Solanum lyratum Thunb., 1784

Knapp, Sandra, 2013, A revision of the Dulcamaroid Clade of Solanum L. (Solanaceae), PhytoKeys 22, pp. 1-432 : 163-170

24. Solanum lyratum Thunb. , Fl. Jap. 92. 1784 Figure 61

Solanum lyratum Thunb. ex Murray, Syst. Veg., ed. 14: 224. 1784. Type: Based on Solanum lyratum Thunb.

Solanum dichotomum Lour., Fl. Cochinch. 129. 1790. Type: China. "Cantone Sinarum" (no specimens cited).

Solanum dulcamara L. var. pubescens Blume, Bijdr. 698. 1825. Type: Based on Solanum lyratum Thunb.

Solanum dulcamara L. var. chinense Dunal, Prodr. [A.P. de Candolle] 13(1): 79. 1852. Type: China. "Kianang Prov.", L. Macartney s.n. (lectotype, designated here: BM [BM000942424]).

Solanum dulcamara L. var. lyratum (Thunb.) Seibold & Zucc., Observ. Bot. 4: 317. 1915. Type: Based on Solanum lyratum Thunb.

Solanum septemlobum Bunge var. indutum Hand.-Mazz., Oesterr. Bot. Z. 83: 234. 1934. Type: China. S-Schansi [Shanxi], Sunnandschen, 23 Jul 1916, E. Licent 2286 (holotype: W [W-1933/0006499]; isotype: K [K000658180], BM [BM000942501]).

Solanum lyratum Thunb. var. filamentaceum Hayashi, J. Geobot. 22: 4. 1974. Type: Japan. Hondo: prov. Musashi, Mt. Takao, 23 Sep 1973, R. Mineo s.n. (holotype: TNS [originally cited from TUAT], not located).

Solanum cathayanum C.Y.Wu & S.C.Huang, Fl. Reipubl. Popul. Sin. 67(1): 84. 1978. Type: Based on Solanum dulcamara L. var. chinense Dunal

Solanum kayamae T.Yamaz., J. Jap. Bot. 68: 339. 1993. Type: Japan. Ryuku: Okinawa Island, Mabuni, 1 Dec 1992, K. Kayama 6658 (holotype: TI [n.v.]; isotype: BM [BM000647002]).

Solanum lyratum Thunb. forma purpuratum Konta & Katsuy., Bull. Natl. Sci. Mus., Tokyo, B 31(1): 23. 2005. Type: Japan. Honshu: Shizuoka, Suzaki, Shimoda City, Suzaki Peninsula, ca. 40 m, 2 Sep 2003, F. Konta 23381-a (holotype: TNS [photograph]; isotype: TNS [n.v.]).

Type.

Japan. Nagasaki, C. Thunberg s.n. (holotype: UPS).

Description.

