower surface. Leaf stalks are long, pale red, sparsely covered with dark-brown pubescence, later becoming glabrous.

Figure 2.65  Stem and leaves of Actinidia hemsleyana.

Pistillate flowers Axillary cyme inflorescences with two to three flowers, sometimes single. Calyx has five sepals, brown, pubescent on outer surface. Flowers are white to light purple, but darker purple at petal bases, becoming paler as flowers age. Numerous stamens with purplish black anthers. Ovary is globose, c. 6 cm in diameter, with dense brown tomentum. Staminate flowers Cyme inflorescence, one to three flowers. Calyx has five sepals, 0.8-1.2 cm long, broad ovate, apex short acuminate, outer surface has dense, short brown tomentum. Corolla has five petals, ovate, 1-1.5 cm long, white to pale pinkish. Numerous stamens, anthers 0.7-1 cm long, purple. Rudimentary ovary tomentose. Fruit Cylindrical, short globose or irregular short cylindrical. Fruit are 1.8-3.2 cm long and 1.5-2.6 cm in diameter. Fruit apex sometimes shallowly concave. Skin has inconspicuous lenticels, densely covered with grayish, ­yellow-brown bristle-like hairs, becoming glabrescent when mature. Persistent reflexed sepals and remnant styles. Fruit stalk s dark-brown, pubescent, 1.1-1.5 cm long, medium diameter. Fruit flesh is yellowish green to green. The fruit are acid sweet, juicy, and have a strong aroma. Seed are reddish-brown, smaller than those of A. chinensis, oblong, several hundred seed per fruit, about 0.8 g per 1000 seed. Vc content is 12.8 mg per 100 g fresh weight, soluble solids content is 6.9% and total acidity is 1.05% (Figure 2.66).

Diagnostic characters and species relationships The species is distinguished by its oblanceolate leaves which are glaucous underneath and the floral branchlets which have sparse rusty-brown strigose hairs.

Morphologically, Actinidia hemsleyana was considered by Li (1952) to be more related to A. melliana, A. henryi, and A. arisanensis than other species. Subsequently, A. arisanensis was merged into A. callosa by Li et al. (2007a,d). However, these relationships were not supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. hemsleyana in the National Germplasm Repository for Actinidia at WBG were 0.0079 and 0.0115, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58

2.15  Actinidia henryi 蒙自猕猴桃 meng zi mi hou tao

Actinidia henryi Dunn in Bull. Misc. Inform. Kew 1906: 1. 1906. TYPE: China. Yunnan: Mengzi, A. Henry 10381 (Holotype, K; Isotype, US). Actinidia henryi var. polyodonta Handel-Mazzetti in Symb. Sin. 7: 391. 1933. TYPE: China. Yunnan: Fumin, May 18, 1915, Sandstein 6113. Actinidia carnosifolia C.Y. Wu in Fl. Yunnan. 1: 60. 1977. TYPE: China. Yunnan: Malipo, Laojunshan, May 25, 1962, K.M. Feng 22768 (Holotype, KUN). Actinidia carnosifolia var. glaucescens C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 318. 1984. TYPE: China. Guangxi: Longsheng, alt. 1140 m, July 24, 1955, Guangfu Exped. 996 (Holotype, IBK). Distribution: China: Yunnan, Guangxi, Guangdong, Guizhou, and Hunan (Figure 2.67).

Chapter 2 Species

Figure 2.66  Flowers (pistillate, staminate) and fruit of Actinidia hemsleyana.

Figure 2.67  Natural range of Actinidia henryi.

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Morphological characteristics Medium to large, semi-deciduous climber. Stems  One-year-old shoots are c. 3.3 cm long, light green, densely covered with coarse brownish purple hairs. Twoyear-old stems are light brown, densely covered with long, coarse brown hairs; lenticels are usually sparse, oblong, light brownish yellow; pith is lamellate, white. Leaves Chartaceous to leathery, 7-14 cm long and 3-6.5 cm broad, oblong, round to shallowly cordate at base; bilaterally symmetrical; acuminate apex. The upper leaf surface is green, generally glabrous, slightly rough. The midvein is indistinct, brown to black, with sparse, short, downy brown to black hairs, or glabrous, slightly concave. Many small serrations on the leaf margin are spreading, awns are darkbrown. The lower surface is pale green, usually light silvery green between secondary veins, glabrous except for veins, 9-10 secondary veins on each side of prominent midvein. Petioles are relatively short, 2.5-3.0 cm long, covered with short, coarse, rusty-brown hairs. Pistillate flowers  Cyme inflorescence with two to three flowers. Pedicels are c. 7 mm long, light green, covered with dense light brown and white shaggy hairs. Calyx has five ovate to elliptical sepals, 7 mm long, 6 mm broad, white, covered with dense rusty-brown shaggy hairs on the outer surface, parallel strips on inner surface spreading toward the margins. Corolla has five petals, obovate, 7 mm

long and 6 mm broad, white to pink. About 30 stamens, white filaments, 3 mm long, anthers are yellow, sagittate. Styles are white with slightly swollen stigma; ovary is yellowish green, with dense, white pubescence, globose to short cylindrical, c. 3.5 cm in long diameter, 2.2 cm in short diameter. Staminate flowers Cyme inflorescence, usually with three to four flowers. Pedicels are slightly slender, 9-12 mm long, yellowish green, densely covered with yellowish-brown pubescence. Calyx has four to five sepals, mostly five, 5 mm long, 3 mm broad, milky-white, densely covered on outer surface with light yellow pubescence. Corolla has five to six petals, predominantly five, 7 mm long, 5 mm broad, white to pink, obtuse at base, ruffled edges. About 40 stamens, filaments are milky-white, 2.5 mm long; anthers are golden-yellow, sagittate, 1.8 mm long. Ovary is rudimentary, tiny, conical, covered with yellowish-brown pubescence. Fruit Cylindrical to long conoid, narrowing to apex, wider at shoulders. Skin is glabrous, green, numerous large yellowish-brown lenticels, round, elliptical or irregular ­ shaped, conspicuous. Sepals and stylar remains persistent. Stalk, 9 mm long, brown, covered with downy, brown hair. Fruit are 16 mm long and 9 mm in diameter, average weight 4 g. Fruit flesh is green with pale green core. Numerous seed are yellow, smooth and ovoid. The fruit have a light acid flavor without astringency. Vc content is 4.40 mg per 100 g fresh weight, soluble solid content is 5.6% and titratable acidity is 0.77% (Figure 2.68).

Figure 2.68  Leaves and fruit of Actinidia henryi.

Diagnostic characters and species relationships

2.16  Actinidia holotricha

The species is distinctive because the floral branchlets have dense brown strigose hairs, the coriaceous leaves are glaucous underneath, and the fruit are often cylindrical.

Actinidia holotricha Finet & Gagnepain in Bull. Soc. Bot. France 52 (Mém. 4): 18. 1906. TYPE: China. Yunnan: Outchay, May 18, 1882, Delavay s.n. (Holotype, P).

Actinidia henryi appears to be closely related to A. hemsleyana and A. rudis (Li, 1952).

Distribution: China: Yunnan (endemism) (Figure 2.69).

全毛猕猴桃 quan mao mi hou tao

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95

Figure 2.69  Natural range of Actinidia holotricha.

Morphological characteristics Large, deciduous climber. Stems  One-year-old shoots are green, covered with light brown hispid hairs; lenticels are not distinct, white and elliptical; internodes are c. 1-3 cm long. Twoyear-old stems are dark purple-brown, covered with darkbrown hispid hairs; lenticels are dense, grayish-white, linear or elliptical; internodes are c. 10 cm long; pith is lamellate, white. Leaves  Chartaceous and nearly round or ovate, 8.513.5 cm long and 6.5-13.5 cm broad mostly bilaterally symmetrical at base of leaf, short acuminate at apex. Upper surface is green with strigose hairs; midvein and lateral veins are pale green, sparsely covered with white pubescence, denser at the base of the midvein. Lower leaf surface is pale green, covered with sparse strigose hairs; midvein and lateral veins are greenish-white. Leaf margin has, large teeth alternating with small teeth, tips green, spreading, more frequent toward leaf apex. Petiole,

3.5-6.5 cm long, is purple-red, densely covered with light brown to red hispid hairs. Pistillate flowers  Usually single; pedicel, c. 2.5 cm long, is green, covered with white and light brown pubescence. Calyx of five to six sepals, mostly six or occasionally seven, c. 7 mm long and 7 mm broad, elliptical or ovate, light brown, outer surface covered with light brown pubescence. Flowers are c. 5 cm in diameter, light yellow, six petals, occasionally seven, c. 2.5 cm long and 1.8 cm broad, obovate with longitudinal strips. About 150 stamens, filaments are white, slender, 9 mm long, anthers are yellow and ellipsoid, c. 9 mm long. Numerous styles, stigma slightly swollen, ovary is short cylindrical to globose, c. 6 mm in diameter, densely covered with white hairs.

Diagnostic characters and species relationships The species is characterized by suborbicular leaves with cordate base, strigose hairs on both leaf surfaces. Actinidia holotricha appears to be closely related to A. rudis and A. rubus (Liang, 1984).

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2.17  Actinidia hubeiensis 湖北猕猴桃 hu bei mi hou tao

Actinidia hubeiensis H.M. Sun & R.H. Huang in J. Wuhan Bot. Res. 12(4): 321. 1994. TYPE: China. Hubei: Wuhan, April 27, 1988, H.M. Sun 8805 (Holotype, HIB). Distribution: China: Hubei (endemism) (Figure 2.70).

Morphological characteristics Vigorous, deciduous climber. Stems One-year-old shoots are yellowish green, the apical parts are purplish-red, glabrous; lenticels are light yellow;

Figure 2.70  Natural range of Actinidia hubeiensis.

Figure 2.71  Stem and leaves of Actinidia hubeiensis.

buds are densely covered with dark rusty-brown tomentum. Two-year-old stems are brown, glabrous; lenticels are light yellow, dot-like or linear; pith is lamellate, light brown. Leaves Chartaceous; ovate, long ovate or obovate, leaf blade is 6-14 cm long and 5-13 cm broad; bilaterally symmetrical; rounded or mucronate at apex, rarely emarginated, cordate at base; leaf margins have fine hairs, like eyelashes, on serrations. The upper leaf surface is dark green, the lower surface is light green, sparsely covered with white stellate hairs, readily shed; veins are conspicuous, six to nine secondary on each side of midvein, purplish-red when young. The petiole is light purplish-red, densely covered when young with downy hairs, gradually shed (Figure 2.71).

Chapter 2 Species

Pistillate flowers Predominantly single. Pedicel is c. 0.7-1.5 cm long, covered with short downy hairs. Calyx has three to six sepals, ovate, c. 3-5 mm long and 2-4 mm broad, with dense brown tomentum. Flowers are c. 2 cm in diameter; five to six petals which are obovate and c. 7-8 mm long, 5-8 mm broad, white at apex, and pale pink-purple at base, aging to pale yellow. Numerous stamens, filaments white, 4-5 mm long, anthers are yellow, kidney-shaped, 1.2-1.6 mm long. Styles are string-like, milky-white, usually 30-36 per flower, 4-5 mm long with slightly swollen stigma. Ovary is cone-shaped, 4.5 mm in diameter, densely covered with gray-white pubescence.

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Fruit  Ovate, conical or nearly spherical; c. 2-3 cm long and 1.2-2.5 cm in diameter; averaging 7 g. Fruit skin is dark green; densely spotted, lenticels yellowish-brown; hairs shed so nearly glabrous at maturity. Fruit shoulder is round with persistent calyx reflexed. Fruit stalk is c. 2.0-2.5 cm long. Flesh green with small, whitish core. Red-brown seed. The fruit ripen in October, sweetly acid flavor, juicy and fine texture. VC content is 51-60 mg per 100 g fresh weight; soluble solids content is 14% and total acidity is 2.24% (Figure 2.72).

Figure 2.72  Pistillate flowers and fruit of Actinidia hubeiensis.

Diagnostic characters and species relationships The species is characterized by its leaves being narrowly ovate to broadly orbicular obovate with apex abruptly acute to rounded, and very sparse tomentum of stellate hairs on the lower leaf surface, fruits ovoid to conical to subglobose with dense spots, glabrous. Actinidia hubeiensis is closely related to A. chinensis and may have originated by natural hybridization between A. chinensis and another species, such as A. callosa (Sun and Huang, 1994): this is supported by molecular

genetic data (Huang et  al., 2002a; Li et  al., 2002a; Chat et al., 2004; Li, 2006). Chromosomes: 2n = 2x = 58

2.18  Actinidia indochinensis 中越猕猴桃 zhong yue mi hou tao

Actinidia indochinensis Merrill in J. Arnold. Arbor. 19: 53. 1938. Actinidia callosa Lindley var. indochinensis (Merrill) H.L. Li in J. Arnold. Arbor. 33: 48. 1952. TYPE: Vietnam. Chapa, A. Petelot 5938 (Holotype, A).

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Kiwifruit: The Genus ACTINIDIA

2.18a  Actinidia indochinensis var. indochinensis 中越猕猴桃(原变种) zhong yue mi hou tao (yuan bian zhong)

Actinidia glabra H.L. Li in J. Arnold. Arbor. 33: 44. 1952. TYPE: China. Guangxi: Nanning, in shrubs, alt. 390 m, October 15, 1928, R C. Ching 7875 (Isotype, PE). Actinidia sabiifolia auct. non Dunn: Chun in Sunyatsenia 4(3-4): 190. 1940, excl. S.P. Ko 53651.

Actinidia flavofloris H.Z. Jiang in J. Southwest Agric. Univ. 17(2): 94. 1995. TYPE: China. Yunnan: Hekou, May 25, 1989, S.S. Sun et al. 89-3002 (Holotype, Herbarium, Institute of Horticulture, Yunnan Academy of Agricultural Sciences). Distribution: China: Yunnan, Guangxi, Guangdong, Fujian, and probably extending to Vietnam (Figure 2.73).

Figure 2.73  Natural range of Actinidia indochinensis.

Morphological characteristics Large vigorous deciduous climber. Stems  One-year-old shoots are yellow-brown, apical parts with grayish yellow, downy hairs; pith is solid and light green. Two-year-old stems are reddish-brown, glabrous, lenticels are white, linear, sparse; pith is lamellate and light brown.

Figure 2.74  Stem, emerging shoots, and leaves of Actinidia indochinensis.

Leaves Chartaceous when young becoming leathery when old, oblong or long round, lanceolate, 4-10 cm long and 3.5-5 cm broad. Cuneate or broadly cuneate at base, acuminate or acute at apex, long tip obliquely. Leaf upper surface is green and glabrous. New leaves are relatively smooth. Veins are brown and glabrous. Leaf margin is entire in lower part, sparse serrations in apical part of undulating margin, serrulate tips are brown.

Chapter 2 Species

The lower surface is glaucous, and glabrous, midvein and lateral veins are light brown; the basal parts of veins on new leaves are covered with glandular tomentum seldom seen on older leaves; six to seven secondary veins on each side of midvein; anastomosing veins are dense and indistinct The petiole is slender, brown, c. 2.5-4.0 cm long, in new leaves covered with fine rusty-brown pubescence, becoming glabrous (Figure 2.74). Pistillate flowers  Usually single, 1.5-1.8 cm in diameter; pedicel is 2-3 cm long; calyx has five to six sepals, ovate, 3-5 mm, covered with fine rusty-brown tomentum; corolla has five to six petals, usually white, obovate, c. 1 cm long; ovary, 3-4 mm in diameter, round, densely covered with rusty-brown tomentum; styles 7-8 mm long.

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Staminate flowers Cyme inflorescence, 1-11 flowers; pedicel is c. 7 mm long, yellow-brown, peduncle is c. 2.1 cm long, relatively slender and yellow-brown. Calyx has five sepals, oblong, c. 6 mm long and 2 mm broad, light green, covered with fine tomentum. Numerous stamens, c. 35, anthers yellow, ovoid. Fruit Short ellipsoid; c. 4-5 cm in diameter, averaging 6.08.0 g, skin is yellowish green but becomes tan brown when mature, russet-like, dense lenticels distinct, ­yellowish-brown, Fruit flesh green, medium core, numerous seed. The fruit has sweet-acid flavor, juicy. Vc content is 16.5-41.5 mg per 100 g fresh weight, soluble solids content is 7.0-9.4%, sugar content 5.7-6.4%, and acid content 2.0% (Figure 2.75).

Figure 2.75  Flower and fruit of Actinidia indochinensis.

Diagnostic characters and species relationships

Chromosomes: 2n = 2x = 58, 4x = 116

The species is distinct in that its leaves are glaucous underneath and basal obcuneate, glandular tomentum on leaf midveins and fruits globose to nearly globose.

2.18b  Actinidia indochinensis var. ovatifolia

A. indochinensis was considered by Liang (1984) on morphological grounds to be a sister species of A. chrysantha, and this is supported by molecular genetic data on paternal lineages (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006)

Actinidia indochinensis var. ovatifolia R.G. Li, X.G. Wang & L. Mo in Guihaia 22: 386. 2002. TYPE: China. Guangxi: Guilin, April 4, 1993, R.G. Li 9302 (Holotype, IBK).

卵圆叶猕猴桃(变种) luan yuan ye mi hou tao

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Kiwifruit: The Genus ACTINIDIA

Figure 2.76  Natural range of Actinidia indochinensis var. ovatifolia.

Distribution: China: Guangxi (endemism) (Figure 2.76).

Diagnostic characters and species relationships

Morphological characteristics

A. indochinensis var. ovatifolia differs from A. indochinensis var. indochinensis in its leaf blade being broadly ovate to orbicular and the upper leaf surface having sparse, very fine, downy hairs.

Leaves Ovate to orbicular; the upper surface is covered with very fine downy hairs; hairs on the lower surface gradually shed later. Flowers White. Fruit The apex of the fruit is covered by short yellowish tomentum, persistent stylar remains (Figure 2.77).

As a variety of A. indochinensis, its relationships with other species are probably the same as those of A. indochinensis var. indochinensis.

2.19  Actinidia kolomikta 狗枣猕猴桃 gou zao mi hou tao

Figure 2.77  Fruit of Actinidia indochinensis var. ovatifolia. Photograph by Li Jiewei.

Actinidia kolomikta (Maximowicz & Ruprecht) Maximowicz in Mém. Acad. Imp. Sci. St.-Pétersbourg Divers Savans 9: 63. 1859.—Prunus? kolomikta Maximowicz & Ruprecht in Bull. Cl. Phys.-Math. Acad. Imp. Sci. SaintPétersbourg 15: 129. 1856. TYPE: Amur River, C.J. Maximowicz s.n. (Isotype, NY). Actinidia kolomikta var. gagnepainii (Nakai) H.L. Li in J. Arnold Arbor. 33: 19. 1952. Actinidiagagnepainii Nakai in Bot. Mag. Tokyo 47: 258. 1933. TYPE: China. Hubei, A. Henry 8806 (Syntype, GH) Actinidia tetramera Maximowicz var. maloides (H.L. Li) C.Y. Wu in Fl. Yunnan. 1: 69. 1977. Actinidia maloides H.L. Li in J. Arnold Arbor. 33: 25. 1952. TYPE: China. Sichuan: Yuexi, T.T. Yu 927 (Isotype, PE). Actinidia leptophylla C.Y. Wu in Fl. Yunnan. 1: 63. 1977; auct. non C.F. Liang in Fl. Reipubl. Popularis Sin.

Chapter 2 Species

49(2): 229. 1984. TYPE: China. Yunnan: Yongshan, alt. 2300 m, June 21, 1932, H.T. Tsai 51071 (Holotype, KUN; Isotype, PE). Actinidia maloides H.L. Li f. cordata C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 311. 1984. TYPE: China. Sichuan: Leibo, alt. 2100 m, July 1, 1959, Sichuan Eco. Pl. Exped. 1035 (Holotype, PE). Distribution: China: widely occurs in Yunnan, Hunan, Jiangxi, Sichuan, Chongqing, Hubei, Gansu, Shaanxi, Henan, Shanxi, Hebei, Beijing, Liaoning, Jilin, Heilongjiang. Japan, Korea and Far East region of Russia (Figure 2.78).

