ABSTRACT: Specimens in several large herbaria were studied for a worldwide revision of the genus Euonymus L. (Celastraceae). These specimens included 14 new species, Euonymus kengmaensis, E. vaganoides, E. aculeola, E. potingensis, E. hukuangensis, E. ficoides, E. subcordata, E. chengii, E. dolichopa, E. laxicymosa, E. hui, E. parasimilis, E. wui, and E. pereoriacea, from east and south Asia (mostly from China).

Species E. bockii Loesener ex Diels similis, sed foliis basi attenuatis, acuminatis vel acutis, capsula immaculata differt; species E. theifolii Wallich ex Lawson similis, sed foliis chartaceis vel papaverinis, margine crenulatis, nervis lateralibus curvis prorsum, non extensis ad margine, paniculatis-cymis longior, usque ad 8 cm longis, 6 cm diam. differt.

Shrubs (scandent) to small trees, to 8 m tall; stems gray-green to gray-brown, terete, glabrous; twigs green to light green, terete; winter buds with several purple-brown scales. Leaves chartaceous or papyraceous, ovate to elliptic, 8-12 cm long, 3-5 cm wide, glabrous, acuminate or acute at both ends; margin crenulate; lateral veins 4-6 pairs, slightly curved upward, disappearing before reaching margin; petiole ca. 4 mm long. Panicle-cymes terminal or axillary, many flowered, to 8 cm long, 6 cm in diam. Flowers 4-merous, ca. 7 mm in diam.; petals greenish-yellow or yellowish-green, nearly round. Capsule globose or subglobose, brown or yellow-brown, opening by 4 lobes at maturity. Seeds 2 in each locule, dark brown, covered by orange aril.

Distribution: China (endemic to Yunnan).

Additional specimens examined: CHINA. Yunnan: Kengma Xian, C. W. Wang 72919, 72914 (KUN); Jingdong Xian, M. K. Li 3338, 3390 (KUN), Cangyuan Xian, C. W. Wang 73224 (A), 76835 (A, KUN); Longchuan Xian, S. K. Wen 580602 (KUN); Longling Xian, J. Chen 683 (KUN), H. T. Tsai 55652 (A).

This species is similar to E. bockii Loesener ex Diels in leaf size, but differs in the attenuate, acuminate or acute leaf base and capsules without small spots. It is also similar to E. theifolia Wallich ex Lawson in the color and shape of the fruit, but differs from it by the larger, papyraceous leaves with crenulate margins, the lateral veins curving forward and ending before extending to the margin, and in the long panicle-cymes to 8 cm long and 6 cm in diam.

Species E. vagani Wallich similis, sed foliis crassis coriaceis, maginis, margine paucis crenulatis, capsula magina, globosa, maculatis albis differt.

Shrub-like, climbing; stems and branches terete; twigs sturdy. Leaves clustered at apex of branches, elliptic-obovate, 8-10 cm long, 3-5 cm wide, thick coriaceous, base wide-attenuate, apex cuspidate; margin sparsely crenulate on margin, lateral veins 5-7 pairs, prominent on lower surface; petiole sturdy, ca. 3 mm long. Flowers not seen. Capsules at apex of branches, axillary, solitary or paired, nearly globose, 6-9 mm in diam., purple, densely small white spotted when dry. Seeds 2 in each locule, covered by orange aril.

Distribution: China: Guangxi, Hunan, Yunnan.

Additional specimens examined: CHINA. Guangxi: Guanyang Xian, Z. Y. Chen 52179 (KUN); Hunan, Xinning Xian, L. H. Liu 15027, 15339 (KUN); Yunnan: Pingbian Xian, Bai Mu Tan, 1250 m, C. W. Wang 82658 (KUN); Yongshan Xian, C. W. Wang 84851 (KUN).

This species is similar to E. vagans Wallich in the texture of the leaves, which are more thickly coriaceous and have fewer crenulations on margin; the large globose capsules are white spotted.

Proxima E. echinati Wallich, sed frutices pumiles vel nanis, foliis non petiolis, capsula echinata, echiatis tenuis vel filiformis differt.

Dwarf shrubs, ca. 30 cm tall; young branches 4-angled, green-brown. Leaves sessile, ovate, 3-5 cm long, 1-1.5 cm wide, margin serrulate. Flowers not seen. Capsules axillary, solitary or 2 or 3 clustered, brown to pink, nearly globose ca. 5(-8) mm in diam., with very small, slender, needle- or thread-like spines; seeds obovoid, black, covered by aril.

Distribution: China, endemic to Yunnan.