Sprawling shrub or herbaceous vine, woody at the base. Stems flexuous, not winged, sparsely to densely pubescent with translucent, glandular, simple uniseriate trichomes to 4 mm long, with 4-6 cells, the gland 1-celled, these overtopping shorter glandular simple trichomes ca. 0.5 mm long, the trichomes weak and tangled; new growth densely pubescent with trichomes like those of the stems. Bark of older stems pale yellowish tan, glabrescent, the longer trichomes usually breaking off and only the shorter ones remaining. Sympodial units plurifoliate. Leaves simple or pinnatifid, usually only with 2 lyrate lobes at the base, 2-6(-9) cm long, 0.5-5(-7) cm wide, cordate to lyrate, widest in the basal third, thin and membranous, both surfaces uniformly pubescent with weak, translucent simple uniseriate trichomes to 4 mm long, these usually glandular like those of the stems; primary veins 5-7 pairs, usually yellowish in dry material; base cordate, occasionally truncate, not decurrent on the petiole; margins entire or lobed, the lobes usually only a pair at the base, these sometimes completely divided and the leaf apparently pinnate, occasionally with up to 4 pairs of lobes; apex acuminate or acute; petiole 1-3 cm long, pubescent like the stems, twining. Inflorescences terminal or lateral, 2.5-10 cm long, open and many times branched, with 10-40+ flowers, usually only a few open at a time, pubescent with simple uniseriate glandular trichomes like those of the stems; peduncle 2-6 cm long; pedicels 7-11 mm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, slender and spreading, glabrous or short-glandular pubescent, articulated at the base in a small sleeve 1-2 mm long; pedicel scars irregularly spaced 2-9 mm apart, more congested distally. Buds ellipsoid, the corolla strongly exserted from the calyx tube before anthesis. Flowers all perfect, 5-merous. Calyx tube 1-1.5 mm long, conical, the lobes 0-1 mm long, deltate or mere enations on the calyx rim, glabrous, papillate on the tips. Corolla 8-13 mm in diameter, white to pale lavender, with green and white spots at the base of each lobe, stellate, lobed 3/4 of the way to the base, the lobes 3-6 mm long, 2-3 mm wide, strongly reflexed at anthesis, the tips and margins densely papillose. Filament tube minute, the free portion of the filaments 1-1.5 mm long, glabrous; anthers 3-3.5 mm long, 1-1.5 mm wide, ellipsoid, loosely connivent, often dark blue or black, the base sagittate, poricidal at the tips, the pores lengthening to slits with age. Ovary glabrous; style 5-7 mm long, glabrous; stigma capitate, the surface minutely papillate. Fruit a globose berry, ca. 1 cm in diameter, bright red, translucent and shiny when ripe, the juice staining scarlet; fruiting pedicels 1-1.5 cm long, ca. 0.75 mm in diameter at the base, spreading. Seeds & gt;30 per berry, ca. 2.5 mm long, ca. 1.5 mm wide, flattened reniform, pale yellowish tan, the surfaces minutely pitted, when mature the seed apparently hairy from the elongate lateral testal cell walls, these to 0.5 mm. Chromosome number: not known.

Distribution

( Figure 62 ). Solanum lyratum occurs in a wide variety of habitats in China, Japan and north Vietnam, from sea level to 1500 m.

Ecology.

Usually growing in secondary situations along roadsides and in waste ground; often in urban areas.

Common names:

China: bai ying ( Zhang et al. 1994); Japan: murasaki-hiyodorijogo ( Konta et al. 2007).

Conservation status.

Least Concern (LC); EOO>100,000 km2 (LC) and AOO>10,000 km2 (LC). See Moat (2007) for explanation of measurements.

Discussion.

Like the other temperate zone species in the Dulcamaroid clade ( Solanum dulcamara , Solanum pittosporifolium ), Solanum lyratum is widely distributed, somewhat weedy and grows in many different habitats. It is sympatric with and very similar to Solanum pittosporifolium ; the species can be differentiated by leaf pubescence. Solanum lyratum always has long glandular trichomes on all vegetative parts, especially the new growth, while Solanum pittosporifolium is glabrous or has very short uniseriate trichomes on the leaves and young stems. Leaf shape in Solanum lyratum is extremely variable; most specimens have lyrate leaf bases and ternate leaves with a very large central lobe (see Figure 2 ), but simple leaves are common and some specimens (e.g., Licent 2286, the type of Solanum septemlobum var. indutum) have pinnatifid leaves with 5-7 pairs of leaflets. Pubescence, however, is very consistent and is an excellent diagnostic character for Solanum lyratum . Flower color also varies from dark purple to white, but there are always green spots at the base of the corolla lobes. Although most Solanum species have bright yellow anthers, Solanum lyratum occasionally has blackish purple anthers; it is not clear whether this is genetic or is due to an environmental effect. Anthers in Solanum lyratum are never tapered like those of Solanum dulcamara ; due to pubescence differences these two species are not difficult to distinguish (but see Solanum pittosporifolium ).