Morphological characteristics Large deciduous climber.

Figure 2.78  Natural range of Actinidia kolomikta.

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Stems One-year-old shoots are grayish brown and glabrous, or sparsely covered with white or light brown downy hairs, lenticels not obvious. Two-year-old stems are y­ ellowish-brown or dark-brown, glabrous; numerous lenticels are distinct, round to linear, yellowish white; pith is lamellate, yellowish-brown. Leaves Membranous to thin papery, ovate to elliptical, 6-15 cm long and 5-10 cm broad. Leaf base is cordate, occasionally nearly round; apex is acuminate or acute. Leaf upper surface is yellowish green, glabrous; leaf margin is serrated into simple teeth to sharp spine-like teeth. The lower surface is grayish-green, sparsely covered with short grayish brown pubescence, often in axils of veins. Leaves can often be variegated with white or pink blotch toward the apex, fading as leaves age. The petiole is 2.5-5 cm long, pale yellow to reddish-brown, initially with sparse dusty pubescence, later shed (Figure 2.79).

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Kiwifruit: The Genus ACTINIDIA

Figure 2.79  Stem and leaves of Actinidia kolomikta.

Pistillate flowers  Predominantly single, sometimes in a few inflorescences with two to three flowers; 10-15 mm in diameter. Pedicel is c. 5-6 mm long, green, usually glabrous. Calyx has four to five sepals, nearly round, green with red blush. The corolla has five to seven petals, 6-10 mm long, white, sometimes greenish toward base, obovate, sometimes with fine sparse pubescence but usually glabrous. Stamens, c. 14-16 per

flower, filaments 2-3 mm long, anthers almost round or oblong, yellow. Styles are white, usually 16-18 per flower, 3-6 mm long. Ovary is spherical, light yellowish green and glabrous. The pistillate flowers produce a sweet fragrance. Staminate flowers  Single or as cyme inflorescences with three flowers. The size, shape, and color of staminate flowers are similar to those of pistillate flowers. Usually 16-18 stamens per flower, yellow anthers. Vestigial ovary. Fruit Small, c. 3 cm long and 1.5 cm in diameter averaging 1.9-5.0 g, long cylindrical or nearly globose, fruit apex sometimes with a beak. Calyx and stylar remains often persistent. Fruit skin is green with vertical stripes on the surface, becoming yellowish green on maturity, glabrous; lenticels usually absent, if present are very inconspicuous, nearly round. The fruit stalk is about 2.5 cm long. Fruit flesh is dark green, fine texture with small core, juicy, very sweet acid flavor with fruity aroma. VC content is 100-200 mg per 100 g fresh weight. Soluble solids content is 11% and total acidity 1.3%. In Wuhan, the fruit ripen in November (Figure 2.80).

Figure 2.80  Pistillate flowers and fruit of Actinidia kolomikta.

Chapter 2 Species

Diagnostic characters and species relationships

Morphological characteristics

The species is characterized by its broadly ovate leaves, unequal at the base and the spotless fruit.

Medium vigorous deciduous climber.

Actinidia kolomikta is a very distinct species and morphologically does not appear to be closely related to any other species (Liang, 1984; Li, 1952; Li et al., 2007a,d), a conclusion supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). Chromosomes: 2n = 2x = 58, 4x = 116

2.20  Actinidia laevissima 滑叶猕猴桃 hua ye mi hou tao

Actinidia laevissima C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 317. 1984. T YPE: China. Guizhou: Jiangkou, alt. 1700 m, June 3, 1964, Z.S. Zhang et al. 402236 (Holotype, PE). Actinidia jiangkouensis S.D. Shi & Z.C. Zhang in Acta Bot. Yunnan. 16(4): 345. 1994. TYPE: China. Guizhou: Jiangkou, alt. 530 m, October 20, 1992, Q.B. Wang et al. 92025 (Holotype, GBG; Isotype, KUN). Actinidia laevissima var. floscula S.D. Shi in Acta Bot. Yunnan. 16(4): 346. 1994. TYPE: China. Guizhou: Yinjiang, alt. 1040 m, S.D. Shi 92013 (Holotype, GBG; Isotype, KUN). Distribution: China: Yunnan, Guizhou, Hunan, and Hubei (Figure 2.81).

Figure 2.81  Natural range of Actinidia laevissima.

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Stems  One-year-old shoots are brownish-green; glabrous; lenticels are distinct, yellow. Two-year-old stems are ­reddish-brown; lenticels are relatively large and distinct, linear or oblong; pith is lamellate, yellow-brown. Leaves Chartaceous; oblong, leaf blade is c. 8.4-9.0 cm long and 3.6-3.9 cm broad; basal predominantly cuneate; apex acuminate, slightly oblique. The upper surface is green. Almost entire margin with tiny white serrulates and brown ciliate. Secondary veins are indistinct. The lower surface is white-green; main and secondary veins are distinct, green and anastomosing. The petiole is green or purple green, slightly covered with ­yellow-brown pubescence and about 1.7-2.0 cm long. Flowers  Predominantly single, 8-12 mm in diameter, pink or white; corolla has c. five to seven petals. Four sepals at calyx are 4-5 mm long, oblong, pubescent abaxially near margin of inner surface, over whole outer surface. Filaments are 2.5-3.5 mm long. Anthers are yellow color, ovoid. Ovary is subglobose, 3 mm in diameter, covered with thinly grayish pubescent. Pedicel is about 10-15 mm long, glabrous. Fruit  Small, almost spherical to cylindrical, dark olivegreen, 15 mm long, 8 mm in diameter, glabrous, numerous ­yellow-brown lenticels, prominent stylar remains, some sepals persistent. Seed 1-1.5 mm (Figure 2.82).

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Kiwifruit: The Genus ACTINIDIA

Figure 2.82  Stem, leaves, and fruit of Actinidia laevissima.

Diagnostic characters and species relationships The species is characterized by its lenticels conspicuously raised on branchlets, leaves glaucous abaxially and spotted glabrous fruits. Actinidia laevissima was considered by Liang (1984) to be closely related to A. glaucocallosa.

Figure 2.83  Natural range of Actinidia lanceolata.

2.21  Actinidia lanceolata 小叶猕猴桃 xiao ye mi hou tao

Actinidia lanceolata Dunn in J. Linn. Soc., Bot. 38: 356. 1908. TYPE: China. Fujian: Nanping, April-June 1905, Hongkong Herb. 2399 (Holotype, HK). Distribution: China: Guangdong, Guizhou, Hunan, Jiangxi, Fujian, Zhejiang, and Anhui (Figure 2.83).

Chapter 2 Species

Morphological characteristics Relatively slight, deciduous climber. Stems  Young shoots are densely covered with short, reddish-brown or dark-brown tomentum. One-year­ old shoots are light to dark-brown, densely covered with coarse brown hairs or long dark-brown hairs; pith is small, ­yellowish-brown, lamellate. Older stems are gray or darkbrown, glabrous; lenticels are sparse, inconspicuous, grayish brown, oblong, elliptical- or diamond-shaped. Leaves  New leaves and their petioles often pinkish, covered with brownish hairs. Older leaves are thick and chartaceous, lanceolate, oblanceolate, ovate-lanceolate or round cuneate, c. 4-15 cm long and 1.5-5 cm broad; short acuminate at apex and cuneate or round cuneate at base. The apical part of the leaf margin has small teeth ending in short, thin spines. The upper leaf surface is dark green, often red on margins, lustrous, glabrous; distinct veins are pale green; the lower leaf surface is glaucous, grayish-­green, densely covered with g­ rayish-white or grayish yellow-­brown, stellate hairs. The petiole is 0.9-3.1 cm long, purplish-red, sometimes densely c­ overed with short yellowish-­brown hairs (Figure 2.84).

small, downy, reddish-brown hairs, the inner surface sparse fine hairs. Greenish-white to white corolla has five petals, oblong to spatulate-obovate, 4-5.5 mm in length. Numerous stamens; filaments 1-4 mm long; anthers ­yellow, oblong, c. 1-1.5 mm long. Styles are string-like, c. 1.5 mm long. Ovary nearly globose to ovoid, c. 1.5 mm in diameter, densely covered by dense, reddish-brown tomentum. Fruit Fruit are very small, averaging c. 1 g; mostly long, cylindrical, globose or ovoid, 8-10 mm in diameter. Fruit skin is light brown and covered with fine, yellowish-brown hairs,

Flowers  Inflorescences cymose, two-branched, seven-­ flowered; peduncles 3-6 mm; pedicels 2-4 mm; bracts linear, 1-1.5 mm. Calyx has three to five sepals, ovate to oblong, 3-4 mm in length; the outer sepal surface has

Figure 2.84  Stem, over-wintering bud, and leaves of Actinidia lanceolata.

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Figure 2.85  Pistillate flowers and fruit of Actinidia lanceolata.

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Kiwifruit: The Genus ACTINIDIA

readily shed on maturity; lenticels (fruit spots) are distinct, round, grayish-white; reflexed sepals are persistent, fruit shoulder is slightly retuse. The fruit flesh is dark green, the small round core is pale green. Small number of seed, brown or dark-brown, long, ovate. Fruit ripen in November, acid flavor, juicy. VC content is 33 mg per 100 g fresh weight, soluble solids content is 12% and total acidity 1.2% (Figure 2.85).

Diagnostic characters and species relationships The species is distinct in its leaves elliptic-ovate to e­lliptic-lanceolate, leaf abaxially dense very short, grayish stellate hairs. Actinidia lanceolata was considered by Liang (1984) to be related to A. styracifolia and A. latifolia, but this is not supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. lanceolata resources in the National Germplasm Repository for Actinidia at WBG were 0.0513 and 0.0749, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58

2.22  Actinidia latifolia 阔叶猕猴桃 kuo ye mi hou tao

Actinidia latifolia (Gardner & Champion) Merrill in J. Straits Branch Roy. Asiat. Soc. 86: 330. 1922. Heptaca? latifolia Gardner & Champion in Hooker’s J. Bot. Kew Gard. Misc. 1: 243. 1849. TYPE: China. Hongkong, J.G. Champion s.n. (Holotype, not known).

Figure 2.86  Natural range of Actinidia latifolia var. latifolia.

2.22a  Actinidia latifolia var. latifolia 阔叶猕猴桃 (原变种) kuo ye mi hou tao (yuan bian zhong)

Actinidia championii Bentham in Fl. Hongk. 26. 1861. Actinidia gnaphalocarpa Hayata in Icon. Pl. Formos. 9: 97. 1922. TYPE: not designated. Actinidia latifolia var. tonkinensis (H.L. Li) H.L. Li in J. Arnold Arbor. 33: 52. 1952. A.tonkinensis H.L. Li in J. Arnold Arbor. 24: 366. 1943. TYPE: Vietnam. Tonkin, W.T. Tsang 29907 (Isotype, K) Actinidia guilinensis C.F. Liang in Guihaia 8(2): 129. 1988. TYPE: China. Guangxi: Guilin, May 21, 1987, C.F. Liang 34519 (Holotype, IBK). Distribution: China: Yunnan, Guangxi, Guangdong, Hainan, Guizhou, Hunan, Jiangxi, Fujian, Zhejiang, Sichuan, Hubei, and Hong Kong (Figure 2.86).

Morphological characteristics Large, vigorous deciduous climber. Stems  One-year-old shoots are grayish-green, covered with dense white tomentum; 2-year-old stems are reddish-brown, glabrous; lenticels are oblong-lanceolate to linear, pale; pith is light green or white, solid in younger stems, becoming hollow later. Leaves  Large, chartaceous; long elliptical to oblong ovate, 15-22 cm long and 10-13 cm broad; bilaterally asymmetrical; usually rounded at base, predominantly mucronate at apex, occasionally obtuse; margin has small, inconspicuous teeth. The upper leaf surface is dark glossy green, glabrescent or sparsely covered with appressed white pubescence; the veins are light green, covered with brown p ­ ubescence; secondary veins are

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Figure 2.87  Stem, emerging shoot, and leaves of Actinidia latifolia var. latifolia.

slightly elevated, same color as the rest of the leaf. The lower surface is light green, densely covered with white stellate hairs or pubescence; main and secondary veins are conspicuous and elevated, light greenish yellow. The petiole, c. 3-5 cm long, is light grayish brown and grayish-green, densely covered with white and brown tomentum (Figure 2.87). Pistillate flowers  Dichasium, up to 14 flowers per inflorescence. Primary peduncle is c. 14 mm long, secondary ­peduncle c. 10-12 mm long, pedicel c. 2 mm long; light green,

Figure 2.88  Flower and fruit of Actinidia latifolia var. latifolia.

covered with short grayish yellow pubescence. Flowers, c. 1.4 cm in diameter, usually in a tight spherical bunch. Calyx has two to three sepals, triangular, grayish-green, glabrous on inner surface, brown tomentum on outer surface, becoming reflexed after flower opens. Corolla has five petals, c. 9 mm long. reflexed, almost spindle-shaped, greenish-white, cream to warm pink. Numerous stamens, filaments are pale green, anthers are light yellow, nearly round, slightly pointed at one apex. Styles, usually 27-33 per flower, 3 mm long, pale green, with slightly swollen stigma. Ovary is nearly ellipsoid,

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2.3 mm in diameter, covered with short whitish pubescence. The pistillate flowers produce a sweet fragrance. Flowering is later than in most other Actinidia species. Staminate flowers  Dichasium, up to 70 flowers or more in inflorescence. Calyx with five sepals, long elliptical, reflexed, similar to those of pistillate flowers; with large gap between the petals. Stamens, 30-50 per flower, filaments are pale green, anthers are light yellow, short elliptical. Vestigial ovary is small, pale green, nearly globose. Fruit  Small, averaging c. 4 g, mostly cylindric or nearly round, 19-27 mm long and 11-14 mm in diameter. Fruit skin is brown, glabrous, with very dense and distinct fruit spots; persistent calyces; circle of persistent stylar remains sometimes shed. The stalk is short and stout. Fruit flesh is emerald-green with small, long elliptical, creamy green core. Numerous seed, brown, long elliptical. The fruit have an acid flavor, juicy. Vc content is very high, 670-2140 mg per 100 g fresh weight, soluble solids content is 10% and total acidity 1-2%. The fruit ripen in November in Wuhan (Figure 2.88).

by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The natural PIC of A. latifolia ranged from 0.644 to 0.830 revealed by SSR markers (Liu et al., 2007), while the average genetic diversity (h) and haplotype genetic diversity (H) based on cpSSR markers was 0.057 and 0.349, respectively (Zhang et al., 2007). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. latifolia in the National Germplasm Repository for Actinidia at WBG were 0.0197 and 0.0288, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58

2.22b  Actinidia latifolia var. mollis 长绒猕猴桃 chang rong mi hou tao

Actinidia latifolia var. mollis (Dunn) Handel-Mazzetti in Symb. Sin. 7: 391. 1931. Actinidia championii Benth. var. mollis Dunn in J. Linn. Soc., Bot. 39: 407. 1911. TYPE: China. Yunnan: Simao, A. henry 12041 (Isotype, A).

Diagnostic characters and species relationships

Distribution: Yunnan and Guangxi (Figure 2.89).

The species is distinct with its multibranched inflorescences forming dense clusters of flowers and its densely spotted fruit. As currently described, it is a very variable ­species in leaf characteristics, inflorescence size and branching and in fruit size.

Morphological characteristics

Actinidia latifolia is morphologically similar to A. lanceolata (Liang, 1984), but a close relationship is not supported

Figure 2.89  Natural range of Actinidia latifolia var. mollis.

Stems One-year-old shoots are light green, densely covered with brilliant red-yellow pubescence; pith is lamellate and off-white. Two-year-old stems are brown; the surface exposed to the sun is densely covered with coarse brown hairs in clusters, occasionally also with pubescence; lenticels are sparse, ovate or linear. Older stems

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Figure 2.90  Stem and leaves of Actinidia latifolia var. mollis.

are brown, covered with coarse brown hairs; lenticels are distinct and white, elliptical or round; pith is lamellate and pale green.

Leaves  Thick and chartaceous, c. 12-15  cm long and 9-11 cm broad; auriculate-cordate with about 18 mm deep retuse at base, covered with very fine downy hairs; apex mucronate. The upper surface is dark green, densely covered with minute, straight white, downy hairs; dense and small emerald-green dots alike on leaf blades; midvein is green, densely covered with long reddish-brown tomentum. The leaf margin is nearly entire but with some brown mucronate spines like spreading awns. The lower surface is grayish-green, very densely and thickly covered with white stellate hairs, almost like felt, the midvein and secondary veins are reddish yellow, densely covered with pubescence and long downy hairs. The petiole is light green, c. 3.74.6 cm long and 2 mm in diameter, densely covered with long hairs (Figure 2.90). Fruit Small, averaging 4-5 g, mostly short ellipsoid; five sepals are persistent; fruit skin is covered with short ­yellow tomentum; numerous small seed. The fruit ripen in ­ November. VC content is >50 mg per 100 g fresh weight; soluble solids content is c. 5% and total acidity c. 1% (Figure 2.91).

Diagnostic characters and species relationships It is distinguished from A. latifolia var. latifolia by the upper surface of the leaves being densely covered in soft downy hairs and the lower surface being very densely covered with long, puffy, stellate hairs. As a variety of A. latifolia, its relationships with other species are probably the same as those of A. latifolia var. latifolia.

2.23  Actinidia liangguangensis 两广猕猴桃 liang guang mi hou tao

Actinidia liangguangensis C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 322. 1984. TYPE: China. Guangxi: Jinxiu, LuoYun, alt. 680 m, June 25, 1956, H F. Qin 700307 (Holotype, IBK). Figure 2.91  Pistillate flowers and fruit of Actinidia latifolia var. mollis.

Distribution: China: Guangxi, Guangdong, and Henan (Figure 2.92).

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Kiwifruit: The Genus ACTINIDIA

Figure 2.92  Natural range of Actinidia liangguangensis.

Morphological characteristics Large, evergreen climber. Stems  The length of flowering shoots varies considerably from several cm to 40 cm or more, 2-5 mm in diameter; short shoots are densely covered with yellowish-brown tomentum, whereas longer shoots are sparsely covered with yellowish pubescence; occasional lenticels are small but very distinct. Two-year-old stems, c. 3.5-7 mm in diameter, are dark reddish-brown to black, glabrous; pith white, lamellate.

Figure 2.93  Stem and leaves of Actinidia liangguangensis.

Leaves  Soft leathery, ovate or oblong; 7-13 cm long and 4-9 cm broad; bilaterally symmetrical; base of leaves blunt, rounded or tending to cordate, apex acute or acuminate with spike (cuspidate), margin sometimes reddish, serrate with hard, sharp teeth. The upper leaf surface is green, glabrous, lustrous. The lower surface is grayish-green, densely covered with light yellowish-brown stellate hairs, tightly appressed; color of the tomentum darker on veins; eight to nine secondary veins on each side of midvein; lateral veins are relatively slender, but distinct; sub-lateral veins are hardly visible. The petiole, c. 2-7 cm long, is sparsely covered with dark-brown, downy hairs (Figure 2.93).

Chapter 2 Species

Flowers Cyme inflorescence with one to three flowers, predominantly single flowers. Peduncle is c. 2-7 mm long and pedicel c. 5-6 mm long, bracts are small, lanceolate, covered with soft, shaggy, yellow-brown hairs. Flowers are white, c. 1.5 cm in diameter. Calyx has five sepals, c. 4-5 mm long, oblong, dense tomentum on outer surfaces but inner surfaces nearly glabrous. Corolla has five petals, ground-­ladleobovate shaped, c. 9-10 mm long, rounded at apex and attenuate at base. Numerous, 34-36, stamens, filaments are slender, linear, c. 4-6 mm long, anthers are golden-­yellow, ovate to sagittate. Ovary of pistillate flowers is globose, covered with dense y­ ellow tomentum, styles are 4 mm long;

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in staminate flowers ovary is vestigial. Flowering is in late April to early May. Fruit  Small, averaging 2.4 g; mostly long globose c. 1.83.1 cm long, 1 cm in diameter; slightly protruding at apex, flat or slightly retuse at base; calyx persistent. Fruit skin is green; the fruit spots are small and round. Fruit initially densely covered with coarse hairs becoming partially shed on maturity. Fruit stalk is c. 2 cm long. Fruit flesh dark green, medium-sized core, numerous elliptical seed. Fruit have acid flavor, juicy and soft texture. VC content is 1056 mg per 100 g fresh weight, soluble solids content is 7% and total acidity 1.0%. In Wuhan, fruit ripen in November (Figure 2.94).