Two specimens at "herb. Banks" (Macartney s.n. and Staunton s.n. both now at BM) were cited by Dunal in his description of Solanum dulcamara var. chinense. The Macartney collection (BM000942424) has here been selected as the lectotype in order to preserve usage, as it is a specimen of Solanum lyratum ; the Staunton sheet (BM000942515) is Solanum septemlobum .

Specimens examined.

China. Anhui: Wangshan, Ching 8698 (US); Hsi, 3 Jun 1935, Migo s.n. (A); Huoshan County, Fozlingshuiku, 12 Sep 1985, South Team 1429 (MO); Baijianshan, Xigou, Xuancheng, 400 m, 2 Nov 1998, Wang E-8 (MO); Chongqing: Chengkou County, 600 m, 31 Aug 1958, Dai 102263 (MO); Chengkou County, Qiganshan, 1750 m, 6 Sep 1958, Dai 102476 (MO); Chengkou County, Shuanghekou, Baichishan, 1100 m, 17 Sep 1958, Dai 102779 (MO); Fujian: Huxingshan, 2 May 1959, Huang 190030 (MO); Hinghwa and vicinity, 1 Sep 1926, Lin Pi 6434 (A); O-Boih Village, Hinghwa and vicinity, 11 Sep 1926, Lin Pi 6549 (A); Shaowu, Wong To plain, 18 Aug 1926, Niao 9475 (A); Mount Wuyi, road from Sangang to Hongdu, Wuyi Mt, 700 m, 2 Aug 1980, Wuyi Shan Team 80-0529 (MO); Gansu: Wudu County, Weizigou forest district, 2250 m, 27 Aug 1989, Ying et al. 0907 (MO); Guangdong: Ruyuan County, Tianjingshan Forest District, 21 Sep 1974, Anonymous 93909 (MO); Ruyuan County, Baishuizhai, Tixia village, 14 Aug 1935, Chen 10764 (MO); Lokchong, Kwangtung: Lokchong District, 25 Nov 1930, Chen 42264 (MO); 10 li from Nan Shung, 15 Dec 1927, Chun 5700 (A); Changjiangxiang, Renhua County, 800 m, 4 Sep 1958, Deng 7490 (MO); Meikengxiang, Xinlai County, 300 m, 1 Nov 1958, Deng 8306 (MO); Baishikeng, Jiu keng, Dinghu Mt, 200 m, 11 Jun 1964, Ding & Shi 1512 (MO); Boluo County, Luofushan, 26 Apr 1978, Guangdong Prov. 1978 Team 5908 (MO); Gaoyao County, Yankengxiang, 29 Mar 1957, Huang 162696 (MO); Samshui, near Dam-shui, May 1875, Lamont 1084 (BM); Yung-Yun City, Kai ngan tsz, 29 Nov 1932, Lau 635 (A); Mo Po Lo, Tung Koon, 6 Apr 1935, Lau 20766 (MO); Longmeng County, Shangping, vicinity of Shangping village, Nankunshan, Yonghan, 26 Dec 1956, Li Xuegen 200363 (MO); Wengyuan, Wengyuan County, 22 Sep 1935, Liu 24502 (MO); Gaoyao County, Hetaixiang, 6 Sep 1958, Liu 01965 (MO); Lian County, Tianguangshan, Yao’anxiang, 2 Nov 1958, Tan 60191 (MO); Yangshan County, Daping, Xiluxiang, 700 m, 27 Nov 1958, Tan 60451 (MO); Ding-hu Shan, 11 Jun 1964, Ting 1512 (A); Yam Na Shan [Yit nga shan] (Mei [Kaying] District), 4 Aug 1932, Tsang 21496 (GH, K); Lokchong, 18 Oct 1928, Tsiang 1403 (K); Jingyingsuo, Laoshan, Tianlin County, 600 m, 26 Nov 1957, Zhang 11000 (MO); Dongxing, 15 May 1959, Zhou 02801 (MO); Guangxi: Damiaoshan, Xiaosangxiang, Antai district, 480 m, 16 Nov 1958, Chen 17286 (MO); Danyashan, Guilin City, 27 Oct 1958, China Germany Team 026 (MO); Guilin City, 28 Oct 1958, China Germany Team 135 (MO x2); Nor Yut, Tai Ching Shan, 19 Jun 1935, Ko 55334 (A); Dabu, Guilin City, 130 m, 24 Oct 1993, Li 13078 (MO); Lingui County, 185 m, 25 Oct 1994, Li 14226 (MO); Haiyang, Haiyang shan, Lingchuan City, 500 m, 10 Jul 1995, Li 14773 (MO); Nanbianshan, Lingui, 185 m, 7 Aug 1997, Li 15968 (MO); Zhongfengxiang, Ziyuan county, 230 m, 13 Oct 1998, Li 489 (MO); Mianliang Cun, Bai sheng xiang, Napoxian, 800 m, 7 Nov 1998, Qin 2394 (MO); Ting-hsing-chai Shan, opposite Liang mao-shan, vicinity Pai-chou city, 20 Aug 1937, Taam Ying-Wah 39 (A); Changtan, Longjin County, 30 Nov 1959, Tan 57159 (MO); Daqingshan District, Longjin County, 550 m, 20 Dec 1957, Tan 57478 (MO); Tang Lung Village, Shap Man Taai Shan, SW of Shang-sze, Kwangtung [Guangdong] border, 3 Sep 1934, Tsang 24190 (A); Yao Shan, 1 Dec 1936, Wang, C.W., 40508 (A); Longsheng County, Huaping, Huaping Forest district, 780 m, 20 Sep 1984, Wei Yuzhong & Lu Qinghua 20474 (MO); Jingxi County, Wupingxiang, Wupingxiang, 29 Dec 1958, Zhang 14857 (MO); Guizhou: Yinjiang County, Zhangjiaba, Yinjiang Xian, along the trail between Zhangjiaba and Huguoshi on the W side of Fanjang Shan mountain range, 1200 m, 25 Sep 1986, Bartholomew et al. 1651 (BM); 29 Jul 1936, Deng Shiwei 90622 (MO); Jiangkou Xian, Heiwan River, on the SE side of the Fanjing Shan mountain range in the vicinity of the Ecological Station of the Guizhou Academy of Sciences, 560 m, 20 Aug 1986, Sino-American Guizhou Bot. Expedition 78 (A); Songtao Xian, Lengjiaba, in the vicinity of the confluence of the Xiaohe and Dahe Rivers, NE side of the Fanjing Shan mountain range, 820 m, 5 Oct 1986, Sino-American Guizhou Bot. Expedition 2159 (A); Tsunyi Xian, Liang Feng Yah, 10 Aug 1931, Steward et al. 263 (A); Tsao-Feng-San, Tsingchen, 29 Jul 1936, Teng 90622 (A); Henan: Teng-feng, Shao lin ssu im Kreise Teng fong, 670 m, Aug 1907, Schindler 165 (BM, G, L, S); Xinyang County, Huoshao Temple, 600 m, 17 Jul 1985, South Team T-0120 (MO); Xinyang County, Yuhuangshan, Yuhuangshan, 400 m, 17 Jul 1985, South Team T-0180 (MO); Shangcheng, Mao’ershishan, Yangqiaoxiang, 200 m, 8 Sep 1998, Wang G-73 (MO); Hong Kong: Chinese University of Hong Kong, 2 Nov 2004, Li Ming 43 (A); Hubei: Yichang, Ichang, (Patung District), Henry 110 (K); Yichang, Ichang, Henry 635 (K); Yichang, Nan-t’o and mountains to northward. Ichang, Feb 1887, Henry 2198 (K, US); Wuhan, along road near Wuhan Botanic Garden, 46 m, 30 Sep 2007, Knapp et al. 10142 (BM); Songbai, Songbaizhen, Shennongjia Natural Reserve, 950 m, 2 Sep 1996, Shi Shigui S-0950 (MO); Shennongjia Forest District, between Biacaoping and Bajiaomiao, 1400 m, 20 Sep 1980, Sino-American Expedition 1576 (A); Jianshi County, 800 m, 15 Aug 1997, Wang 97004 (MO); Hunan: Yizhang, Liyuan bao, 100 m, 13 Sep 1942, Chen 2392 (MO); Dafobin, 20 Aug 1950, Dahlström 255 (S); Xinning, Ma-Ling-Tung, Sinning Hsien, 21 Sep 1935, Fan & Li 523 (A, BM, ECON); Hangsha, Datankeng, Hangshan, 980 m, 21 Aug 1964, Huang 112416 (MO); Dongan, Shunhuangshan, 280 m, 1 Sep 2004, Liu Jin-Kui 667 (US); Yongshun County, west of Hunan Province, 600 m, 15 Aug 1958, Liu 9582 (MO); Hengshang, Fangguang Temple, Hengshan (Nanyue), 560 m, 11 Aug 1948, Liu 00373 (MO); Yizhang, Mt. Mangshan, 700 m, 9 Aug 2005, Xiao Bai-Zhong 4525 (K, US); Wugang County, Yunshan, Sanliting, Yunshan, 4 Sep 1950, Zhang 4495 (MO); Jiangsu: Soochow, 23 Sep 1926, Chiang 8236 (BH); Nanjing, Kiangsu province, 22 Oct 1966, Chiao 12926 (BH, G, K); Chiu-yong, 21 Jul 1926, Ling 12166 (GH); Nanjing, Nanking Kiangsu, 3 Oct 1929, Sun 206 (K); Nanjing, Nanking Kiangsu, 4 Oct 1930, Sun 727 (K); Jiangxi: Pailou County, Jiujiang, 110 m, 3 Sep 1995, Ce-ming 95683 (BM); Jiujiang, Lushan, Kiangsi Prov. Lu Shan. Huang Yen Ssu, 4 Sep 1932, Cheo 251 (ECON, K); Lu-shan-hsien, 1100 m, Chu 4085 (BM); Lushan, Shimenjian Gorge, W slopes of Lushan Mountains, 698 m, 27 Sep 2007, Knapp et al. 10139 (BM); Lushan, San Die Quan, (3 Steps Waterfall), Lushan Mountain, 661 m, 28 Sep 2007, Knapp et al. 10141 (BM); Kiennan, Sai Hang Cheung, near Tung Lei Village, 1 Aug 1934, Lau 4279 (BM, G, GH, S, US); Kiennan, Sai Hang Cheung, near Tung Lei Village, 1 Sep 1934, Lau 4379 (BM, G, GH, S, US); Wuning County, Wuningyan, Wuningyan, 350 m, 23 Oct 1994, Tan 941258 (MO); De-an Xian, Dingfeng, 160 m, 6 Nov 1996, Tan 11003 (MO); Fenyi City, Dagangshan, Fenyi city, 26 Aug 1985, Yao 9233 (A, K); Shaanxi: Mian County, from Changbazi to Xiasbianhe, 750 m, 2 Oct 1952, Fu 6130 (MO); T’ai-hua Shan, 1 Aug 1932, Hao 3832 (K); Huashan, Shensi, Hua-shan, 26 Aug 1932, Hao 4192 (K); Zhenba County, 820 m, 20 Aug 1999, Wang 522 (MO); Zhouzhi Xian, 1200 m, 28 Jul 1999, Zhu et al. 2386 (MO); Yang Xian, 1600 m, 8 Jul 1999, Zhu et al. 2909 (MO); Taibai Xian, 1800 m, 22 Jul 2000, Zhu et al. 3166 (MO); Shandong: Meng Shan, Fei Hsien, 350 m, 25 Jul 1936, Cheo & Yen 184 (BM, G, GH); Tai Ming Hu, 4 Sep 1920, Chiao 3030 (B, K, PE, US); Shan Ling, Gaaoshan, 500 m, 29 Sep 2006, Guo Chengyong 5470 -5 (US); Ningyang, Guo Chengyong 51386 -1 (HITBC); Zoucheng, Guo Chengyong 52414 -1 (HITBC); Yantai, Bamboo temple, 8 Oct 1980, Herb. Forbes 8780 (BM); Shanghai: British Consulate, 26 Sep 1982, Carles s.n. (BM); Shanghai, 1887, Faber 8 (K); Shanxi: Tongshan gongshe, Yuangu County, 800 m, 14 Oct 1982, Fu 18757 (MO); Yüan-chü Distr., Ye-cho-shan, Fu-chis-ho, 1200 m, 18 Jul 1924, Smith 6432 (MO); Sichuan: Yajiang Xian, along first tributary fo the Yalong Jiang E of the city of Yajiang, 541 m, 6 Aug 2006, Boufford et al. 35933 (A); Xinlong Xian, confluence