Diagnostic characters and species relationships The species is characterized by the leaves being densely stellate tomentose underneath, and dark green and glabrous above and the fruit being initially tomentose and always distinct lenticels. Actinidia liangguangensis was considered by Liang (1984) to be closely related to A. fulvicoma (Liang, 1984), and this is supported by molecular genetic data in the paternal lineage (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). Chromosomes: 2n = 2x = 58

2.24  Actinidia lijiangensis 漓江猕猴桃 li jiang mi hou tao

Actinidia lijiangensis C.F. Liang & Y.X. Lu in Guihaia 9(1): 21. 1989. TYPE: China. Guangxi: Guilin, May 1988, C.F. Liang 34544 (Holotype, IBK). Distribution: China: Upper reaches region of the Lijiang River, Guangxi (endemism) (Figure 2.95).

Morphological characteristics Large deciduous climber. Stems  One-year-old shoots are pale yellowish-brown densely covered with pale yellowish-brown stellate-downy hairs. Two-year-old stems are dark-brown, hairs on these or older stems are reddish-brown to dark-brown, but gradually shed, and eventually stems are glabrous; lenticels are oblong, conspicuous, pale yellow; pith is lamellate and white to pale brown.

Figure 2.94  Flower and fruit of Actinidia liangguangensis.

Leaves  Chartaceous; broad ovate-obovate or wide ovate, leaf blade is 4.5-12 cm long and 4-12.5 cm broad; cordate at base, short acuminate or acute at apex. The leaf margin is denticulate, hard. The upper surface of young leaves is lustrous, sparsely covered with stellate, downy hairs; the lower leaf surface is densely covered with ash-white, stellate, downy hairs. The hairs on older leaves are readily shed. The

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Kiwifruit: The Genus ACTINIDIA

Figure 2.95  Natural range of Actinidia lijiangensis.

veins on the lower leaf surface are conspicuous, raised, six to nine secondary veins on each side of midvein. The petiole is 3-6.5 cm long, covered with pale yellowish-brown stellate, downy hairs when young, and gradually shed (Figure 2.96). Pistillate flowers  Cyme inflorescence with one to three flowers. The peduncle is c. 1.5-2 cm long; pedicel is c. 2 cm long. Calyx has four to five sepals, broadly ovate, densely covered with powdery tomentum on both inner and outer surfaces. Flowers are yellowish white; five to six petals, obovate, c. 12-15 mm long. Numerous, c. 25-28, stamens, filaments c. 5 mm long; anthers are yellow. Usually 16-25 styles, 7 mm long, connected at the base. Ovary is globose, 5 mm in diameter, densely covered with pale yellow-white pubescence. Fruit Mostly cylindrical, 4-5 cm long, 2-2.5 cm in diameter; average weight 28 g. Fruit skin is green, initially covered with light yellow-white tomentum but readily shed; lenticels are dense, pale brown, like spots on fruit surface. Calyx is persistent and reflexed. Fruit stalk is c. 2.0-2.5 cm long. Fruit flesh is dark green with medium-sized core; lightly acid flavor and slightly juicy; Vc content is 60 mg per 100 g fresh weight; soluble solids content is 14% and total acidity 1.1%. The fruit ripen in December (Figure 2.97).

Diagnostic characters and species relationships

Figure 2.96  Stem and leaves of Actinidia lijiangensis.

The species is distinct in its leaves are broadly ovate to broadly obovate to orbicular, underneath sparsely stellate tomentose, readily becoming glabrous, and its fruit are narrowly cylindrical.

Chapter 2 Species

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2.25  Actinidia linguiensis 临桂猕猴桃 lin gui mi hou tao

Actinidia linguiensis R.G. Li & X.G. Wang in Guihaia 23(3): 200. 2003. TYPE: China. Guangxi: Guilin, September 2001, R.G. Li 0110 (Holotype, IBK). Actinidia wantianensis R.G. Li & L. Mo. in Guihaia 23(3): 201. 2003. TYPE: China. Guangxi: Guilin, September 2001, R.G. Li 0111 (Holotype, IBK). Distribution: Endemic to Guangxi, China.

Morphological characteristics Large deciduous climber. Stems  One-year-old branchlets are velutinous becoming glabrescent; lenticels conspicuous. Two-year-old shoots are glabrous; lenticels conspicuous; pith brown, lamellate. Leaves  Large, 8-14 cm long, 4.5-10 cm broad; ovate to orbicular, papery; base tends toward cordate; leaf margin is serrate; apex is acuminate; eight lateral veins on each side of midvein. The upper leaf surface is bright green, initially pubescent, but hairs shed becoming glabrous; the lower leaf surface has stellate tomentum, largely lost becoming glabrescent. The petiole is 4-6 cm long, velutinous becoming glabrescent. Flowers  Inflorescences contain one to three flowers; velutinous; peduncle is 0.6-1.4 cm; pedicel 1-1.8 cm. Calyx has four to six sepals, broadly ovate, 5-7 mm long, 4-6 mm wide, velutinous. Petals five to seven, ovate or narrowly ovate, 1013 mm long, 7-9 mm wide, pink or pinkish yellow. Anthers yellow. Ovary globose, c. 4 mm in diameter, white tomentum. Flowering April-June. Fruit  Fruit oblong to ovoid, c. 1.7 cm long; dense tomentum of stiff hairs; persistent sepals. Fruit mature OctoberNovember (Figure 2.98).

Diagnostic characters and species relationships The species is characterized by leaves which are ovate to orbicular, with sparse stellate tomentum underneath; fruit are oblong to ovate, densely tomentose. Figure 2.97  Pistillate flowers and fruit of Actinidia lijiangensis.

Actinidia lijiangensis was considered by Liang and Lu (1989) to be closely related to A. chinensis, and this is supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. lijiangensis in the National Germplasm Repository for Actinidia at WBG were 0.0907 and 0.1325, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58

Actinidia linguiensis is considered to be closely related to A. zhejiangensis (Li, 2003).

2.26  Actinidia longicarpa 长果猕猴桃 chang guo mi hou tao

Actinidia longicarpa R.G. Li & M.Y. Liang in Guihaia 23(3): 198. 2003. TYPE: China. Guangxi: Guilin, September 2001, R.G. Li 0108 (Holotype, IBK). Actinidia rubrafilmenta R.G. Li & J.W. Li in Guihaia 23(3): 198. 2003. TYPE: China. Guangxi: Guilin, September 2001, R.G. Li 0109 (Holotype, IBK).

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Kiwifruit: The Genus ACTINIDIA

Leaves Large, 6.2-19 cm long, 5.4-11.5 cm wide; ovate to orbicular; papery; leaf margin finely serrate; apex acuminate to shortly caudate; six to eight secondary veins on each side of midvein. Leaf upper surface lustrous, glabrous; lower leaf surface has stellate tomentum largely lost to become glabrescent. Petiole 3.5-6 cm, initially velutinous, becoming glabrescent. Flowers  Inflorescences have one to seven flowers; peduncle c. 1.2 cm; pedicel 0.9-2 cm, velutinous. Calyx has three to six sepals, ovate, 6-7 mm long, 3-5 mm wide, velutinous. Corolla has five petals, narrowly ovate, 1.4-1.7 cm long, 0.8-1.3 cm wide; petals pink, or apical part white and lower part red. Anthers yellow. Oblong ovary, white tomentum. Flowers April-June. Fruit  Oblong, 2-4 cm long, dense soft tomentum, sometimes with persistent reflexed sepals. Fruit mature October (Figure 2.99).

Figure 2.98  Stem, leaves, pistillate flowers, and fruit of Actinidia linguiensis.

Distribution: Endemic to Guangxi, China.

Figure 2.99  Stem, leaves, flowers, and fruit of Actinidia longicarpa.

Morphological characteristics Large deciduous climber.

Diagnostic characters and species relationships

Stems One-year-old branchlets initially velutinous, becoming glabrescent, lenticels subconspicuous; 2-year-old stems glabrous, lenticels subconspicuous; pith white, lamellate.

The species is characterized by ovate to orbicular leaves, sparsely stellate tomentose on lower surface, and fruit which are usually oblong and densely tomentose.

Chapter 2 Species

Actinidia longicarpa appears to be closely related to A. eriantha (Li, 2003).

2.27  Actinidia macrosperma 大籽猕猴桃 da zi mi hou tao

Actinidia macrosperma C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 311. 1984. T YPE: China. Zhejiang:

115

Tianmushan, October 18, 1958, Hangzhou Bot. Gard. 31236 (Holotype, PE).

2.27a  Actinidia macrosperma var. macrosperma 大籽猕猴桃 (原变种) da zi mi hou tao (yuan bian zhong)

Distribution: China: Guangdong, Guangxi, Zhejiang, and Hubei (Figure 2.100).

Figure 2.100  Natural range of Actinidia macrosperma var. macrosperma.

Morphological characteristics Small to medium-sized, deciduous climber. Stems Current season’s canes are light green, about 5-20 cm long, 2-2.5 mm in diameter; glabrous or sparsely covered with rusty-brown pubescence on basal part of the canes; lenticels are indistinct or just visible; occasionally, spine-like remains of broken pedicels can been seen in the leaf axil. Buds are glabrous. Two-year-old stems are greenish-brown; lenticels are small and sparse, hardly visible; pith is solid, white. Leaves Young growing leaves are membranaceous, coriaceous when they are fully mature; ovate or nearly oblong, 3-8 cm long; basal part is wide cuneate or round; bilaterally symmetrical or slightly asymmetrical; apex is acuminate, acute to obtuse. Leaf

margin is serrated as spreading teeth or round dull teeth; margin is entire when leaves are mature. Upper surface of leaf is green, glabrous, glossy; lower surface is light green, occasionally with bristly hairs in the vein axils, veins are not very distinct, four to five secondary veins on each side of the midvein. Petiole is light red, 10-22 mm long, glabrous (Figure 2.101). Pistillate flowers  Predominantly single, c. 3 cm in diameter. Peduncles are about 9-15 mm long, glabrous or partially covered with sparse pubescence. The bracts are lanceolate or linear, minute, 1-2 mm long, with a few marginal glandular hairs. Calyx has two to three sepals ovate or long ovate, 6-12 mm long, green; ­glabrous on both surfaces. Corolla of 5-12 petals, reflexed orbicular-obovate, 10-15 mm long, pure white. Ovary is bottle-shaped, 6-8 mm long, about

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Kiwifruit: The Genus ACTINIDIA

Figure 2.101  Stem, emerging shoot, and leaves of Actinidia macrosperma var. macrosperma.

7 mm in diameter, pale green, glabrous, with 5 mm long styles. Numerous stamens, anthers yellow. Flowers are very fragrant. Flowering in mid-May. Staminate flowers White, 2 cm in diameter.

Fruit  Ovoid or ovoid-globose shaped, 3-3.5 cm long, with small beak at apex; sepals are sometimes persistent; glabrous. No fruit spots. Seeds are particularly large, 4-5 mm long. Fruit ripen early-mid October to become bright orange, fruit flesh is also orange when ripe (Figure 2.102).

Figure 2.102  Pistillate and staminate flowers and fruit of Actinidia macrosperma var. macrosperma.

Chapter 2 Species

Diagnostic characters and species relationships The species is characterized by its spotless fruit, orange when ripe, and the very large seed, 4-5 mm long. Actinidia macrosperma has been considered to be a distinct species, not morphologically close to any other species (Liang, 1984; Li et al., 2007a,d). However, a close relationship between A. macrosperma and A. valvata is indicated by molecular genetic data in maternal lineage (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. macrosperma in the National Germplasm Repository for Actinidia at WBG were

117

0.0276 and 0.0403, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 4x = 116

2.27b  Actinidia macrosperma var. mumoides 梅叶猕猴桃 mei ye mi hou tao

Actinidia macrosperma var. mumoides C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 312. 1984. TYPE: China. Zhejiang: Hangzhou, 1958, S.Y. Chang 2159 (Holotype, PE). Distribution: China: Yunnan, Jiangxi, Zhejiang, Anhui, and Jiangsu (Figure 2.103).

Figure 2.103  Natural range of Actinidia macrosperma var. mumoides.

Morphological characteristics Stems  One-year-old stems are light green, glabrous, some light brown at connection of petioles and stems; sparse lenticels, linear, white; pith is lamellate or solid, white; internodes are 1.2-2.7 cm long. Two-year-old stems are dark reddish-brown, relatively hard, glabrous and smooth; lenticels are long to linear, pale gray-yellow. Current season’s shoots are relatively hard but slim. Shoots, when carrying flowers, are c. 3-15 cm long. Leaves Chartaceous, long elliptical, occasionally broad lanceolate, 6-12 cm long, 3-5 cm broad. Leaves on apical part of current shoots are lanceolate, caudate with a long, tapering tail; basal leaves are green, younger leaves at stem tip are pale purple-brown. Upper leaf surface is smooth, primary veins are light green, glabrous. The midvein has sparse brown pubescence. The basal part (c. 1/3) of leaf margin is entire, the apical part (two thirds) is serrated with small teeth,

light green or brown spines, appressed. Lower leaf surface is smooth, green, glabrous. Veins distinct, raised, six to eight secondary veins on each side of midvein. Petiole, c. 2 cm long, slender, reddish color, sparsely covered with short, light brown pubescence (Figure 2.104). Pistillate flowers  Flowering shoots are c. 10-20 cm long. Pedicel is c. 11-12 mm long, slender, light green, sparsely covered with pale brown pubescence. Flowers single, white, 13 mm in diameter. Calyx has five sepals, c. 4 mm long, 2 mm broad, nearly oblong, obtuse at apex, light green, sparsely covered with pale brown or white pubescence. Corolla has five to six petals, predominantly five, c. 8 mm long and 6 mm broad, obovate, truncate at base, spatulate, pale green at base, otherwise white, with radiating stripes, wider in middle part of petal, then branching. Numerous, c. 30, stamens, filaments are 3 mm long, slender, anthers yellow, long oblong, c. 1 mm long. Ovary is spherical, slightly

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Figure 2.104  Stem and leaves of Actinidia macrosperma var. mumoides.

flattened, c. 2.5 mm long, 3 mm broad, white densely covered with white pubescence, c. 27 styles, pale yellow-green, slightly swollen stigmas. Full bloom was observed on April 12, 1988 in Pingbian, Yunnan (Figure 2.105). Staminate flowers Cyme inflorescence, one to three flowers per inflorescence, usually three, rarely one, arising from leaf axil. Calyx has four to seven sepals, green, sparsely covered with white and light brown pubescence, ovate, apex acuminate. Flowers are 13 mm in diameter; four to six, mostly five, petals, 9 mm long, 6 mm broad, broad obovate, spatulate, acuminate at apex, with radiating stripes from the basal part, the stripes wider in middle part of petal and branching at their ends. Pedicel is slender, c. 5-8 mm long, light green, sparsely covered with light brown pubescence. The pedicel of the apical (king) flower is relatively longer. Peduncle is about 9-12 mm long. Numerous, 26-29 stamens, filaments greenish-white, c. 4 mm long, anthers are yellow, elliptical, 1 mm long. Pistil is vestigial, ovary is very small, spherical, densely covered with white pubescence. Full bloom was observed on April 9, 1989 in Pingbian, Yunnan.

Fruit Nearly globose, c. 2.6 cm long, diameter 2.7 cm, average weight c. 11 g; green when unripe; glabrous. The fruit apex is almost flat, stamens and sepals shed. Fruit stalk is 1.8 cm long, brown, glabrous, with sparse y­ ellowish-brown spots. Fruit flesh when unripe is green, small white core. Seed are particularly large, oblong. The fruit have a very acid taste, not astringent, but with tart and chili hot tastes, juicy. VC content is 36 mg/100 g fresh weight, soluble solids content is 5.4% and total acidity 0.98%.

Diagnostic characters and species relationships A. macrosperma var. mumoides differs from A. macrosperma var. macrosperma in its leaf midvein and petiole being sparsely setose. As a variant of A. macrosperma, its specific relationships with other species are probably the same as those of A. macrosperma var. macrosperma. Chromosomes: 2n = 4x = 116

2.28  Actinidia melanandra 黑蕊猕猴桃 hei rui mi hou tao

Actinidia melanandra Franchet in J. Bot (Morot). 8: 278. 1894. TYPE: China. Chongqing: Chengkou, R.P. Farges s.n. (Holotype, P; Isotype, K).

2.28a  Actinidia melanandra var. melanandra 黑蕊猕猴桃 (原变种) hei rui mi hou tao (yuan bian zhong)

Figure 2.105  Pistillate flowers of Actinidia macrosperma var. mumoides.

Actinidia hypoglauca C. Pei & Y.W. Law in Bot. Bull. Acad. Sin. 2: 27. 1948. TYPE: China. Sichuan: Ebian, T.H. Chao 304 (Holotype, NAS). Actinidia changii P.S. Hsu in Act. Phytotaxon. Sin. 11(2): 197. 1966. TYPE: China. Zhejiang: Ruichang, May 3, 1959, Chekiang Prov. Econ. Bot. Exped. 2583 (Holotype, HZBG). Actinidia viridiflava P.S. Hsu in Act. Phytotaxon. Sin. 11(2): 198. 1966, excl. S.Y. Chang 6126. TYPE: China. Zhejiang: Tianmushan, May 26, 1957, Y.Y. Ho 21869 (Isotype, HZBG).

Chapter 2 Species

Actinidia henanensis C.F. Liang in Guihaia 2(1): 1. 1982. TYPE: China. Henan: Lushi, Xiong’ershan, June 1981, J.X. An & S.G. Zheng 8017 (Holotype, IBK). Actinidia melanandra var. cretacea C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 310. 1984. TYPE: China. Hubei: Xingshan, Sanchakou, July 15, 1957, H.J. Li 1088 (Holotype, IBSC; Isotype, PE). Actinidia melanandra var. subconcolor C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 310. 1984. TYPE: China.

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Zhejiang: Tiantaishan, Hangzhou Botanical Garden 0281 (Holotype, IBK). Actinidia globosa C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 309. 1984. TYPE: China. Guangxi: Ziyuan, October 5, 1937, J.X. Zhong 83575 (Holotype, IBSC). Distribution: China: Yunnan, Guizhou, Hunan, Jiangxi, Fujian, Zhejiang, Sichuan, Chongqing, Hubei, Gansu, Shaanxi, and Henan (Figure 2.106).

Figure 2.106  Natural of Actinidia melanandra var. melanandra.

Morphological characteristics Medium-sized to large, deciduous climber. Stems  One-year-old shoots are grayish brown; lenticels are oblong, white or green. Two-year-old stems are grayish brown; lenticels are oblong, grayish brown; pith is lamellate, greenish-brown. Leaves Chartaceous to thin leathery, ovate to long ovate; leaf is 7-11 cm long and 3.5-4.5 cm broad; base cuneate and apex acute to acuminate. The upper surface is dark green and glabrous, lustrous; apical half of leaf margin is densely serrate. The lower surface glaucous, glabrous except for tufts of hairs in the axils of the lateral veins (barbate). The petiole is light red, 3-6 cm long (Figure 2.107). Pistillate flowers  Cyme inflorescence with one to three flowers; pedicel is c. 1.7 cm long, green, glabrous. Calyx has five sepals, oblong, c. 7 mm long, 3 mm broad; light ­purplish-red, long pubescence on outer surface, glabrous

on inner surface. Flowers are c. 2.2 cm in diameter; five petals, greenish to ivory white, oblong, c. 13 mm long, without radiating stripes on the surface. Numerous stamens, c. 63, filaments c. 3 mm long, white, anthers are black. Styles are white, 2.5 mm long with slightly swollen stigma. Ovary is bottle shaped, glabrous, 3 mm in diameter. In Wuhan, flowering is in May. Staminate flowers Cyme inflorescence, three flowers, pedicel is c. 1-2 cm long. Calyx has five sepals; corolla is usually smaller than that of pistillate flowers, c. 1.5-2.0 cm ­diameter, five petals, white, broad ovate. Numerous (37) stamens, ­filaments are 3 mm long, anthers black, oblong. Ovary is very small, vestigial, green. Fruit Fruit globose to oblong with long beak; 2-4.5 cm in length; average weight 15 g; fruit skin is dark purple and glabrous; no lenticels; calyx shed. The stalk is c. 2.6 cm long. Fruit flesh is green with small, elliptical, whitish core. Numerous seed are purplish-red to black, elliptical,

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Figure 2.107  Stem, emerging shoots, leaves, and fruit of Actinidia melanandra.

smooth seed coat. The fruit ripen in September, sweet flavor, medium juicy. VC content is c. 100-200 mg per 100 g fresh weight, soluble solids content is 14% and total acidity 0.9% (Figure 2.108).

Figure 2.108  Pistillate and staminate flowers and fruit of Actinidia melanandra.

Diagnostic characters and species relationships The species is distinct in its leaves papery to thin leathery, glaucous underneath, black anthers and spotless fruit.

Chapter 2 Species

Morphologically, Actinidia melanandra appears to be closely related to A. arguta and A. hypoleuca (Li, 1952; Liang, 1984; Li et al., 2007a,d), and this is supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. melanandra in the National Germplasm Repository for Actinidia at WBG were 0.0079 and 0.0115, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58, 4x = 116

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2.28b  Actinidia melanandra var. glabrescens 无髯猕猴桃 wu ran mi hou tao

Actinidia melanandra var. glabrescens C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 310. 1984. TYPE: China, Hunan: Hengshan, June 1943, Z.D. Chen 40 (Holotyoe, IBSC). Distribution: China: Yunnan, Guizhou, and Hunan (Figure 2.109).

Figure 2.109  Natural range of Actinidia melanandra var. glabrescens.

Morphological characteristics Stems  One-year-old shoots are russeted and glabrous, lenticels are elliptical or linear, pale yellow, and protrude from the surface; internodes 8-17 mm long. Two-year-old stems are reddish-brown, glabrous; lenticels are elliptical, ­yellow-brown; pith is lamellate, brown. Leaves  Chartaceous, narrowly elliptical; leaf blade is 5-11 cm long and 3.5-4 cm broad; bilaterally symmetrical; base wide cuneate; acuminate or acute at apex. The upper leaf surface is green and glabrous; midvein and secondary veins are yellowish green; leaf margin is serrated into small teeth densely appressed, green. The lower leaf surface is

glaucous and glabrous; midvein and secondary veins are light green, glabrous, no hairs in axils of veins. The petiole is purplish-red, glabrous, 2.5-3.5 cm long. Fruit Fruit are c. 2.8 cm long and 2.3 cm in diameter, average weight c. 8 g; short cylindrical with beak; fruit skin ­purplish-red, glabrous and without spots. Fruit flesh is pale green, core pale yellow; good flavor with distinctive aroma. Seed are small. In Leishan, Guizhou, the fruit ripen in September.

Diagnostic characters and species relationships It differs from A. melanandra var. melanandra in that on the lower surface, there are no hairs in the axils of the veins.

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As a variant of A. melanandra, its specific relationships with other species are probably the same as those of A. melanandra var. melanandra.

2.29  Actinidia melliana 美丽猕猴桃 me li mi hou tao

Actinidia melliana Handel-Mazzetti in Anz. Akad. Wiss. Wien, Math.-Naturwiss. Kl. 59: 57. 1922. TYPE: China. Guangdong: Longtoushan, alt. 800 m, April 5, 1917, R. Mell 22 (Isotype, A). Distribution: China: Guangxi, Guangdong, Hainan, Hunan, and Jiangxi (Figure 2.110).

Morphological characteristics Medium-sized, semi-evergreen to deciduous climber. Stems Flowering shoots are 2-4 cm long, 2 mm in diameter, elongating shoots 30-40 cm long. One-year-old shoots and 2-year-old stems are both densely covered with long, stiff, rusty-brown hairs, 6-8 mm long; lenticels are conspicuous, pale yellow; pith is lamellate, white.

Figure 2.110  Natural range of Actinidia melliana.

Leaves  Membranaceous to hard chartaceous, 2-year-old leaves are coriaceous, long oblong, long-lanceolate or long-obovate, 6-15 cm long, 2.5-9 cm broad. The leaves are shallow and subcordate to auriculate-subcordate at base, short acuminate to long acuminate at apex, and with small stiff teeth on the margin that is usually reflexed toward the apex. Hairs on both leaf surfaces, sometimes becoming reduced in older leaves to mainly along veins. The upper leaf surface has long stiff hairs, and the lower surface is green or glaucous with short, stiff appressed hairs. Midvein and secondary veins are conspicuous, but reticulate veinlets are hardly visible. Petioles are 1018 mm long, covered with long, hard, rusty-brown hairs (Figure 2.111). Flowers  Axillary cymose inflorescences with up to 10 flowers. Peduncle is 3-10 mm long, two-branched, covered with long, stiff, rusty-brown hairs. Calyx has five sepals, 4-5 mm long, oblong ovate, obtuse to acute at apex, thinly covered with downy hairs on outer surface. Corolla has five petals, 8-9 mm long, 5-7 mm broad, obovate, with rounded apex, pale pink to white. Numerous stamens, filaments are c. 2.5 mm long, anthers are yellow, 1 mm long. Ovary is almost globose, densely covered with

Chapter 2 Species

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Figure 2.111  Stem and leaves of Actinidia melliana.

rusty-brown tomentum, styles are 3 mm long. Flowering in May-June. Fruit Small fruit, 1.7-2.9 cm long and 0.9-1.2 cm in diameter, average weight 2 g, maximum weight 3.7 g; cylindrical, almost flat at stylar end or slightly depressed in center. Sepals sometimes are shed but stylar remains persistent. Skin is light green, thin, with many conspicuous pale brown spots, round; covered with sparse, short, coarse hairs but becoming glabrescent. Fruit stalk is 1.5-2 mm long. Fruit pulp is green, soft, with medium-sized core, not juicy, without aroma. Seeds are flat ovate, dark-brown, c. 500 seed per fruit. Vc content is 45 mg per 100 g fresh weight, sugar content is 1.46% and total acidity is 2.51% (Figure 2.112).

Diagnostic characters and species relationships The species is characterized by its branchlets, petioles and leaves having stiff, rusty-brown hairs and the underneath of the leaves being clearly glaucous. Actinidia melliana was thought by Li (1952) to be more closely related to A. hemsleyana than any other Actinidia species, but this is not supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. melliana in the National Germplasm Repository for Actinidia at WBG were 0.0473 and 0.0691, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58

2.30  Actinidia obovata 倒卵叶猕猴桃 dao luan ye mi hou tao

Actinidia obovata Chun ex C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 323. 1984. TYPE: China. Guizhou: Qingzhen, May 21, 1936, S.W. Deng 90347 (Holotype, IBSC). Figure 2.112  Pistillate flowers and fruit of Actinidia melliana.

Distribution: China: Yunnan and Guizhou (Figure 2.113).

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Figure 2.113  Natural range of Actinidia obovata.

Morphological characteristics Large, deciduous climber. Stems  One-year-old shoots are green, sparsely covered with pale brown hairs; lenticels are inconspicuous, elliptical or linear, pale brown; internodes 3-5 mm; buds densely covered with yellow-brown hairs. Two-year-old stems are brown, sparsely covered with short, coarse hairs; lenticels are round or oblong, yellow-brown; pith is lamellate, brownish white. Leaves Chartaceous; lamina is 9.5-13 cm long, and 8-10 cm broad; obovate or nearly round to obtriangular; bilaterally symmetrical or asymmetrical; rounded, subcordate or cuneate at base; cuspidate or truncate at apex. The upper surface is dark green and almost glabrescent; midvein and secondary veins are pale green and covered with light brown tomentum; hairs on the basal part of main veins are denser; dentation on the apical part of leaf margin ends in sharp thorns which are green, brown at tip. The lower surface is pale green, densely covered with white tomentum; midvein and secondary veins are green, densely covered with white and pale brown tomentum. The petiole, c. 4.5-7 cm long, is purplish-red, covered with pale brown tomentum (Figure 2.114). Pistillate flowers Predominantly single. Calyx has five, rarely six, sepals, c. 8 mm long and 6 mm broad, elliptical or ovate,

Figure 2.114  Leaf of Actinidia obovata.

both surfaces covered with white or pale brown tomentum. Flowers are c. 4.7 cm in diameter, five petals, c. 20 mm long, 18 mm broad, nearly round or obovate; white becoming yellow or apricot-yellow. Numerous, c. 125, stamens; filaments are white, 10 mm long; anthers are yellow, elliptical to sagittate. Styles are white, usually 31, 5 mm long with slightly swollen stigma. Ovary is short, cylindrical, 8 mm in diameter, covered with white or pale brown pubescence. Pedicel is c. 2 cm long, green.

Chapter 2 Species

Staminate flowers Cyme inflorescence with three flowers, sometimes single. Calyx has five or six sepals, 7 mm long, 5 mm broad; elliptical or ovate; brown. Opening flowers are white, changing into apricot-yellow after 1 day; corolla is smaller than that of pistillate flowers, c. 4.0 cm diameter; five, occasionally six or seven petals, c. 2 cm long, 1.7 cm broad; obovate, with longitudinal stripes. Numerous, c. 120, stamens; filaments are white; anthers are yellow, oblong to sagittate. Vestigial ovary is covered with white or pale brown pubescence. Pedicel, c. 2 cm long, is green, covered with light brown pubescence. Fruits Cylindrical fruit, c. 2.7-4.5 cm long, 2.3-4.5 cm in diameter, average weight c. 8-23 g. Fruit apex is with truncate beak, stylar remains persistent; shoulders rounded, sepals persistent. Fruit skin is grayish brown, initially with brown tomentum but becoming glabrous. Fruit stalk, 1.7-3.4 cm long, is green or greenish-brown, glabrous. Fruit flesh is green or yellow; small white core; acid flavor, juicy, fine texture. Fruit ripen in September.

Diagnostic characters and species relationships The species is distinct in its leaves are obovate to obovate-obtriangular, the lower surface of the leaves has sparse stellate hairs, readily shed, and the mature fruit are glabrous. Actinidia obovata was thought by Liang (1984) to be related to A. callosa var. henryi, A. chinensis, and A. sorbifolia.

2.31  Actinidia persicina 桃花猕猴桃 tao hua mi hou tao

Actinidia persicina R.G. Li & L. Mo in Guihaia 23(3): 197. 2003. TYPE: China. Guangxi: Guilin, September 2001, R.G. Li 0107 (Holotype, IBK). Distribution: Endemic to Guangxi, China.

Figure 2.115  Stem, leaves, and fruit of Actinidia persicina.

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Morphological characteristics Large, semi-evergreen to deciduous climber. Stems One-year-old branches are covered with brown velutinous hairs, lenticels conspicuous; 2-year-old branches are sparsely velutinous to glabrescent; lenticels oblong or ovate, grayish; pith initially white becoming brown, lamellate. Leaves  Lamina, papery to almost leathery in older leaves; 9-20 cm long, 5-9.5 cm broad; long ovate to broadly ovate; base rounded to cordatulate; apex acuminate to mucronate; margins finely serrate. Upper leaf surface is green, lustrous, sparsely covered with short, coarse hairs, denser on midvein and veins but becoming lost, older leaves are glabrous above. Underneath, leaves are glaucous toward the margins; sparse stellate hairs initially but then lost, soon glabrescent; six to eight veins on each side of midvein. Petiole 2.5-6 cm long, densely velutinous when leaf is young, becoming glabrescent; green or pale purple-pink. Pistillate flowers Inflorescences one- to three-flowered, brown velutinous; peduncle 3-7 mm; pedicel 6-13 mm. Calyx has three to five sepals, 5 mm long, 3 mm broad, ovate, both surfaces with dense brown tomentum. Flowers are 3.5-4.5 cm in diameter, five petals, c. 9 mm long, 5 mm broad, bright pink fading toward margins. Anthers yellow. Ovary almost spherical, c. 6-8 mm in diameter, white tomentum. Flowering in April. Fruit Ovoid or oblong, cylindrical, c. 3-3.4 cm long, 2.4-2.8 cm in diameter; weight 13-18 g; sparsely velutinous; persistent sepals not reflexed. Fruit mature October (Figure 2.115).

Diagnostic characters and species relationships The species is distinct in that the underneaths of its leaves are glaucous near the margin, stellate tomentose when young, becoming glabrescent, and fruit ovoid or oblong, with sparse, short brown hairs.

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Actinidia persicina is related to A. rongshuiensis and A. eriantha (Li, 2003). Chromosomes: 2n = 2x = 58

2.32  Actinidia pilosula 贡山猕猴桃 gong shan mi hou tao

Actinidia pilosula (Finet & Gagnepain) Stapf ex HandelMazzetti in Symb. Sin. 7: 390. 1933. Actinidia callosa Lindley var. pilosula Finet & Gagnepain in Bull. Soc. Bot. France 52 (Mém. 4): 19. 1906. TYPE: China. Yunnan: Gongshan, June 1893, R.P. Soulie 1396 (Isotype, NY). Actinidia kungshanensis C.Y. Wu & S.K. Chen in C.Y. Wu in Fl. Yunnan. 1: 71. 1977. TYPE: China. Yunnan: Gongshan, alt. 2200 m, in forest, September 17, 1938, T.T. Yu 23001 (Holotype, KUN; Isotype, PE). Distribution: Endemic to Yunnan, China (Figure 2.116).

Morphological characteristics Medium-sized, deciduous climber. Stems  One-year-old shoots are greenish-brown, sparsely covered with pale purple downy hairs; lenticels are relatively sparse, inconspicuous, linear, yellow; internodes 5-25 mm

Figure 2.116  Natural range of Actinidia pilosula.

long. Two-year-old stems are reddish-brown, glabrous and smooth; lenticels are abundant, small, grayish-white, elliptical; pith is lamellate, white to brownish. Leaves  Thick and chartaceous, long elliptical; 7.5-15 cm long and 4.5-6.8 cm broad; broadly truncate at base, often unequal; mucronate or long attenuate at apex. The upper leaf surface is bright green, lustrous, glabrous. Midvein and secondary veins are green, depressed, sparsely covered with light purplish-red, downy hairs when leaf is young, later glabrous. The margin is dentate, teeth large; on middle and apical parts of leaf, dentation is alternate with one to three small teeth between larger teeth; teeth are purple, mostly appressed; basal part of leaf margin entire. The lower leaf surface initially covered with white hairs, either simple or branching, partially lost. The midvein, secondary veins, 7-11 per side, and veinlets are whitish-green, elevated, densely covered with white hairs. Secondary veins anastomose at leaf margin. The petiole, c. 2.6-3.5 cm long, is purplish-red, sparsely covered with purplish-red and white tomentum, becoming glabrous (Figure 2.117). Pistillate flowers  Cyme with more than two subsidiary axes, three to seven flowers. Peduncles are thick, 1.5-2.2 cm long, sparsely covered with pale purplish-red tomentum; pedicel is light purplish-red or green, densely covered with light purplish-red downy hairs, c. 0.5 cm long. Calyx has

Chapter 2 Species

127

Figure 2.117  Stem, over-wintering bud, and leaves of Actinidia pilosula.

three to five sepals, c. 8 mm long and 6 mm broad, elliptical, both inner and outer surfaces glabrous to densely covered with purplish-red downy hairs. Flowers are c. 3.3 cm in diameter; six petals, c. 15 mm long, 9 mm broad, elliptical, reflexed; broad petal base and slightly retuse; light yellow, densely covered with yellow-red, radiating stripes to the margins, the stripes forked at tip. Numerous stamens, c. 26, filaments, 4 mm long, are white, anthers are yellow, sagittate or elliptical. Usually 30 styles, 5 mm long, with slightly swollen stigma. Ovary is nearly globose, 4 mm long and 5 mm in diameter, densely covered with pale purplish-red downy hairs. Staminate flowers Cyme inflorescence, five to seven flowers. Peduncles are thick, 7 mm long, green, densely covered with pale purplish-red tomentum; pedicel, c. 6 mm long, is pale purplish-red, densely covered with pale purplish-red downy hairs. Calyx has five sepals, c. 4 mm long, 2 mm broad, elliptical, green. Opening flowers are pale yellowish white; corollas are smaller than those of pistillate flowers, c. 2.0 cm in diameter, five petals, c. 10 mm long, 5 mm broad, obovate. Numerous stamens, 25-28, filaments are white, c. 5 mm long, anthers are yellow, elliptical, c. 2 mm long. Ovary is vestigial, densely covered with pale purplish-red pubescence.

Figure 2.118  Pistillate flowers and fruit of Actinidia pilosula.

Fruit Globose to long cylindrical, c. 2.3 cm in diameter; fruit skin is greenish-brown and becoming glabrous; lenticels are white to pale brown, round. Styles and sepals are usually persistent. Seeds are small, 2 mm long. Fruit flesh and core green. Fruit ripen at the end of September to early October (Figure 2.118).

Diagnostic characters and species relationships The species is distinct in the lower surface of its leaves having dense white tomentum, the midvein and lateral veins being conspicuous and the fruit being spotted, without hairs.

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Actinidia pilosula is thought to be related to A. venosa and A. trichogyna (Li, 1952; Liang, 1984).

2.33  Actinidia polygama 葛枣猕猴桃 ge zao mi hou tao

Actinidia polygama (Siebold & Zuccarini) Maximowicz in Mém. Acad. Imp. Sci. St.-Pétersbourg Divers Savans 9: 64. Trochostigma polygama Sieb. & Zucc. in Abh. Math.Phys. Cl. Königl. Bayer. Akad. Wiss. 3(2): 728. 1843 [“polygama”]. Actinidia polygama var. lecomtei (Nakai) H.L. Li in J. Arnold Arbor. 33: 22. 1952.-Actinidia lecomtei Nakai in Bot. Mag. (Tokyo) 47: 253. 1933.

Figure 2.119  Natural range of Actinidia polygama.

Distribution: China: Yunnan, Guizhou, Hunan, Fujian, Zhejiang, Sichuan, Chongqing, Hubei, Anhui, Gansu, Shaanxi, Henan, Shandong, Beijing, Liaoning, Jilin, Heilongjiang. Large part of Japan (Figure 2.119).

Morphological characteristics Large, deciduous climber. Stems  One-year-old stems are slender but hard, brown, glabrous and smooth, densely covered with lenticels which are round, oblong or linear, white or yellowish white; internodes are c. 2.4-4.6 cm. Two-year-old stems are dark-brown, glabrous and smooth, slender but hard; numerous small

Chapter 2 Species

l­enticels are rounded, oblong or lanceolate, grayish-white; pith is white, solid. Leaves  Membranaceous to chartaceous, nearly ovate, 7.110.7 cm long, 4.6-9 cm broad; base is truncate, bilaterally symmetrical; apices acuminate with short tips. Upper leaf surface green, glabrous, lustrous. The whole leaf or the apical part can be white or yellowish in early Spring, becoming greener with age. Midvein and secondary veins are

Figure 2.120  Stem, emerging shoot, and leaves of Actinidia polygama.

Figure 2.121  Pistillate and staminate flowers and fruit of Actinidia polygama.

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lighter green, depressed, glabrous or sometimes with sparse hairs. Leaf margin is serrated with little spines appressed or spreading, but not distinct, light purple-red. Lower leaf surface is light green, glabrous. Midvein and secondary veins are prominent, whitish-green, glabrous or sometimes with sparse hairs, five or six secondary veins on each side of midvein, reticulate toward leaf margin. Petiole, c. 2 mm in diameter, c. 2.5-3.3 cm long, is green, smooth, usually glabrous (Figure 2.120).