of the nearly dry tributary and Yalong Jiang at intersection of road from Ganzi to Xinlong (highway 217) and road to Baiyu, just downstream from village of Acha (Echa), 1244 m, 25 Aug 2006, Boufford et al. 37306 (A); Omei Shan, 1100 m, 22 Aug 1938, Chiao & Fan 358 (A, US); Omei Xian, Mt. Omei, Dec 1942, Chow 7261 (A); Mt. Omei, Tapingssu, 16 Aug 1938, Chow 8152 (A); Kiating, Pasientung, 8 Dec 1938, Chow 8986 (A); Kiating, outside of the west gate, 20 Nov 1938, Chow 8920 (A); Mount Omei, Omei Hsien, 18 Aug 1928, Fang 3164 (A, K, US); Mowchow, Mow Hsien, 20 Sep 1928, Fang 5503 (A, K, US); Lifan, Mung-twin-ko, Aug 1941, Hu 1982 (A); Taking, 1958, Li 78542 (A); Ta-tien, 1900 m, 2 Jul 1922, Smith 2265 (A, MO); Chengkow, 1958, Tai 102263 (A); Dujiangyan, Qingchengshan, Si Chuan, Guan Xian, 800 m, 20 Aug 1987, Wang Zhong-tao 870423 (K); Kwang-yun Hsien, and vicinity, 11 Oct 1930, Wang 22655 (GH); Mount Emei, 800 m, 2 Sep 1995, Xu 4773 (MO); Jeui-shen Xian, 10 Jun 1932, Yu 1015 (A); Dujiangyan, Zhu Dahai 2232 (HITBC); Tibet: Chi-na-tung, Tsa-wa-rung [Sikang province], 2500 m, Aug 1935, Wang 65289 (A); Yunnan: Momien, 30 May 1868, Anderson s.n. (K); Lijiang, Sashiba, 29 Aug 1939, Ching 21378 (A); Lan ngy Tsin, 14 Jul 1904, Brother Ducloux 2728 (A); Muli, Consinling near Ngerya on the border of Chungtien, in side valley, 2300 m, 24 Aug 1939, Feng 2778 (A); Si-chour Xian, Faa-doou, 1400 m, 13 Sep 1947, Feng 11662 (A); Mar-li-po, Hwang-jin-in, 1100 m, 13 Nov 1947, Feng 13234 (A); Lichiang, 2743 m, Jun 1910, Forrest 5922 (BM, K); Tengyueh, hills around Tengyueh, 1829 m, Jul 1912, Forrest 8630 (K); Gongshan, Bingzhonglou, Nujiang Lisa Aut. Pref., Dan Dong village outskirts on road to Bingzhonglou, near Gongshan, 1450 m, 9 Sep 1997, Gaoligong Shan Expedition 1997 8738 (E); Mephyr, 1524 m, Henry 11855 (A, K, US); Chengjiang Xian, 18 Jun 1940, Hou 74591 (MO); Lijiang, Judian, Qingzhang Team, 425 (HITBC); Lijiang, Yangtze watershed, prefectural district ft Likiang [Ligiang], slopes of Likiang Snow Range, May 1922, Rock 4311 (A, US); Ho-ch’ing, inter Hoching et Singquch, 2600 m, 27 Sep 1914, Schneider 2705 (GH, K); Lan-ping Xian, 17 Aug 1933, Tsai 53712 (A); Chiu-kang, W of Chamutung, 2000 m, Oct 1935, Wang 67159 (A); Wei-si Xian, Yeh-Chih, 2500 m, Jul 1935, Wang 67927 (A); A-tun-tze, 2700 m, Sep 1935, Wang 70376 (A); Lung-kai, Cheng-kiang, 2350 m, 4 Aug 1939, Wang 41503c (A); Atuntze, Mt Kaakerpu, 2800 m, 20 Sep 1937, Yu 10384 (A, BM); Muli, Muli, Wachin, Buraton, 2900 m, 2 Nov 1937, Yu 14713 (A, BM); Chungtien, Haba, 2600 m, 22 Nov 1937, Yu 14941 (A, BM); Salwen Valley, Dara, 1800 m, 24 Jun 1938, Yu 22007 (A); Zhejiang: Hangzhou City, Oct 1957, Jin Yilang 322 (MO); Hupaoma’er shan, Hangzhou City, 25 Sep 1953, Wu 1306 (MO); Hangzhou, Zhu Qiugui 292 (HITBC).