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Kiwifruit: The Genus ACTINIDIA

Pistillate flowers  Usually single, in leaf axils, rarely two to three flowers in an inflorescence. Pedicel is c. 5-15 mm long, sometimes pubescence on pedicel near sepal attachment. Calyx of five sepals, occasionally four or six, 5-7 mm long, 4.5 mm broad, nearly round with acute apex, green, glabrous, persistent. Flowers are c. 1.8-2.5 cm in diameter, five to six petals, 0.8-1.3 cm long, obovate, white, aging to yellowish white, There are c. 20 stamens, white filament, yellow anther but in plants from western China, anthers can be more brown. Ovary is bottle-shaped, yellow-green, glabrous; styles, c. 18-20, 4-5 mm long, white, stigma is white. The flowers are fragrant. Full blooming is in middle July. Staminate flowers Predominantly single, occasionally cyme inflorescence. Size, color and number of petals similar to those of pistillate flowers. There are c. 20-24 stamens in staminate flowers, yellow or brown anthers. Vestigial ovary and styles (Figure 2.121). cm long, 2.1  cm in diameter; fresh Fruit  Fruit, c. 3.4  weight c. 5-9 g, almost globose to ovoid to cylindric- or oblong-ovoid, shoulders are flat or round, stylar end pronounced, sharp beak. Fruit skin is green when unripe, becoming yellow when ripe, glabrous and smooth, with white powdery surface, no fruit spots. Calyx is persistent, five sepals, wider than the shoulders, oblong, greenish-white, and no pubescence on either surface. Fruit talk is green, glabrous and smooth, 2 mm in diameter, 1.8 mm long. Fruit flesh is smooth, pulpy not juicy, yellow when ripe, relatively large yellow core. Seeds dark-brown to black, smooth coat, flat, oblong. The fruit begin to ripen in late August. The unripe fruit have a very strong, peppery flavor, lost on ripening. VC content is 70-90 mg/100 g fresh weight, soluble solids 17.5% and total acidity 0.22-1.28%.

Diagnostic characters and species relationships The species is characterized by large solid pith and flowers with five sepals. Morphologically, Actinidia polygama is closely related to A. tetramera and A. valvata (Li, 1952; Liang, 1984; Li et al., 2007b,c), and this is supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. polygama in the National Germplasm Repository for Actinidia at WBG were 0.0828 and 0.01209, respectively, as revealed by AFLP markers (Li, 2006).

Distribution: Endemic to Guangxi, China.

Morphological characteristics Large deciduous climber. Stems Current branchlets brown velutinous, becoming glabrescent, lenticels subconspicuous; lenticels conspicuous on 2-year-old stems; pith white, laminate. Leaves  Leaf lamina chartaceous, 7-21 cm long, 4-11 cm broad, ovate-oblong to broadly ovate, base tends to cordate, apex acuminate, leaf margin is finely serrate. Upper leaf surface has fine downy hairs, soon lost to become glabrous. Lower leaf surface has stellate tomentum, partly lost to become glabrescent. On each side of midvein eight or nine secondary veins, reticulate veins prominent. Petiole is 2-4.5 cm long, velutinous, glabrescent. Flowers  Inflorescences are one to three-flowered, velutinous; peduncle 1-2 mm; pedicel 1.2-1.5 cm. Calyx has three to six sepals, ovate, 6-7 mm long, 3-5 mm broad, ovate. Corolla has five or six petals, 11-14 mm long, 8-11 mm wide, broadly ovate, red. Ovary globose, whitish tomentum. Flowers June. Fruit  Cylindrical, 1.5-3 cm long, 1.2-1.5 cm in diameter, tomentose; persistent sepals not reflexed. Fruit mature October-November.

Diagnostic characters and species relationships The species is distinct in the underneath of its leaves having sparse stellate tomentum, but readily shed, with prominent reticulate veins. Fruit are cylindrical with sparsely brown tomentum. Actinidia rongshuiensis is thought to be related to A. persicina and A. eriantha (Li, 2003).

2.35  Actinidia rubricaulis 红茎猕猴桃 hong jing mi hou tao

Actinidia rubricaulis Dunn in Bull. Misc. Inform. Kew 1906: 2. 1906. SYNTYPES: China. Yunnan: Yuanyang, A. Henry 10696 (A); Mengzi, A. Henry 11334 (A).

2.35a  Actinidia rubricaulis var. rubricaulis 红茎猕猴桃 (原变种) hong jing mi hou tao (yuan bian zhong)

Chromosomes: 2n = 2x = 58, 4x = 116

Distribution: China: Yunnan, Guangxi, Guizhou, Hunan, Chongqing, and Hubei (Figure 2.122).

2.34  Actinidia rongshuiensis

Morphological characteristics

融水猕猴桃 rongshui mi hou tao

Actinidia rongshuiensis R.G. Li & X.G. Wang in Guihaia 23(3): 196. 2003. TYPE: China. Guangxi: Guilin, September 2001, R.G. Li 0106 (Holotype, IBK).

Large, semi-evergreen climber. Stems Young shoots are brownish or grayish-green, covered with white pubescence, pubescence shed as stems

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Figure 2.122  Natural range of Actinidia rubricaulis.

age; lenticels are white, irregular shaped. Older stems are reddish-brown, with alternating pale green and brown stripes along the surface; pith is small, lamellate or solid, green. Leaves Thick chartaceous, long-ovate or nearly oblong, or broad lanceolate, c. 9-13 cm long, 4-6 cm broad. Base is cordate or round; apex is acuminate, occasionally acute. Leaf margin is densely serrated with small teeth appressed, purplish-red. Upper surface of leaf is dark green; basal part of midvein is reddish purple or pale green, with sparse white hispid hairs. Lower surface is much lighter green, covered with white pubescence. On each side

Figure 2.123  Stem and pistillate flowers of Actinidia rubricaulis.

of midvein, there are six or seven secondary veins. The petiole, c. 3-3.5 cm long, is purplish-red, covered with ­purple-red pubescence. Pistillate flowers  Single or in inflorescences of up to five flowers. Pedicels are glabrous, c. 1.5 cm long. Calyx has five sepals, c. 4 mm long, 3 mm broad, oblong, obtuse to round at apex, margins usually ciliate. Corolla has five petals, c. 7.5 mm long, c. 4.5 mm broad, long ovate, round at apex, white. Numerous stamens, filaments are c. 3.5 mm long, anthers, c. 1 mm long, ovoid, yellow. Numerous styles. Ovary is oblong-ovoid, c. 2.5 mm long, 1 mm in diameter, pubescent becoming glabrescent (Figure 2.123).

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Figure 2.124  Fruit of Actinidia rubricaulis.

Fruit  Fruit ovoid to oblong cylindrical, 1.5-2  cm long, 1.3 cm in diameter, average weight 1-3 g. Fruit skin is dark olive-green, abundant uniform-sized spots, y­ellowish-brown, ­ irregular shaped. Calyx of five sepals, persistent or shed later, brown, broad ovate to long lanceolate. Fruit talk is c. 1-12 mm long, pale green; mostly single fruit, or two or three fruits in a cluster. Fruit flesh is dark green; small core, pale green. Vc content is 16.8 mg/100 g fresh weight, 2.7% total sugars, 2.61% total acids. Fruit mature in September to October (Figure 2.124).

Diagnostic characters and species relationships The species is characterized by its slender 1-year-old reddish branches with dense small lenticels, inflorescence often one-flowered.

Figure 2.125  Natural range of Actinidia rubricaulis var. coriacea.

Chromosomes: 2n = 2x = 58, 4x = 116

2.35b  Actinidia rubricaulis var. coriacea 革叶猕猴桃 ge yi mi hou tao

Actinidia rubricaulis var. coriacea (Finet & Gagnepain) C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 224. 1984. Actinidia callosa Lindl. var. coriacea Finet & Gagnepain in Bull. Soc. Bot. France 52 (Mém. 4): 19. 1906. TYPE: China. Yunnan, M. Delavay 5152 (Isotype, K). Actinidia coriacea (Finet & Gagnepain) Dunn in J. Linn. Soc., Bot. 39: 405. 1911.

Chapter 2 Species

Distribution: China: Yunnan, Guangxi, Guangdong, Guizhou, Hunan, Jiangxi, Sichuan, Chongqing, Hubei, and Henan (Figure 2.125).

Morphological characteristics Large, semi-evergreen climber. Stems  One-year-old stems are slender, glabrous and smooth, green; abundant lenticels are distinct, yellow, usually linear but sometimes oblong; short internodes, c. 8-12 mm long. Young shoots are covered with purplish red-brown hairs; pith lamellate, greenish-white. Twoyear-old stems are relatively slender, but hard, brown, glabrous but rough surface, densely covered with lenticels which are usually linear, occasionally oblong or round,

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g­rayish-­ white and protruding from the surface; pith lamellate, brown. Leaves Thick leathery (coriaceous), c. 6-8 cm long, 3-5 cm broad, long-ovate to broad lanceolate; base is broad cuneate, bilaterally symmetrical, apex is acuminate or short acute. Upper surface of leaf is oily green, glabrous and smooth, lustrous. Midvein and secondary veins are glabrous. Apical part of leaf margin is wavy, densely serrated with small teeth having purple-red or brown tips, appressed, basal part leaves is entire. Lower leaf surface is pale yellowish green, glabrous, lustrous. Midvein is whitish-green, glabrous; secondary veins are pale green, glabrous; five to seven on each side of midvein. Petiole, c. 1.9-3 cm long, is purplish-red, glabrous, medium stout (Figure 2.126).

Figure 2.126  Stem and leaves of Actinidia rubricaulis var. coriacea.

Pistillate flowers  Pedicels are long, c. 21-26 cm, ­whitishgreen, glabrous. Calyx has five sepals, c. 6 mm long, 4 mm broad, elliptical, slightly curled inward like ladle, elliptical, green with pale purple-red pubescence, sparse on inner surface, but dense on outer surface. Flower c. 2 cm in diameter, five petals, c. 10 mm long, 6 mm broad, obovate to nearly elliptical, petal margin is slightly curled inward, narrow at base with light red radiating stripes extending to apex, the stripes usually having one or two branches, some are reticulate. Petals are often picotee, deep red at base, white toward margins. Numerous stamens, 23; filaments slender, c. 6 mm long, changing in color from white to purple-red. Anthers are yellow, kidney-shaped or oblong, c. 2 mm long, c. 1 mm broad, with deep, sometimes curving, longitudinal dehiscing grooves. Numerous styles, 24, white, c. 6 mm long, relatively slender, with slightly swollen, pale purple-red stigma. Ovary is nearly globose, about 3.5 mm in diameter, densely covered with pale purple-red pubescence, retained until fruitlets are set. Full blooming was observed on May 19, 1989 in Longling, Province.

Staminate flowers  Very floriferous, inflorescence a trichasium, averaging 22 flowers per inflorescence, usually five inflorescences on lower branching point of the trichasium; as many as 63 inflorescences on a 1.8 m long shoot. Primary peduncle is about 3-5 mm, covered with white pubescence. Primary pedicel of the apical flower is about 7-8 mm long, of other flowers c. 3 mm. Calyx has five sepals, long oblong, very pale red. Petals, usually five, pink, obovate, truncate at base, are pink to red, often with white margins. Numerous stamens, 25-35; filaments c. 5 mm long, anthers are yellow, slightly blackish yellow at apex. Ovary and styles vestigial. Full blooming was observed on May 16, 1989 in Malipo, Yunnan Province. Fruit Ovoid or globose, 2 cm long, 1.3 cm in diameter; average weight c. 1.8 g. Fruit skin is greenish-brown to darkbrown with yellow-brown spots on surface; glabrous when fruit are ripe. Fruit flesh is green, small core pale green, acid taste. Vc content is 24 mg/100g fresh weight, soluble solids is 13% and total acidity 0.57%. Fruit ripen mid-November, in Jiangkou, Taoshong, Guizhou Province (Figure 2.127).

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Figure 2.127  Pistillate and staminate flowers and fruit of Actinidia rubricaulis var. coriacea.

Diagnostic characters and species relationships

Morphological characteristics

It differs from A. rubricaulis var. rubricaulis in its deep red flowers and its coriaceous leaves and the ovary retaining white pubescence until young fruit are formed.

Medium-sized, deciduous climber.

As a variety of A. rubricaulis, it is naturally very closely related to A. rubricaulis var. rubricaulis. In addition, it ­appears to be related to A. fortunatii (Li, 1952). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. rubricaulis var. coriacea in the National Germplasm Repository for Actinidia at WBG were 0.0316 and 0.0461, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58

2.36  Actinidia rubus 昭通猕猴桃 zhao tong mi hou tao

Actinidia rubus H. Léveillé in Repert. Spec. Nov. Regni. Veg. 12: 282. 1913. TYPE: China. Yunnan: Zhaotong, June 1912, E.E. Maire s.n. (Holotype, A). Distribution: China: Yunnan and Sichuan (Figure 2.128).

Stems  One-year-old stems are green, covered with light brown, downy hairs; lenticels are white, round or oblong. Two-year-old stems are dark purplish brown, covered with reddish-brown, coarse hairs; lenticels are gray, oblong; internodes are c. 2.5-10 cm long; pith is lamellate, white. Leaves Thick chartaceous, broadly oblong-ovate to broadly oblong-obovate, 8.0-11.5 cm long, 6.5-8.5 cm broad, base shallowly cordate, symmetrical, apex acute. Upper leaf surface is green, almost glabrous, rarely strigillose; veins are pale green, purple toward base. Leaf margin is densely serrate, small teeth, shortly ciliate, appressed, green but with purplish brown tips. Lower surface is glabrous, pale green; veins are pale green, glabrous. On each side of midvein eight or nine secondary veins. Petiole is 2.5-7.0 cm long, pale purple, with sparse, pale brown, downy hairs. Pistillate flowers Single; peduncle is green, glabrous, 2.5 cm long. Calyx has five sepals, elliptical, 7 mm long and 5 mm broad, greenish-white, glabrous on outer surface. Flower 3 cm in diameter, five petals, obovate, 1.5 cm long, 1.0 cm

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Figure 2.128  Natural range of Actinidia rubus.

broad, yellowish white. Numerous stamens, c. 50; filaments white, c. 6 mm long; anthers yellow, sagittate. Styles, 36, white, 5 mm long, with white, slightly swollen stigmas. Ovary is flat cylindrical, 4 mm in diameter, with dense, white, downy hairs. Fruit Nearly globose; average weight c. 7-10 g; fruit apex flat, shoulders round. Fruit skin is green, glabrous; numerous lenticels are conspicuous, small, brown, round. Sepals are shed or can be persistent. Fruit talk is slender, yellowish-brown, 2.3 cm long, 2.1 cm in diameter. Fruit flesh is jade-green, core is white. Numerous seeds are small, purple, smooth coat, generally suborbicular or elliptical. The fruit are bland, juicy and soft texture. Vc content is 30 mg/100 g fresh weight, soluble solids content is 6.62% and acid content is 0.60%.

2.37  Actinidia rudis 糙叶猕猴桃 cao ye mi hou tao

Actinidia rudis Dunn in J. Linn. Soc., Bot. 39: 408. 1911. TYPE: China. Yunnan: Mengzi, A. Henry 11307 (Isotype,NY).

2.37a  Actinidia rudis var. rudis 糙叶猕猴桃(原变种)cao ye mi hou tao (yuan bian zhong)

Actinidia petelotii Diels in Notizbl. Bot. Gart. Berlin 11: 213.1931. TYPE: Vietnam. Chapa: A. petelot 3829 (Isotype, NY). Distribution: Endemic to Yunnan, China (Figure 2.129).

Diagnostic characters and species relationships

Morphological characteristics

The species is distinct in its leaves broadly oblong-ovate to broadly oblong-obovate with base truncate to cordatulate, leaf margin irregularly setose-serrulate, and branchlets densely reddish-brown strigose

Large deciduous climber.

Actinidia rubus is thought to be related to A. holotricha (Li, 1952; Liang, 1984).

Stems  One-year-old shoots are light green, densely covered with long, light purple hairs, denser and redder toward shoot apex. Two-year-old stems are greenish-brown, densely covered with long yellow or ginger-brown, stiff hairs; sparse lenticels are nearly elliptical, pale yellowish

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Figure 2.129  Natural range of Actinidia rudis var. rudis.

gray; internodes of 2-year-old stems are 6.0-6.5 cm long; pith is lamellate, white. Leaves  Chartaceous and mostly elliptical, 9.5-13.5  cm long, 5.5-7.5 cm broad; base nearly cordate or occasionally rounded; apex predominantly acute with long tip, 14-23 mm long. Serration on leaf margin is distinct, large spreading teeth alternating with one to three small teeth, appressed, purple-red ciliated. Upper leaf surface is green and densely covered with stiff, appressed purple-red hairs, relatively rough on the surface. Midvein is purple-red, even dark red toward base; six to eight secondary veins, light green, on each side of midvein; midvein and lateral veins are densely covered with stiff, appressed purple-red hairs, longer hairs on the midvein. The lower surface is light green, covered with white-downy hairs, more on the junctions of anastomosing veins. Petiole is c. 2-5.0 cm long, purple-red on exposed surface, and pale green on other surface, densely covered with purple-brown hispid hairs, relatively stout. Pistillate flowers  Cyme inflorescence, one to three flowers, predominantly three. Pedicel is c. 10 mm long, pale green, densely covered with purple-red and white

­ ubescence Calyx of five sepals, yellowish white, elliptical, p acute at apex, covered with purple and white pubescence. Flowers are c. 1.5 cm in diameter, five petals are gourdlike shape, white, c. 8 mm long, 6 mm broad, smooth margin, with parallel longitudinal stripes. Numerous stamens, 26, filaments white, c. 2 mm long, anthers yellow, sagittate. Styles white, 25 per flower, c. 4 mm long, with white, slightly swollen stigma. Ovary is nearly globose, c. 2.5 mm in diameter, green, and covered with dense white pubescence. Staminate flowers  Cyme inflorescence, two or three flowers, predominantly three. Calyx of five sepals, c. 8 mm long, white, light brown and purple pubescence especially toward apex. Flowers c. 1.5 cm in diameter, white, five petals are gourd-like or obovate, c. 7 mm long, 5 mm broad. Numerous stamens, c. 27; filaments white, c. 3 mm long; anthers are yellow, sagittate. Vestigial ovary, long cylindrical, yellowish green, covered with white pubescence. Fruit Small; long cylindrical; c. 2 cm long, 0.8 cm in diameter; average weight 1-5 g; shoulders rounded; apex often with

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small beak. Fruit skin is olive-green, tomentose when young, then glabrous; many lenticels, small, pale yellowish-brown, round or elliptical. Calyx is often persistent. Fruit talk is green and covered with brown downy hairs, more stout in basal part and slender in upper part, c. 1-2 cm long. Fruit flesh green; small, round core pale green. Numerous seed, elliptical. Fruit have a relatively strong acid flavor, numbing but not astringent, not juicy. VC content is c. 5 mg/100 g fresh weight, soluble solids content is 4.8% and acid content is 1.0% (Figure 2.130).

Diagnostic characters and species relationships It is distinct in that branchlets are densely strigose and leaves are densely strigose on both surfaces. Actinidia rudis is related to A. holotricha (Liang, 1984).

2.37b  Actinidia rudis var. glabricaulis 光茎猕猴桃 guang jing mi hou tao

Actinidia rudis var. glabricaulis C.Y. Wu in Fl. Yunnan. 1: 57. 1977. TYPE: China. Yunnan: Sichou, Nanchang, May 10, 1959, Q.A. Wu 8065 (Holotype, KUN). Distribution: Endemic to Yunnan, China (Figure 2.131).

Morphological characteristics Stems  One-year-old shoots are brownish-green, sparsely covered with pale purple downy hairs; lenticels are sparse, large, yellow, linear; internodes are 0.5-3.0 cm long. Twoyear-old stems are red, smooth, glabrous, medium diameter; lenticels are sparse, yellowish white, oblong or linear; pith is large, white. Leaves  Chartaceous, 6.2-11.0 cm long, 2.9-5.2 cm broad; base cuneate or nearly round, bilaterally symmetrical or asymmetrical, apex acute. Leaf margin is entire at base, above densely serrate with purple awns, appressed. Young leaves are covered with pale purple, downy hairs, which are readily shed with age. Lower surface of young, apical leaves is densely covered with white stellate hairs also readily shed with age. Upper surface of mature leaf is dark green, with sparse long white hairs, appressed. Midvein is concave, green, densely covered with pale purple, downy hairs or sparsely covered with long white hairs, appressed; secondary veins are green, sparsely covered with long white hairs, appressed. Lower leaf surface is green, glabrous; main and secondary veins are whitish-green, sparsely covered with short flosses; seven or eight secondary veins on each side of midvein, reticulately anastomosing at margin. Petiole is 2.03.7 cm long, purple, medium diameter, sparsely covered pale purple-red flosses, usually twisted at base.

Diagnostic characters and species relationships Figure 2.130  Stem, leaves, and fruit of Actinidia rudis var. rudis.

This variety is distinguished from A. rudis var. rudis by 1-year-old branches having brown hairs and 2-year-old

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Kiwifruit: The Genus ACTINIDIA

Figure 2.131  Natural distribution of Actinidia rudis var. glabricaulis.

branches being glabrous. Hairs on the lower leaf surface are restricted to the veins. As a variant of A. rudis, it also is related to A. holotricha (Liang, 1984).

2.38  Actinidia rufa 山梨猕猴桃 shan li mi hou tao

Actinidia rufa (Siebold & Zuccarini) Planchon ex Miquel in Ann. Mus. Bot. Lugduno-Batavi 3: 15. 1867. Trochostigma rufum Siebold & Zuccarini, Abh. Math.Phys. Cl. Königl. Bayer. Akad. Wiss. 3(2): 727. 1843 [“rufa”]. Actinidia arguta (Siebold & Zuccarini) Planchon ex Miquel var. rufa (Siebold & Zuccarini) Maximowicz in Bull. Acad. Imp. Sci. St. Petersb. 31: 1886. Actinidia callosa Lindley var. rufa (Siebold & Zuccarini) Makino in Bot. Mag (Tokyo) 16: 147. 1901. D i s t r i b u t i o n : Ta i w a n , C h i n a ; Ko r e a a n d Jap a n (Figure 2.132).

Morphological characteristics Large deciduous climber. Stems  Young shoots and floral branchlets have fine rustybrown pubescence. One-year-old shoots are ­grayish-green and glabrous. Two-year-old stems are brown; lenticels are small, elongated, conspicuous; pith is lamellate, greenish-brown. Leaves Chartaceous; 4-16 cm long, 3.5-9.5 cm broad; ovate to long ovate; leaf base rounded to truncate or shallowly cordate; apex obtuse to mucronate or long acuminate; margin shallowly mucronate-serrate, teeth produced by tips of veinlets, reddish, glandular. Both leaf surfaces glabrous, green; upper surface lustrous; lower surface initially has rusty-brown pubescence on veins, soon lost, tufts of hairs remaining in axils of veins. Upper surface has sunken veins; lower surface veins conspicuous, six or seven secondary veins on each side of midvein; reticulate veinlets conspicuous with subconspicuous parallel cross-bars. Petiole, 2-4 cm long, initially slightly pubescent, glabrous when mature (Figure 2.133).

Chapter 2 Species

Figure 2.132  Natural range of Actinidia rufa.

Figure 2.133  Stem and leaves of Actinidia rufa.

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Kiwifruit: The Genus ACTINIDIA

Flowers Cyme inflorescence;1-3 flowers in pistillate cymes, up to eight in staminate cymes. Calyx has five sepals, ovate, c. 5 mm long, both surfaces are covered with rusty-brown velutinous pubescence. Flowers are c. 1.5 cm in diameter, five petals, obovate, c. 10 mm long, white sometimes with pink-purple blush toward base of petals. Numerous stamens, 38 filaments, white, anthers yellow. Ovary is globose, c. 6 mm in diameter, dense brownish tomentum, c. 30 styles. In Wuhan, flowering is in May.

Fruit  Ovoid to oblong; 1.5-3 cm long, average weight 5-10 g. Stylar end often indented. Fruit skin is green; small, inconspicuous yellow-brown lenticels; very fine brown pubescence easily lost. Fruit flesh is green, small core, numerous black seed. Fruit ripen in \, acid, numbing and hot flavor, juicy. Vc content is 13.5 mg per 100 g fresh weight, soluble solids content is 7.5% and total acidity 0.6% (Figure 2.134).

Figure 2.134  Pistillate and staminate flowers and fruit of Actinidia rufa.

Diagnostic characters and species relationships It is characterized by fine brownish pubescence on floral branchlets bud readily shed, three to nine or more flowers in staminate inflorescences, single-flowered pistillate inflorescences, and fruits ovoid to oblong. Although morphologically quite distinct, Actinidia rufa was once thought to be related to A. arguta, and was even treated as a variety of A. arguta (Li, 1952). It is now recognized as a clearly distinct species (Ohwi, 1965; Walker, 1976; Li et al., 2007a,d). A close relationship between A. rufa and A. arguta is not supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The

Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. rufa in the National Germplasm Repository for Actinidia at WBG were 0.0789 and 0.1152, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58

2.39  Actinidia rufotricha 红毛猕猴桃 hong mao mi hou tao

Actinidia rufotricha C.Y. Wu in Fl. Yunnan. 1: 73. 1977. TYPE: China. Yunnan: Malipo, alt. 1400 m, May 13, 1964, S.Z. Wang 284 (Holotype, KUN).

Chapter 2 Species

2.39a  Actinidia rufotricha var. rufotricha 红毛猕猴桃 (原变种) hong mao mi hou tao (yuan bian zhong)

Distribution: Endemic to Yunnan, China (Figure 2.135).

Morphological characteristics Medium-sized, semi-deciduous climber. Stems  One-year-old shoots are densely covered with yellowish red villose hairs. Two-year-old stems are ­greenishbrown, covered with erect, relatively long, coarse, brown hairs. Lenticels are sparse, large, oblong; pith is lamellate, whitish-green.

141

Leaves  Thick chartaceous tending toward leathery, 14.217.8 cm long, 5.7-7.2 cm broad; broad lanceolate; base round or broad cuneate, predominantly symmetrical; apex acuminate, slightly oblique. The upper leaf surface is deep green, smooth, mostly glabrous. Midvein and secondary veins are greenish-white, conspicuous, concave, occasionally with bristles. The lower leaf surface is whitish-green, mostly with the midvein and secondary veins, 9-11 on each side of midvein, protruding and the mesh of veinlets is ­conspicuous. The surface is densely setose and stellate hairy with long cottony hairs mainly on the main veins but these are sparse elsewhere. Leaf margin is shallowly sinuate, sparse serrations ending as small red awns. Petiole, 3.2-4.6 cm long, medium diameter, is red, covered with coarse brown hairs, medium in diameter.

Figure 2.135  Natural range of Actinidia rufotricha var. rufotricha.

Flowers  Cyme inflorescences from nodes one to nine of 1-year-old shoots. Calyx has four sepals, ovate, c. 4 mm long, rusty-brown tomentum on outer surface. Flower has five petals, obovate, c. 5 mm long, apex rounded, pink or white. filaments slender, anthers yellow, ovoid, c. 2 mm long. Ovary in pistillate flowers is almost spherical, c. 1.5 mm in diameter, densely pubescent. Fruits  Ovoid, long cylindrical, 20-23  mm long and c. 10 mm in diameter. Skin olive-green, conspicuous lenticels,

round, yellow-brown. Sepals and stylar remains persistent. Fruit flesh is dark green, circular core whitish-green, many seed. Fruit talk is glabrous, 10-13 mm long. Fruit ripen in November (Figure 2.136).

Diagnostic characters and species relationships It is distinct in branchlets and petioles densely rufous setose, leaves oblong-lanceolate, underneath densely setose and stellate hairs along veins, and stellate hairs elsewhere.

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Figure 2.136  Stem, leaves, pistillate flowers, and fruit of Actinidia rufotricha var. rufotricha.

2.39b  Actinidia rufotricha var. glomerata 密花猕猴桃 mi hua mi hou tao

Actinidia rufotricha var. glomerata C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 319. 1984. TYPE: China. Guangxi: Lingle, May 21, 1960, C.F. Liang 32889 (Holotype, IBK). Distribution: China: Yunnan, Guizhou, and Guangxi (Figure 2.137).

Morphological characteristics Stems  One-year-old shoots are reddish-brown; glabrous; lenticels are conspicuous, white, oblong or linear. Twoyear-old stems are reddish-brown; glabrous; lenticels are conspicuous, grayish-white, protruding from the surface; internodes are 1.2-3.2 cm long; pith is lamellate, brown.

Figure 2.137  Natural range of Actinidia rufotricha var. glomerata.

Leaves  Chartaceous, nearly oblong, 8.5-11.5 cm long, 5.08.0 cm broad; leaf base is rounded, predominantly ­symmetrical; apex is mucronate. The upper leaf surface is dark green, smooth, glabrous, lustrous. Midvein and secondary veins are yellowish-brown, glabrous. Serrations on the margin are small, dense, ending in red, appressed awns. The lower leaf surface is light green; sparely cottony-­tomentum with short stellate hairs, readily shed; the midvein and secondary veins, 9-11 on each side of midvein, are yellowish green, covered with white or yellowish-brown, downy hairs which are readily shed. Petiole, 4-5 cm long, is purplish-red, glabrous (Figure 2.138). Pistillate flowers  Usually single. Calyx has four sepals, 3-4 mm long, ovate, with dense tomentum on outer surface. Flower is 4-5 mm in diameter, five or six petals, 5-7 mm long, ovate, bright pink to red, central part sometimes paler or white. Many stamens, filaments long, 2-4 mm, anthers

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Figure 2.138  Stem and leaf of Actinidia rufotricha var. glomerata.

yellow. Ovary is globose, 2-3 mm in diameter, densely covered with short tomentum. Flower stalk is 6-10 mm long. Fruit Small cylindrical fruit; c. 1.9 cm long, 1.2 cm in diameter; average weight c. 2-3 g; nearly truncate at apex with small, short beak; sepals deciduous. Fruit skin is green, glabrous, with small, round or oblong, yellowish-brown lenticels. Fruit stalk, c. 8 mm long, is green, densely covered with yellow downy hairs. Fruit flesh is green with small, cylindrical, whitish core. Seed are small, oblong, with smooth purple

coat. The fruit have an acid flavor, slightly sweet but with no astringency, juicy. Vc content is 150 mg per 100 g fresh weight, soluble solids content is 10.64% and total acidity 0.99% (Figure 2.139).

Diagnostic characters and species relationships A. rufotricha var. glomerata is distinguished from A. rufotricha var. rufotricha in its leaves having a thick cottony-­tomentum which is not easily observed as it is readily shed early.

Figure 2.139  Pistillate flowers and fruit of Actinidia rufotricha var. glomerata.

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As a variety of A. rufotricha, it is naturally very closely related to Actinidia rufotricha var. rufotricha. In addition, A. rufotricha var. glomerata is closely related to A. farinosa based on molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). Chromosomes: 2n = 2x = 58

2.40  Actinidia sabiifolia 清风藤猕猴桃 qing feng teng mi hou tao

Actinidia sabiifolia Dunn in J. Linn. Soc., Bot. 38: 357. 1908. TYPE: China. Fujian: Nanping, April-June 1905, Hongkong Herb. 2402 (Holotype, HK). Actinidia callosa Lindley var. sabiifolia Dunn in op. cit. 39: 406. 1911. Distribution: China: Hunan, Jiangxi, Fujian, and Anhui (Figure 2.140).

Morphological characteristics Small deciduous climber. Stems  One-year-old shoots are yellow-brown. Two-yearold stems are 3-5 mm in diameter, dark reddish-brown, lenticels are distinct and protruding from the surface. Pith small, brown, lamellate. Small flowering shoots are 3-7 cm

Figure 2.140  Natural range of Actinidia sabiifolia.

long; flowering shoots on vigorous water shoots are c. 25 cm long, 2.5-3 mm in diameter, glabrous; lenticels are distinct. Leaves Thin chartaceous, ovate to long ovate, 4-8 cm long, 3-4 cm broad; base round or obtuse, mostly bilaterally symmetrical; apex round to mucronate or retuse, but in younger leaves at the apex of vegetative shoots, leaf tips are mucronate to acuminate. Leaf margin is slightly serrated as dull and round teeth. Upper leaf surface is dark green, the lower surface is glaucous, much paler, gray green; glabrous on both surfaces; leaf veins are not protruding, very thin, five or six secondary veins on each side of midvein, slightly waved or nearly straight, sub-lateral veins hardly visible, small veins are dense and thin. Petiole, c. 2 mm long, is purplish pink, glabrous (Figure 2.141). Pistillate flowers Cyme inflorescence, two to four flowers; peduncle is c. 5 mm long, pedicel is c. 10 mm long. Bracts are lanceolate, c. 1 mm long. Calyx has five sepals, 2-3 mm long, obovate to oblong; glabrous and smooth except ciliate along the margins. Flowers are white, c. 8 mm in diameter, five petals are ovate to obovate, revolute, 5-6 mm long. Numerous stamens, 34-36; filaments are thread-like, c. 2 mm long; anthers are yellow, ovate. Ovary is globose, c. 2 mm in diameter, covered with reddish-­ brown pubescence, prominent styles c. 2 mm long. Flowering is in May.

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Figure 2.141  Stem, young shoot, and leaves of Actinidia sabiifolia.

Fruit Small, about 1.5-1.8 cm long and 1.0-1.2 cm in diameter, average weight 1-3 g; ovoid; sepals persistent. Skin of ripe fruit is dark olive-green, glabrous with small ­yellow-brown spots; fruit talk is light red, 8-10 mm long; Fruit flesh is

Figure 2.142  Fruit of Actinidia sabiifolia.

green, large core, numerous seeds are 2.5 mm long; Fruit have an acid flavor, not juicy. Vc content is 68 mg per 100 g fresh weight, soluble solids content is 12%, and acid content 1% (Figure 2.142).

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Kiwifruit: The Genus ACTINIDIA

Diagnostic characters and species relationships The species is distinct in that leaves on floral branchlets are often rounded at apex and spotted fruit are glabrous. Morphologically, Actinidia sabiifolia appears to be related to A. callosa and A. trichogyna (Li, 1952) but this is not supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). Chromosomes: 2n = 2x = 58

2.41  Actinidia sorbifolia

Leaves  Thick chartaceous, becoming thinly leathery; 9.514 cm long, 5.5-9.5 cm broad; ovate-oblong to elliptical, bilaterally symmetrical; base shallowly cordate or nearly round; apex round or mucronate. Upper leaf surface is green, glabrous, lustrous; leaf margin is inconspicuously denticulate, teeth small, green, spreading or ciliate; midvein and secondary veins are light green, basal part of main vein has sparse white tomentum. The lower surface is pale green, covered with white-downy stellate hairs; midvein and secondary veins are greenish-white, covered with white tomentum. Petiole, c. 2.5-3.0 cm long, pink to purplish-red (Figure 2.144).

Large deciduous climber.

Staminate flowers Cyme inflorescence, one to three flowers; pedicel c. 7 mm long, green, covered with pale brown tomentum; peduncle green, covered with white and pale brown tomentum. Calyx has five sepals, elliptical or ovate, 5 mm long, 4 mm broad, covered with downy brown hairs on both sides. Opening flowers are reddish or white, c. 2.5 cm in diameter, five or six petals, predominantly five, 14 mm long, 9 mm broad, overlapping, obovate, with longitudinal stripes. Numerous anthers, c. 56, filaments are white, anthers are yellow, sagittate. Vestigial ovary is small, cone-shaped, densely covered with pale brown pubescence.

Stems  One-year-old shoots are green, partly purplish-red, initially velutinous but becoming glabrous; lenticels are very distinct, sparse, round or elliptical, pale yellowish white; internodes 1-3 cm long. Two-year-old stems are purplish brown, glabrous; lenticels are sparse, round, gray-white; pith is brown, lamellate.

Fruit  Oblong to ovoid to almost globose; c. 3 cm long, 2.8 cm in diameter; average weight 13 g. Fruit skin is green with distinct fruit spots; densely covered with brown tomentum which is lost as fruit mature. Vc ­content is 41 mg per 100 g fresh weight, soluble solids content is 11% and total acidity is 1.76%.

花楸猕猴桃 ha qiu mi hou tao

Actinidia sorbifolia C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 324. 1984. TYPE: China. Guizhou: Yinjiang, in dense forest near road, alt. 1270 m, August 22, 1963, C.P. Tsien 31065 (Holotype, PE). Distribution: China: Yunnan and Hunan (Figure 2.143).

Morphological characteristics

Figure 2.143  Natural range of Actinidia sorbifolia.

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2.42  Actinidia stellatopilosa 星毛猕猴桃 xing mao mi hou tao

Actinidia stellatopilosa C.Y. Chang in Acta Sci. Nat. Univ. Szechuan. 1976(3): 75. 1976 [“stellatopilosa”]. TYPE:China, Chongqing: Chengkou, in shrubs, alt. 1200 m, June 7, 1958, T.L. Dai 105066 (Holotype, SZ; Isotype, PE). Distribution: Endemic to Chongqing, China. Endangered (Figure 2.145).

Morphological characteristics Large deciduous climber. Figure 2.144  Leaf of Actinidia sorbifolia.

Diagnostic characters and species relationships It is characterized by thick 1-year-old branchlets with very conspicuous lenticels; leaves thinly leathery, underneath sparsely stellate tomentose, and fruits globose or ovate. Actinidia sorbifolia is related to A. chinensis and A. obovata (Liang, 1984).

Figure 2.145  Natural range of Actinidia stellatopilosa.

Stems Flowering shoots are c. 30 cm or longer, 5 mm in diameter, initially sparsely covered with stiff, hispid hairs, later gradually shed; lenticels are visible. Two-year-old stems, c. 6.5 mm in diameter, still retain visible remnants of scurfy tomentum; lenticels are conspicuous. Pith is lamellate, white. Leaves  Chartaceous, 8-12 cm long, 4-8.5 cm broad; broadly ovate to broadly obovate; bilaterally asymmetrical; base subcordate; apex mucronate to acuminate to obtuse. Leaf margin has fine teeth. Upper leaf surface is green and glabrous except

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Kiwifruit: The Genus ACTINIDIA

sometimes for patches of downy hairs on midvein. Lower surface is glaucous, pale green, sparsely covered with white hairs, simple or branched and imperfect stellate hairs; hairs on the veins are coarser, yellow-brown; the veins are conspicuous, seven secondary veins on each side of midvein, lateral veins are visible, veinlets are minute. The petiole, 3-7 cm long, is pink, with sparse coarse downy hairs or short bristles. Staminate flowers Cyme inflorescence, two or three flowers. Peduncle is c. 3-5 mm long; pedicel is c. 8-12 mm long. Bract is diamond-shaped, 3-4 mm long, sparsely ­covered with ­ yellow-brown downy hairs. Calyx has five s­epals, 5-7 mm long, short ovate. Opening flowers are white, 1.8 cm in diameter; five or six petals, 10-12 mm long. Obovate. Numerous ­ stamens, filaments are 4-6 mm long, anthers are yellow, sagittate, 2.2 mm long. Vestigial ovary is small, round, covered with yellow-brown tomentum. Flowering during the first 10 days of June.

Diagnostic characters and species relationships It is characterized by papery leaves broadly ovate to obovate, unequal at base, obtuse to abruptly acute at apex, with imperfect stellate hairs underneath, and inflorescences with two or three flowers, flowers white. Actinidia stellatopilosa is thought to be related to A. chinensis var. deliciosa and A. trichogyna (Liang, 1984).

2.43  Actinidia styracifolia 安息香猕猴桃 an xi xiang mi hou tao

Actinidia styracifolia C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 321.1984. TYPE: China. Hunan: Zhijiang, May 28, 1959, P.C. Tam 60954 (Holotype, IBK). Distribution: China: Guizhou, Hunan, Jiangxi, and Fujian (Figure 2.146).

Figure 2.146  Natural range of Actinidia styracifolia.