Japan. Hokkaido: Ishikari, 10 Sep 1903, Arimoto s.n. (GH); Honshu: Toyko, Koishikawa, 22 Aug 1893, Anonymous s.n. (US); Toyko, Jul 1887, Anonymous 338 (US); Yamaguchi, Hakusan Shrine, in vicinity of Kika Park, base of east slope of Castle Mountain(Shiro-yama) National Forest, west bank of Nishiki River, Yokoyama Village in Iwakuni City, 25.5 miles west of Hiroshima City, 18 m, 19 Sep 1953, Charette 1409 (S, US); Shizuoka, Mt. Takakusa-yana, Uchinoya, Okabe-cho, Shida-gun, a mountain 5 km from the seashore of the Pacific Ocean and facing the sea, 150 m, 4 Sep 2006, Konta 35512 (US); Tokyo, 18 Aug 1878, Matsumura s.n. (US); Wakayama, Takaike, Kozagawa, Wakayama Pref., Higashimuro-gun, Kozagawa-cho, Takaike, 20 m, 5 Nov 1964, Matsushita 125 (US); Shinano, Okayama-mura, Shimo-minochi-gun, 14 Aug 1953, Mizushima 12628 (S); Kyoto-fu, Oomoto-honbu, Ten’on-kyo, Kameoka-shi, 19 Sep 1991, Murakami 134 (A); Miyagi Pref., Sendai-shi, Katahira-cho, N Honshu, 11 Aug 1972, Sato 72811 (B); route from Mizundo to summit of the Mt. Atago, Saga, Ukyo-ku, Kyoto-shi, 330 m, 10 Oct 1991, Takahashi 2215 (A); Kyoto, Is. Anja, Obase, Nishi-oura-chiku, Maiduru-shi, 10 m, 9 Nov 2000, Takahashi & Sawada 30261 (A); Kii, Mt. Koya, en route from Daimon to Yatate, 650 m, 9 Sep 1962, Tamura 9362 (S); Ishikawa, Oosugidani, Oosugi-uemachi, Kamatsu-shi, 360 m, 11 Oct 1992, Tsugaru & Murata 17048 (A); Kyoto, Ashiodani, Yotsuya, Hiyoshi-cho, Funai-gun, 250 m, 1 Sep 1994, Tsugaru & Takahashi 20836 (A); Kyoto, Mt. Kunimiyama, Onyu, Nishi-Oura-chiku, Maidzuru-shi, 10 m, 9 Nov 1997, Tsugaru & Takahashi 25940 (A); en route from Kobama to Kaketsu, Amino-cho, Takeno-gun, 10 m, 12 Nov 1998, Tsugaru & Takahashi 27273 (A); Mt. Takahura, Okayamayen, 15 Aug 1952, Uno s.n. (A); Kyushu: Miyazaki Pref., Miyakonojo City, Yamanokuchi, 17 Aug 1950, Hatusima 14243 (US); Yoshino, Kugoshima City, Kyusyu, 4 Oct 1930, Naito s.n. (US); Nagasaki, Is. Tsushima, Shimoagata-gun, Izuhara, 22 Oct 1973, Ohashi et al. 27 (A, US); Shikoku: Tokushima, 1919, Krug 239 (B); Tokushima, 1919, Krug 663 (B).