Morphological characteristics Medium-sized, deciduous climber. Stems Flowering shoots are about 8-12 cm long and 2.5-3 cm in diameter, densely covered with dark, downy tomentum; lenticels are small, sparse, and inconspicuous. Two-year-old stems, c. 2.5 mm in diameter, are grayish brown, glabrous or sparsely gray-white scalp like tomentum; lenticels small and inconspicuous. Pith is lamellate, white. Leaves  Chartaceous; c. 6-9  cm long, 4.5-5  cm broad; ­elliptical-ovate or obovate; base broad cuneate, apex short acuminate to acute. The margin has hard sharp teeth. Upper leaf surface is green, sparsely covered with small, short, coarse strigose hairs, later shed. Lower leaf surface is grayish-green, densely covered with white stellate tomentum; midvein has light brown tomentum, usually seven secondary veins on

each side of midvein; lateral veins and veinlets are conspicuous. The petiole, c. 1.2-2 cm long, pale pink, is densely covered with short, dark-brown, velutinous tomentum (Figure 2.147). Pistillate flowers Cyme inflorescence, two to four flowers; peduncle is c. 4-8 mm long, and pedicel c. 5-7 mm long, both densely covered with brown velutinous pubescence. Bracts are diamond-shaped, c. 2.5-3.5 mm long. Calyx has two or three sepals, nearly round or ovate, 5 mm long, both ­surfaces densely covered with brownish velutinous ­ pubescence. Flowers are 1.3-1.5 cm in diameter, five petals, lanceolate or long obovate, 0.8-1.2 cm long, becoming reflexed, revolute, orange-reddish, often white at margins. Numerous stamens, filaments 4-6 mm long; anthers oblong, c. 1.5 mm long, yellow. Ovary globose, 3-4 mm in diameter, covered with dense, white or pale brown, velutinous pubescence. Styles are prominent, 6 mm long.

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Figure 2.147  Stem, emerging shoot, and leaves of Actinidia styracifolia.

Staminate flowers  Dichasium with five to seven flowers, covered with short tea-brown pubescence. Peduncle is c. 4-8 mm long and pedicel is c. 5-7 mm long. Bracts are ­diamond-shaped, c. 2.5-3.5 mm long. Calyx has two or three sepals, nearly round or ovate, c. 5 mm long, completely covered with dark-brown pubescence on both

sides. The flowers are c. 1.3 cm in diameter, five petals, long round, becoming reflexed, revolute, red, sometimes white at margins. Fruit  Mostly cylindrical; c. 3 cm long, 2 cm in diameter; average weight 2-4 g. Fruit skin is dark-yellow to brown,

Figure 2.148  Pistillate and staminate flowers and fruit of Actinidia styracifolia.

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Kiwifruit: The Genus ACTINIDIA

green spots, covered with dense, white, velutinous tomentum; sepals are persistent. Fruit stalk, c. 2.2 cm long, dark-yellow, sparsely covered with pale yellow pubescence. Fruit flesh is yellowish green with small, white core, numerous small seed. Fruit have an acid flavor, juicy. Vc content is c. 640 mg per 100 g fresh weight, soluble solids content is 9% and total acidity 1.1%. Fruit ripen in October (Figure 2.148).

Diagnostic characters and species relationships The species is distinct in its branchlets, petioles and inflorescences have brownish velutinous pubescence, and the cylindrical fruit which are densely tomentose. Liang (1984) thought that Actinidia styracifolia is related to A. lanceolata, but this is not supported by ­molecular

genetic data (Huang et  al., 2002a; Li et  al., 2002a; Chat et al., 2004; Li, 2006). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. styracifolia in the National Germplasm Repository for Actinidia at WBG were 0.0394 and 0.0576, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58

2.44  Actinidia suberifolia 栓叶猕猴桃 shuan ye mi hou tao

Actinidia suberifolia C.Y. Wu in Fl. Yunnan. 1: 73. 1977. TYPE:China. Yunnan: Pingbian, alt. 900 m, in shrubs, April 18, 1954, P.Y. Mao 3879 (Holotype, KUN). Distribution: Endemic to Yunnan, China (Figure 2.149).

Figure 2.149  Natural range of Actinidia suberifolia.

Morphological characteristics Large deciduous climber. Stems  One-year-old shoots are yellowish-brown, sparsely covered with fine brown pubescence; pith is hollow, green. Leaves  Thick and chartaceous; broad elliptical; c. 1014 cm long and 8-10 cm wide; base broad cuneate to rounded; short tip at apex. Margin of basal part of leaf

entire, apical part has small green teeth. The upper surface is green, smooth and slightly glossy, sparsely covered with brown hairs, appressed; leaf veins are brown, distinct, densely covered with brown pubescence. The lower surface is pale green, covered with white stellate hairs, or with white pubescence; veins light grayish brown, distinctly protruding; cinnamon-colored hairs on main veins. Petiole, c. 2.5 cm long, densely covered with rustybrown pubescence.

Chapter 2 Species

Pistillate flowers Cyme inflorescence with two or three flowers. Pedicel is c. 1 cm long, covered with y­ ellowish-brown pubescence. Calyx has five sepals, c. 6 mm long and 3 mm broad, elliptical, both surfaces covered with yellowish-brown pubescence. Flowers are c. 1.3 cm in diameter; five petals, c. 7 mm long, 4 mm broad, long obovate, apricot-yellow to orange. Numerous stamens, c. 60; filaments are 3 mm long;

151

anthers are yellow, ellipsoid. Ovary is globose, covered with yellowish-brown pubescence. Styles are yellow, usually 27, 3 mm long with slightly swollen stigma. Fruit Nearly globose, c. 1.5 cm in diameter; densely covered with tea-brown pubescence and fruits spotted early in development. Fruit flesh green, core pale greenish (Figure 2.150).

Figure 2.150  Stem, leaf, and fruit of Actinidia suberifolia.

Diagnostic characters and species relationships The species is distinct in that the lower leaf surface is densely tomentose with cinnamon-colored hairs on the veins but stellate hairs elsewhere, and the staminate inflorescences are racemose.

2.45  Actinidia tetramera 四萼猕猴桃 si e mi hou tao

Actinidia tetramera Maximowicz in Trudy Imp. S.Peterburgsk. Bot. Sada 11: 35. 1890. TYPE: China. Chongqing: Chengkou, R.P. Farges 530 (Isotype, NY). Clematoclethra giraldii Diels in Bot. Jahrb. Syst. 29(3-4): 472. 1900.

Actinidia tetramera var. badongensis C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 311. 1984. TYPE: China. Hubei: Badong, forest understory, alt. 2400 m, September 24, 1958, F.H. Chen 5083 (Holotype, HIB). Distribution: China: Yunnan, Guizhou, Sichuan, Chongqing, Hubei, Gansu, Shaanxi, and Henan (Figure 2.151).

Morphological characteristics Medium-sized, deciduous climber. Stems  One-year-old shoots are brownish-green, sparsely covered with long, pale, purplish-red downy hairs; lenticels

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Kiwifruit: The Genus ACTINIDIA

Figure 2.151  Natural range of Actinidia tetramera.

are relatively large, sparse, elliptical or linear, white; internodes 1.3-4 cm long. Two-year-old stems are grayish brown, glabrous; lenticels are small, elliptical and indistinct; pith is small, lamellate, brown. Leaves  Chartaceous; 4-8 cm long and 2-4 cm broad; ovate to round; bilateral symmetry or asymmetry; the base round or nearly flat; apex acute. The upper surface is dark green and sparely covered with whitish barbed hairs. Midvein is green, densely covered with white and pale purplish-red pubescence; secondary veins are green, sparsely covered with white pubescence. The leaf margin is sparsely serrated with relatively large teeth, brown, appressed. The lower surface is grayish-green, sparsely covered with white pubescence. Midvein and secondary veins are whitish-green, densely covered with long white pubescence. A characteristic feature is the tufts of white hairs at the axils of the mid- and lateral veins. In some genotypes, leaves may be variegated with white or pink blotches. The petiole, 4.3-6.5 cm long, is

purplish-red, sparsely covered with pale purplish-red pubescence (Figure 2.152). Pistillate flowers  Trichasium inflorescence with three to seven flowers. Pedicel is c. 1 cm long, pale red, covered with pale red tomentum. Calyx has four, sometimes three, sometimes five, sepals, c. 6 mm long and 10 mm broad; ovate; greenish-brown covered with dense purplish-red tomentum on both sides; margins ciliate. Flowers are c. 2 cm in diameter, yellowish white; mainly four or five but up to six or seven petals, c. 9 mm long, 5 mm broad; obovate or oblong, with radiating red stripes from base to apex. Numerous stamens, c. 40, filaments are white, 3 mm long, anthers are yellow, relatively small. Numerous styles, usually c. 27, 6 mm long with slightly swollen red stigma. Ovary is nearly round to cylindrical, 5 mm in diameter, glabrous. Staminate flowers Cyme inflorescence, one to three flowers, sometimes single flowered; calyx has four sepals, rarely

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Figure 2.152  Stem, young shoot, and leaves of Actinidia tetramera. Photographs provided by Zhangying.

Figure 2.153  Pistillate flowers and fruit of Actinidia tetramera. Photographs provided by Zhangying.

five, 4-5 mm long, 2-3 mm broad, oblong, apex acute; pale green but dark reddish-brown on outer ­surface. Opening flowers are c. 12 mm in diameter. Petals four, sometimes five, are oblong or gourd ladle-shaped, white to pink, pale green toward base. Numerous stamens, 40-47, anthers are yellow. Vestigial ovary is pale green, glabrous. Fruit Mostly oblong or ovoid; c. 1.5 cm long and 1.2 cm in diameter; average weight 1-3 g, without beak. Unripe fruit have pale green skin; smooth, glabrous; no lenticels; persistent sepals are reflexed. When ripe, fruit flesh becomes yellow to orange as does external appearance, medium-sized core. Numerous seed. The fruit is sweetly acid and juicy. VC content is 107 mg per 100 g fresh weight, soluble solids ­content 15.5% and total acidity 0.24%. The fruit ripen in August in Wuhan (Figure 2.153).

Diagnostic characters and species relationships The species is characterized by the lamellate, not solid pith, by the flowers often having four sepals and four petals, and the underneath of the leaves often being barbate (having tufts of hairs) at the axils of lateral veins.

Actinidia tetramera was thought to be related to A. kolomikta and A. polygama (Li, 1952; Liang, 1984), and a close relationship between A. tetramera and A. polygama, but not A. kolomikta, is supported by molecular genetic data (Li, 2006). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. tetramera in the National Germplasm Repositor y for Actinidia at WBG were 0.0079 and 0.0115, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58

2.46  Actinidia trichogyna 毛蕊猕猴桃 mao rui mi hou tao

Actinidia trichogyna Franchet in J. Bot. (Morot) 8: 278. 1894. TYPE: China. Chongqing: Chengkou, alt. 1400 m, R.P. Farges 370 (Holotype, P; Isotype: NY). Actinidia callosa Lindley var. trichogyna (Franchet) Finet & Gagnepain. in Bull. Soc. Bot. France 52 (Mém. 4): 20. 1906 (Contr. Fl. As. Or.).

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Kiwifruit: The Genus ACTINIDIA

Figure 2.154  Natural range of Actinidia trichogyna.

Distribution: China: Guizhou, Jiangxi, Sichuan, Chongqing, and Hubei (Figure 2.154).

Morphological characteristics Medium-sized, deciduous climber. Stems  Buds and young shoots are covered with rusty pubescence but soon become glabrescent; lenticels vary from indistinct to relatively distinct. Two-year-old stems are ­reddish-brown, 3-6 mm in diameter, glabrous; lenticels are distinct, whitish, oblong. Pith is lamellate, pale brown. Leaves  Chartaceous when young to soft leathery when older; obovate to oblong, 5-10 cm long, 3-6 cm broad; base obtuse to rounded or cordate, mostly bilaterally symmetrical; apex acute to acuminate. Leaf margin is serrated. Upper leaf surface is green; lower surface is glaucous, completely glabrous on both surfaces. Veins are not well developed; six or seven secondary veins on each side of the midvein, lateral veins are almost invisible, anastomosing veins are dense and thin. The petiole is pale red, 2.5-5 cm long, glabrous. Pistillate flowers  Axillary cymes have one to three flowers. Peduncle is short, 2-3 mm long, pedicel is 7-8 mm long; bracts are narrow, deltoid-shaped, c. 1.5 mm long. Calyx has five sepals, c. 5-6 mm long, oblong, margins of outer surface and whole of inner surface covered with short, grayish yellow pubescence. Flowers are c. 2 cm in diameter; five petals, obovate, c.

9-10 mm long. Numerous stamens; filaments are thread-like, c. 4-6 mm long; anthers are yellow, oblong, 2.5-3 mm long. Ovary is cylindrical or nearly globose, c. 3 mm long, sparsely covered with grayish yellow pubescence; styles are slightly shorter than ovary. Flowering is between late May and early July. Fruit  Predominantly single, rarely two or three fruit in a cluster; nearly spherical, ovoid to long globose to cylindrical; 1.5-3 cm long and c. 1 cm in diameter. Ripe fruit are dark green, glabrous with large brown spots. Seed are c. 2 mm long. Fruit ripen in October.

Diagnostic characters and species relationships It is distinct in its softly leathery leaves, which are glaucous underneath, and its spotted fruit. Actinidia trichogyna is thought to be related to A. venosa, A. pilosula, A. callosa, and A. glaucocallosa (Li, 1952; Liang, 1984).

2.47  Actinidia ulmifolia 榆叶猕猴桃 yu ye mi hou tao

Actinidia ulmifolia C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 314. 1984. TYPE:China. Sichuan: Pingshan, alt. 900 m, May 25, 1934, Science Institute of West China 5417 (Holotype, PE). Distribution: Endemic to Sichuan (Chongqing), China (Figure 2.155).

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155

Figure 2.155  Natural range of Actinidia ulmifolia.

Morphological characteristics

Diagnostic characters and species relationships

Medium-sized, deciduous climber.

The species is distinct in its leaf blade is obovate, the midvein and lateral veins are conspicuous underneath, the leaf margins are ciliate-serrulate, and the flowers have red petals, up to 14 mm long (herbarium specimens).

Stems  Current flowering shoots are soft and tender, c. 2-4 cm long, 2.5 cm in diameter, sparsely covered with pubescence; scattered lenticels are indistinct. Two-year-old stems are sparsely covered with remaining unshed hispid hairs; scattered lenticels are indistinct; pith is lamellate or solid, pale brown. Leaves Obovate, c. 6.5-8.5 cm long, 3.5-5.5 cm broad, base is obtuse, apex acute to round. Both leaf surfaces are glabrous except for a few soft hairs on veins. Leaf margin is serrate with tiny spines, ciliate. Leaf venation is well developed; veins on upper surface are slightly sunken, but protruding on lower surface. On each side of midvein, seven or eight secondary veins are straight, distinct; reticulate branching visible. Petiole, c. 2-2.5 cm long, is glabrous. Staminate flowers  Inflorescence with one to three flowers, peduncle is c. 3-4 mm long, pedicels are c. 10 mm long; bracts are diamond-shaped, c. 2 mm long, densely covered with short yellowish-brown, velutinous pubescence. Calyx has five sepals, 6-7 mm long, ovate to oblong-ovate, densely covered with yellowish-brown pubescence on both surfaces. Flower is c. 2 cm in diameter, five or six petals, c. 12-14 mm long, obovate, like shallow ladle, base very narrow, apex round; pale red. Numerous stamens, filaments are c. 6-8 mm long, anthers are ovate-sagittate, c. 2.5 mm long. Vestigial ovary is small, covered with yellowish-brown pubescence. Flowering is in late May.

2.48  Actinidia umbelloides 伞花猕猴桃 san hua mi hou tao

Actinidia umbelloides C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 313. 1984. TYPE: China. Yunnan: Jindong, alt. 2000 m, November 27, 1939, M.K. Li 2103 (PE).

2.48a  Actinidia umbelloides var. umbelloides 伞花猕猴桃 (原变种) san hua mi hou tao (yuan bian zhong)

Distribution: China: Yunnan, Sichuan, and Xizhang (Tibet) (Figure 2.156).

Morphological characteristics Large, deciduous climber. Stems One-year-old stems are green, glabrous and smooth; sparse lenticels are yellowish white, elliptical; internodes 11-40 mm long. Two-year-old stems are slender, glabrous and smooth; sparse lenticels are elliptical, brown; pith is relatively small, white, lamellate or solid. Older stems are ­reddish-brown, lighter on shaded side, glabrous.

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Kiwifruit: The Genus ACTINIDIA

Figure 2.156  Natural distribution of Actinidia umbelloides var. umbelloides.

Figure 2.157  Stem, leaves, and fruit of Actinidia umbelloides var. umbelloides.

Leaves Chartaceous; c. 6-14 cm long, 4-8 cm broad, almost oblong; base is broad cuneate, emarginate, bilaterally symmetrical; apex acuminate to obtuse. Upper leaf surface dark green, lustrous, glabrous. Primary vein and secondary veins are concave from the surface, green, glabrous. Leaf margin is densely serrated as small, but indistinct teeth, tooth tip is sharp and short, brown, appressed; margin of the basal part of some leaves entire. Lower leaf surface ash-green, glabrous; midvein and secondary veins conspicuous, whitish-­ green, vein axils sparsely covered with ash-like powder; 8-10 secondary veins on each side of midvein, anastomosing at the margin; some veins are branched at their base. Petiole is purplish-red, glabrous, c. 2.5-3.8 cm long.

Pistillate flowers Cyme inflorescences, three to five flowers per inflorescence. Peduncle is stout, c. 2 cm long, light purple-­red, covered with white pubescence; pedicels all of equal length, c. 10 mm, light purplish-red, sparsely covered with light purplish-­red pubescence. Calyx has six or seven sepals, c. 6 mm long, 4 mm broad, elliptical, green, glabrous on inner surface, purple-red pubescence on outer surface. Flowers are c. 15 mm in diameter; five petals, 7 mm long and 4 mm broad, broad elliptical, slightly broader at base, radiating stripes branching at margins, distinct on the surface, pink. Numerous stamens, c. 24, filaments are white, c. 3 mm long; anthers are yellow, sagittate, relatively small, longitudinal dehiscing grooves are not distinct. Numerous styles, c. 20, c. 4 mm long, milky-white, with slightly swollen stigma. Ovary is nearly

Chapter 2 Species

round, c. 3.5 mm long, 3 mm broad, covered with purplish-red pubescence. Flowering starts end of May in Yunlong, Yunnan. Fruit Ovoid, short ovoid, 1.5-2.2 cm long, apex round; fruit skin dark green with rusty-brown lenticels which are yellowish white at edges, relatively dense, protruding; sparse white pubescence. Calyces are persistent, five sepals, densely covered with short brown velutinous pubescence. Fruit stalk is about 1-1.9 cm long. Flesh dark green, core pale green. Seed are small. Fruit ripen in November (Figure 2.157).

Diagnostic characters and species relationships The species is characterized by its infructescence having three to five fruit, umbelliform, all pedicels of equal length, crowded on peduncle.

157

Actinidia umbelloides is thought to be related to A. fasciculoides and A. callosa var. henryi (Liang, 1984).

2.48b  Actinidia umbelloides var. flabellifolia 扇叶猕猴桃 shan ye mi hou tao

Actinidia umbelloides var. flabellifolia C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 314. 1984. TYPE: China. Yunnan: Fohai, alt. 1800 m, July 1936, C.W. Wang 77266 (Holotype, PE). Distribution: Endemic to Yunnan, China (Figure 2.158).

Figure 2.158  Natural range of Actinidia umbelloides var. flabellifolia.

Diagnostic characters and species relationships It is differs from A. umbelloides var. umbelloides in its leaf blade is suborbicular to obovate and is generally smaller, 6-7 cm long, 4-5.5 cm broad. As a variant of A. umbelloides, its relationships with other species is probably the same as those of A. umbelloides var. umbelloides.

2.49  Actinidia valvata 对萼猕猴桃 dui e mi hou tao

Actinidia valvata Dunn in J. Linn. Soc., Bot. 39: 404. 1911. TYPE: China. Jiangxi: Lushan, J.L. Bullock 121(Isotype, K). Actinidia valvata var. boehmeriaefolia C.F. Liang in Fl. Reipubl. Popularis Sin. 49(2): 311.1984. TYPE: China. Jiangxi: Lushan, Yadongping, alt. 1070 m,

158

Kiwifruit: The Genus ACTINIDIA

September 8, 1963,M.X. Nie & S.L. Chen 7927 (Holotype, PE). Actinidia valvata var. longipedicellata L.L. Yu in Guihaia 8(2): 132. 1988. TYPE: China. Zhejiang: Ningbo, L.L. Yu 8163 (Holotype, IBK).