Republic of Korea. Quelpaert, Nov 1906, Faurie 775 (BM); Soonchun, Schulla Prov, 3 Sep 1934, Smith s.n. (GH); sin. loc, 300 m, Oct 1907, Taquet s.n. (G); 27 Aug 1908, Taquet 1146 (K); Quelpaert, 25 Jun 1908, Taquet 1147 (G, K).

Taiwan. Pa-hsien-shan Forest Recreation Area, T’ai-chung Hsien, Ho-p’ing Hsiang, 800 m, 27 Nov 1997, Bartholomew et al. 7613 (GH); Yangmingshan, Taipei County, 18 Feb 1962, Chuang & Lin Pi 4824 (A); Biyoritne [?], 27 Mar 1903, Faurie 322 (A, BM, G); Kaohsiung, Takao [?], Jan 1914, Faurie 638 (BM, G); Hualien, Yen-tzu-k’ou, 14 Dec 1990, Huang 15045 (A); Compartment 114, Wushikon, Taichung, 14 Oct 1957, Liu s.n. (A); sin. loc., 1864, Oldham 340 (BM); Hsiulin Hsiang, Taroko National Park, 500 m west of Loshao, at road marker 153.5 along Prov. Hwy. 8 between Loshao and Songshan Tunnel, 1100 m, 15 Jun 1999, Peng & Anderberg 17285 (S); Mosha, 9 Jul 1912, Price 816 (K).

Vietnam. Cao Bang: Distr. Ha Lang, municipality Dong Loan, vicinity of Ban Lung and Lung Phuc, about 50 km to E from Cao Bang town, 500 m, 25 Nov 1998, Averyanov CBL-657 (MO); Tonkin, base du massif calcaire au pied du camp militaire, Aug 1943, Pételot 8464 (A).

Figures

Figure 2. Leaf variation in members of the Dulcamaroid clade. A Leaf variation along a single short stem, Solanum salicifolium (Barboza et al. 3467, Catamarca, Argentina) B Thick leathery leaves of Solanum macbridei (Nee & Solomon 30182, La Paz, Bolivia) C Uneven leaf lobing in Solanum dulcamara (Knapp IM- 10102, Tuscany, Italy) D Variable membranous leaves of Solanum lyratum (Knapp et al. 10139, Jiangxi, China).

Figure 61. Solanum lyratum Thunb. (A-D drawn from Smith 6432 and Knapp 10142 E-G drawn from Peixing Tan 57179 G drawn from Smith 2265 H drawn from Junsheng Ying 0907). Illustration by Bobbi Angell.

Figure 62. Distribution of Solanum lyratum Thunb.