Distribution: China: Guangdong, Hunan, Jiangxi, Fujian, Zhejiang, Hubei, Anhui, Henan, and Jiangsu (Figure 2.159).

Figure 2.159  Natural range of Actinidia valvata.

Medium-sized, deciduous climber.

g­ rayish-white pubescence, late glabrous, indistinct lenticels. Two-year-old stems are grayish-green, relatively distinct lenticels; pith is solid, white.

Stems  Current bearing canes are light green, 10-15 cm long, c. 2 cm in diameter; initially sparsely covered with

Leaves  Almost membranaceous, obovate to long-ovate, 5-13 cm long, 2.5-7.5 cm broad, base is broad cuneate to

Morphological characteristics

Figure 2.160  Stem, emerging shoot, leaves, and immature fruit of Actinidia valvata.

Chapter 2 Species

159

less than 1 cm long, sparsely covered with pubescence. Bracts are diamond-shaped, linear, minute, 1-2  mm long. Flowers are white, c. 2 cm in diameter. Calyx has two or three unequal sepals, concave, valvate, ovate to ­oblong-ovate, 6-9 mm long, 4-6 mm broad, apex acute, glabrous on both sides, or sometimes with pubescence on the middle part of outer surface. Corolla has seven to nine petals, c. 1-1.5 cm long, 1-1.2 cm broad, oblong to obovate, base attenuate, apex acute. Numerous stamens; filaments ­slender, c. 5 mm long; anthers are golden-yellow, cylindrical to round, 2.5-4 mm long. Ovary is bottle shaped, 5 mm long, green, smooth and glabrous. Styles form a circle of diameter slightly larger than ovary. Flowers in early May (Figure 2.160). Fruit Ovoid-globose; average weight 9 g; 3.1 cm long, 2.3 cm in diameter, rostrate, persistent calyx, mostly three sepals, reflexed. Skin smooth, glabrous, no lenticels. Fruit green when immature, orange when ripe. Fruit flesh also green when immature, orange when ripe. Strong “chili” flavor when immature. Vc content is c. 90 mg/100 g fresh weight, soluble solids 8.1%, total acid 0.31%; Seed are c. 1.8-3.5 mm long, 2.5 mm in diameter (Figure 2.161).

Diagnostic characters and species relationships It is characterized by stems having white, solid pith, and flowers with two or three sepals and five to nine petals. Morphologically, Actinidia valvata has been considered to be closely related to A. polygama and A. tetramera (Li, 1952; Li et al., 2007a,d), and this is supported by molecular genetic data (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). Chromosomes: 2n = 4x = 116, 6x = 174

2.50  Actinidia venosa 显脉猕猴桃 xian mai mi hou tao

Figure 2.161  Pistillate flowers and fruit of Actinidia valvata.

truncate-round, sometimes decurrent (spreading down petiole), can be bilaterally asymmetrical; apex is acuminate to rounded; margin is serrated with small teeth. Upper leaf surface is green, sometimes white variegated, lower surface paler; both surfaces glabrous except sometimes for pubescence in vein axils underneath. Midvein and five or six secondary veins on each side of midvein inconspicuous above, more prominent below. Petiole is pink, glabrous, about 1.52.0 cm long. Flowers  Inflorescences with two or three flowers, sometimes single. Peduncle is c. 1.5 cm long and pedicel is

Actinidia venosa Rehder in Sargent, Pl. Wilson. 2: 383. 1915. TYPE: China. Sichuan: Wenchuan, E.H. Wilson 1029 (Holotype, A). Actinidia venosa var. pubescens (H.L. Li) C.Y. Chang in Acta Sci. Nat. Univ. Sichuan. 1976(3): 76. 1976. A. venosa f. pubescens H.L. Li in J. Arnold Arbor. 33: 42. 1952. TYPE: China. Sichuan: Huili, alt. 2700 m, September 9, 1932, T.T. Yu 1451 (Holotype, A; Isotype, PE). Distribution: China: Yunnan, Guizhou, Hunan, Sichuan, Chongqing, and Xizang (Tibet) (Figure 2.162).

Morphological characteristics Large, deciduous climber. Stems One-year-old shoots are green, sparsely covered with appressed, long, white, downy hairs, older parts partially covered with white powder-like pubescence, soon glabrescent; lenticels are indistinct; internodes about 1.2-3.6 cm long.

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Kiwifruit: The Genus ACTINIDIA

Figure 2.162  Natural range of Actinidia venosa.

Two-year-old stems are reddish-brown, glabrous; smooth and lustrous; lenticels are relatively large, dense, elliptical, yellow-brown; pith is large, lamellate, greenish-white.

Leaves Chartaceous; 5-15 cm long, 3-8 cm broad; oblong or elliptical; base round or broad cuneate, usually oblique; apex acute, occasionally acuminate. The upper surface is dark green and glabrous, or with sparse hairs when young. Leaf margin is serrate with tiny spine-like teeth, purplish-red, appressed. Midvein and secondary veins are conspicuous, pale green, glabrous. Lower leaf surface is paler green but not glaucous, glabrous except for some pale brown hairs on veins, hairs simple or branching, arachnoid (long and delicate); main and secondary veins are very distinct, raised, greenish-white; 8-11 secondary veins on each side of midvein; very prominent paralleled. Petiole, c. 1.5-4 cm long, purplish-red, glabrous. Pistillate flowers Usually single. Pedicel is about 7 mm long, purple-red and glabrous. Calyx has five sepals, c. 5 mm long and 2 mm broad, purplish-red or green, elliptical. Flowers are c. 2 cm in diameter; five petals, oblong, c. 10 mm long, 5 mm broad, inwardly curled, white. Numerous stamens, c. 25, filaments are white, 2 mm long; anthers are yellow, sagittate. Ovary is green, short cylindrical, c. 3 mm in diameter,

Figure 2.163  Stems, leaves, and fruit of Actinidia venosa.

Chapter 2 Species

glabrous; styles are white, c. 17, 3 mm long with slightly swollen stigma. Staminate flowers Up to seven flowers in inflorescence; peduncle 0.5-1.5 cm, pedicels c. 0.9 cm long, pale purplish-red, densely covered with pale purplish-red pubescence. Calyx has five sepals, brownish-green, elliptical, c. 5 mm long, 2 mm broad, densely covered with pale purple-red pubescence on both sides. Corolla is c. 2 cm in diameter, five petals, yellow-white, obovate, about 10 mm long and 4 mm broad, base is red with radiating longitudinal red stripes. Filaments, c. 21, are white, c. 5 mm long, anthers are yellow, sagittate. Vestigial ovary is small, cone-shaped, densely covered with purplish-red pubescence. Fruit Small; average weight 5 g; c. 2 cm long. 1.8 cm in diameter; nearly ovoid or short cylindrical; shoulders round, usually bilaterally symmetrical; apex is truncate to emarginate; reflexed sepals and styles persistent. Fruit skin is green, sparsely covered with short, pale brown tomentum and white powder, easily shed, glabrous when mature; fruit spots are dense, irregularly distributed, yellow-brown, round or elliptical, small. Fruit stalk, c. 7 mm long, is greenish-brown,

Figure 2.164  Natural range of Actinidia vitifolia.

161

covered with brown tomentum. Fruit flesh is green, core pale yellow. Relatively numerous seed, large, elliptical, ­purple-red. Fruit ripen in September, acid flavor, lightly astringent, not juicy, soft texture when ripe (Figure 2.163).

Diagnostic characters and species relationships The species is characterized by midvein and secondary veins being conspicuous on the lower leaf surface and having white arachnoid pubescence. Actinidia venosa is thought to be related to A. trichogyna, A. pilosula, and A. callosa (Li, 1952).

2.51  Actinidia vitifolia 葡萄叶猕猴桃 pu tao ye mi hou tao

Actinidia vitifolia C.Y. Wu in Fl. Yunnan. 1: 59. 1977. TYPE: China. Sichuan: Leibo, Xi’ning Town, Hetaoping, alt. 1600 m, June 8, 1959, Sichuan Eco. Pl. Exped. 520 (Holotype, KUN). Distribution: China: Yunnan and Sichuan (Figure 2.164).

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Kiwifruit: The Genus ACTINIDIA

Morphological characteristics Large, deciduous climber. Stems  One-year-old stems are grayish-green, sparsely covered with coarse brown hairs; lenticels are conspicuous, white or pale brown, oblong or linear; internodes are c. 8-11 cm long. Two-year-old stems are purplish brown, glabrous, lenticels are conspicuous, brown, oblong or linear, protruding from surface; pith is lamellate, white to brown. Leaves  Thick chartaceous, basal leaves on a shoot are obovate, upper leaves are ovate, 5-14 cm long, 4.5-9.5 cm broad; base truncate or cordate, usually symmetrical; apex mucronate or truncate. The upper leaf surface is green, ­glabrous, sparsely setose along the vein. The bases of leaves at the lower part of a shoot are entire, the bases of leaves further up a shoot are sparsely serrate or entire. Serrations are wavy, frequent with one or two smaller denticulations with appressed pale purple awns alternating with larger serrations with appressed brown awns. The lower leaf surface is green, glabrous apart from short, pale purple downy hairs on midvein and secondary veins. Veins are parallel, protruding from the surface, conspicuous. The secondary veins, 6-10 on each side of midvein, anastomose reticulately at the leaf margin. Petiole is 3.5-6 cm long, stout, purplish, densely covered with white and pale purple short soft hairs.

Figure 2.165  Stem, leaves, pistillate flowers, and fruit of Actinidia vitifolia.

Pistillate flowers Cymose inflorescences with up to five flowers. Peduncle c. 8 mm, pedicels up to 3 cm long, tomentose to strigose. Calyx has four to six sepals, ovate, 4-8 mm long, both inner and outer surfaces tomentose. Flower is white or pink, 3 cm in diameter; four to eight petals are spoon-like, obovate, 13-17 mm long, apex rounded. Numerous stamens, filaments are 6-8 mm long, anthers are yellow, sagittate, c. 2 mm long. Ovary is 5 mm in diameter, almost globose to cylindrical, densely covered with ­yellowish-brown ­pubescence, styles are 7-8 mm long. Fruit Globose to short cylindrical to ovoid to obovoid; 3.24.4 cm long and 2.8-3.8 cm in diameter; weight 21-35 g, truncate at apex, shoulders round. Sepals persistent. Fruit skin is green, glabrous with yellow-brownish lenticels. Fruit stalk, 1.5 cm long, is green, glabrous. Fruit flesh is green, relatively large, white core. Seed are small and oblong. Fruit have an acid taste and are astringent (Figure 2.165).

Diagnostic characters and species relationships The species is characterized by branchlets and petioles being sparsely strigose, the pedicel and peduncles of the inflorescence with one to five flowers being tomentose to strigose, and fruits being globose with brownish lenticels. Actinidia vitifolia is morphologically distinct and there are no obviously similar species: A. callosa is the most similar on the basis of its leaves and veins (Liang, 1984).

Chapter 2 Species

2.52  Actinidia zhejiangensis 浙江猕猴桃 zhe jiang mi hou tao

Actinidia zhejiangensis C.F. Liang in Guihaia 2(1): 2. 1982. TYPE: China. Zhejiang: Qingyuan, October 1981, C.F. Gan 3 (Holotype, IBK).

163

Actinidia persicina R.H. Huang & S.M. Wang in J. Wuhan Bot. Res. 13(2): 113. 1995. TYPE: China. Hubei: Wuhan, September 17, 1991, R.H. Huang 9113 (Holotype, HIB). Distribution: China: Jiangxi, Fujian, and Zhejiang (Figure 2.166).

Figure 2.166  Natural range of Actinidia zhejiangensis.

Morphological characteristics Large, deciduous or partially evergreen climber. Stems Flowering shoots are greenish or yellowish-brown, 10-25 cm long, 4-5 mm in diameter; initially sparsely covered with tomentum, but shed later. Vegetative shoots are very long, c. 6 mm in diameter, with short and slender branches, densely covered with persistent y­ ellowish-brown tomentum; pith is lamellate, white or brown. Leaves  Chartaceous; ovate, oblong or long ovate; 5-20 cm long, 2.5-11 cm broad; base subcordate to auriculate, apex attenuate or acuminate. The upper surfaces of leaves from flowering shoots are glabrous, the main veins on leaves from vegetative shoots may have ­remnants of stiff hairs. The lower leaf surface is glaucous and initially may have a dense brownish tomentum of imperfectly stellate hairs which at maturity are largely or completely lost. The veinlets on the lower surface have a silvery tomentum of unbranched hairs

which are readily shed when the leaf is mature. There are usually seven secondary veins on each side of the midvein. These may be sunken below on the upper surface but conspicuously protrude from the lower surface. The petiole, c. 1-4 cm long, of leaves of flowering shoots is glabrous, on leaves from vegetative shoots it is covered with ­yellowish-brown tomentum (Figure 2.167). Flowers Inflorescences may have one or two branches or a single flower but more usually three to seven flowers; peduncles are 4-10 mm long, pedicels are 0.6-1.6 cm long, both covered with yellowish-brown tomentum; bracts are slender and taper to a fine point, 3-6 mm long. Calyx has five or six sepals, 6-8 mm long, ovate to narrowly ovate, both surfaces with a dense brown tomentum. Corolla has five or six petals, c. 10-16 mm long, obovate to narrowly obovate, intense rose pink. Numerous stamens, filaments are 4-8 mm long; anthers are brownish yellow, ovoid, c. 1 mm in diameter, sagittate at base. Ovary is globose, c. 6-8 mm in diameter, densely covered with grayish pubescence; styles are c. 5 mm long.

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Kiwifruit: The Genus ACTINIDIA

Figure 2.167  Stem and leaves of Actinidia zhejiangensis.

Fruit Short cylindrical, to ovoid to almost globose, 3.5-4 cm long, 3 cm in diameter, average weight 20 g, almost truncate at apex and base. Sepals are persistent, reflexed. The fruit skin is greenish yellow, covered with short, scurfy, downy hairs and more obvious, long, white to yellowish-brown hairs. The fruit stalk, c. 3.5 cm long, is covered with yellowish-­brown tomentum. Fruit flesh is green, relatively few seed. Fruit are juicy, with a good balance of sweetness

Figure 2.168  Pistillate flowers and fruit of Actinidia zhejiangensis.

and acidity, Vc content is 290-370 mg per 100 g fresh weight, soluble solids content is 10-12% and total acidity1.5-1.7%. Fruit ripen in October (Figure 2.168).

Diagnostic characters and species relationships The species is distinct in that its leaves have sparse brownish tomentum with imperfectly stellate hairs,

Chapter 2 Species

and the cylindrical-oblong fruit have dense, long, ­yellowish-brown tomentum. Actinidia zhejiangensis is thought to be related to A. eriantha (Liang, 1984), and this is supported by molecular genetic data in the maternal lineage (Li, 2006). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. zhejiangensis in the National Germplasm Repository for Actinidia at WBG were 0.0631 and 0.0921, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58

165

2.53  Actinidia hypoleuca 白背叶猕猴桃 bai bei ye mi hou tao

Actinidia hypoleuca Nakai in Bot. Mag. Tokyo 38: 312. 1904. TYPE: not designated. Actinidia melanandra sensu Fin. & Gagn. in Bull. Soc. Bot. France 52, Mém. 4: 21. 1905. p.p. (quoted Japanese plants), non Franch. Actinidia japonica Nakai in Bot. Mag. (Tokyo) 28: 311. 1914. TYPE: not designated. Distribution: Japan (endemic) (Figure 2.169).

Figure 2.169  Natural range of Actinidia hypoleuca.

Morphological characteristics Deciduous climber. Stems Branches grayish, young branchlets darker, glabrous, lenticels absent or inconspicuous; pith is lamellate or occasionally nearly solid, brown.

Leaves Chartaceous, long ovate or ovate, 2.5-7 cm long, 2.54.5 cm broad; base round, truncate or broad truncate; apex acute or acuminate; margin has small teeth. Upper leaf surface is dark green, glabrous, but underneath is glaucous with white powder-like surface; glabrous except for tufts of brown pubescence in vein axils. Midvein has four or five secondary veins on

166

Kiwifruit: The Genus ACTINIDIA

each side, inconspicuous above, more obvious below. Petiole is 2-3.5 cm long, glabrous, reddish purple (Figure 2.170). Flowers Axillary inflorescence has one to five flowers, glabrous or with pubescence, peduncle is 5 mm and pedicel is 10 mm long. Calyx has five sepals, long ovate to long oblong, c. 6 mm long, 3 mm broad, glabrescent or with very fine pubescence, margins often ciliate, r­eddish-brown. Flower is

Figure 2.170  Stem and leaves of Actinida hypoleuca.

Figure 2.171  Staminate and pistillate flowers and fruit of Actinida hypoleuca.

ivory white, five petals, 12 mm long, 5 mm broad, long ovate, apex rounded. Numerous stamens; slender filaments c. 3 mm long; anthers are p ­ urple-black, oblong, c. 2 mm long. Ovary is bottle-shaped, 5 mm long and 2 mm in diameter, glabrous, pale green; styles are c. 2-4 mm long. Fruit  Fruit are ovoid, c. 1.5 cm in diameter, glabrous, no spots, slightly rostrate at stylar end (Figure 2.171).

Chapter 2 Species

Diagnostic characters and species relationships The species is distinct in its papery leaves which are glaucous underneath and its ovoid spotless fruit. Morphologically, Actinida hypoleuca appears to be closely related to A. arguta and A. melanandra (Li, 1952; Li et al., 2007a,d), and this is supported by molecular genetic data in the maternal lineage (Huang et al., 2002a; Li et al., 2002a; Chat et al., 2004; Li, 2006). The Nei’s genetic diversity and the Shannon’s polymorphic information index of living collection A. hypoleuca in the National Germplasm Repository for Actinidia at WBG were 0.0907 and 0.1325, respectively, as revealed by AFLP markers (Li, 2006). Chromosomes: 2n = 2x = 58

2.54  Actinidia strigosa 尼泊尔猕猴桃 ni bo er mi hou tao

Actinidia strigosa Hooker f. and Thomas in J. Linn. Soc., Bot. 5: 55. 1861. TYPE: Sikkim, J.D. Hooker s.n. (Holotype, K; Isotype, GH).

167

Leaves  Chartaceous, 7-13 cm long, 4-7 cm broad, ovate to long ovate; base obtuse to rounded, often oblique; apex acuminate to long acuminate; margins have small serrations. Upper leaf surface dark green, lustrous, glabrous or sparsely pubescent along veins; lower surface, lighter green, glabrous or with sparse setose hairs along veins; five to seven secondary veins on each side of midvein, prominent. Petioles are 2.5-3 cm long with short, stiff, appressed hairs or rusty-brown pubescence. Flowers  Axillary inflorescences have two to four flowers; peduncle is up to 1 cm long, pedicels 5-10 mm, both with rusty-brown pubescence; bracts small, lanceolate. Calyx has five sepals, ovate, c. 4-5 mm long and 3-4 mm wide, almost glabrous or sparsely tomentose on outer surface. Flowers white, five petals, obovate, 8 mm long and 5 mm broad, rounded at apex. Numerous stamens; filaments 2.3 mm long; anthers yellow, 1-1.5 mm long. Ovary almost globose, 1.5 mm in diameter, with dense, long, shaggy hairs, styles c. 1.5 mm long.

Distribution: Nepal (endemic)

Fruit Ovoid, 3 cm long.

Morphological characteristics

Diagnostic characters and species relationships

Deciduous climber. Stems  New flowering shoots are reddish-brown with dense hispid hairs; rough surface with only occasional, elongated lenticels. Two-year-old stems have some residual pubescence and lenticels more conspicuous; pith is large, whitish, lamellate.

The species is characterized by the strigose branchlets, and the concolored leaves with rounded base and sparse setose hairs along the veins of the lower surface. There is no literature concerning the taxa which might be related to Actinidia strigosa, but Li (1952) compared it with A. petelotii, which has now been merged into A. rudis (Li et al., 2007a,d).