This is a very unusual species in section Echinococcus in being the only dwarf shrub so far known. It is similar to E. echinata Wallich, but differs in its stature (a dwarf shrub), sessile leaves, and the needle- or thread-like spines of the capsules.

Species E. acanthocarpi Franchet similis, sed capsula pausis echinatis vel spinis, foliis margine integris, minutis revolutivis differt.

Shrubs or small trees or climbing. Stems and branches brown or dark brown, obscurely angled when young. Leaves oblong-elliptic to elliptic, ca. 6-10 cm long, 3-4.5 cm wide, attenuate at both ends, margin entire, lateral veins 5-7 pairs, disappearing before reaching margin; petiole ca. 4 mm long. Cymes several flowered, peduncle 3-4 cm long; flowers 4-merous, disc large, perianth not seen. Capsules with several sparse spines. Seeds immature.

Distribution: China, endemic to Hainan.

Additional specimens examined: China: Hainan: Baoting (Poting) Xian, ca. 1100 m, F. C. How 72803 (A).

This species is similar to E. acanthocarpa Franchet, but differs from it by having fewer spines on the fruit and entire leaves with somewhat revolute margins. This is the only species from Hainan Island with spines on the capsule.

Species E. hainanensis Chun & How similis, sed foliis margine intergris, glabris superficiebus ambabus, nervis lateralibus obscuris differt.

Shrubs to small trees, to 6 m tall; stems and branches terete, brown, twigs sturdy. Leaves obovate-elliptic to elliptic or obovate-elliptic, 6-9 cm long, 2.5-4 cm wide, base acuminate, cuneate or acuminate, apex short apiculate; margin entire; lateral veins 7-8 pairs, obscure or invisible, disappearing before reaching margin; petiole 4-6 mm long. Peduncles axillary to 9 cm long; cymes 1-several flowered. Flowers 4-merous, sepals nearly round, petals nearly round. Fruit 1-several, with pedicel to 9 cm long. Capsules 1-1.5 cm in diam., lobes 4, opening at maturity. Seeds 2 in each locule, nearly round, ca. 4 mm in diam., red when fresh, red or black when dry, covered by aril.

Distribution: China: Guangdong, Hunan.

Additional specimens examined: China: Guangdong: Ruyuan Xian, S. P. Ko 53571 (MO); L. Deng 5538 (MO). Hunan: Yizang Xian, S. H. Chen 2843 (MO); P. H. Liang 85338 (MO); Mangshan, M. X. Huang 111407, 111823 (MO).

This species is similar to E. hainanensis Chun & How, but differs from it in the entire leaves being glabrous on both surfaces, and in the obscure lateral veins.

Species E. rhederiani Loesener similis, sed foliis ovatis vel ovatis-elliptis, crassis coriaceis, capsula ala, alis brevis differt.

Shrubs 1-2 m tall, evergreen; stems and branches terete, purple or black; twigs sturdy, brown to purple. Leaves persistent, thick coriaceous, ovate or ovate-elliptic, 9-12.5 cm long, 4-6 cm wide, base nearly round, margin entire, apex acute to acuminate; lateral veins 8-11 pairs, curved forward near margin; petiole sturdy, to 1 cm long. Flower solitary, axillary, 5-merous, pedicel to 10 cm long. Capsules nearly globose, ca. 1.5 cm in diam., with 5 obscure wings at maturity. Seeds 2 in each locule, globose, covered by orange aril.

Distribution: China, endemic to Yunnan.

Additional specimens examined: China: Yunnan: Malipo Xian, Pan Chia Chu, K. M. Feng 12709 (A, PE), K. M. Feng 13556 (A, PE).

This species with large coriaceous leaves and large smooth fruit with five obscure wings is similar to E. rhederiana Loesener, but differs from it by its mainly ovate or ovate-elliptic, thick coriaceous leaves and capsules with very short wings.

Species E. chloranthoidis Yang similis, sed capsula 4 lobis, foliis basi cordatis, superior impressinervis differt.

Shrubs ca. 2 m tall; stems and branches terete, 4 angled or wing-like when young, twigs sturdy. Leaves sessile, ovate-oblong, 13-15 cm long, 4-6 cm wide, base cordate, margin coarsely serrate, apex acuminate, lateral veins somewhat impressed on upper surface, prominent on lower surface. Flowers not seen. Fruit very young, lobes 4, wing-like.

Distribution: China, endemic to Guangxi.

This species is similar to E. chloranthoides Yang, but differs by having a 4-merous capsule, cordate leaves without a petiole, and the nerves impressed on the upper surface. It is still in need of further study, since there are no specimens besides the type, which has only very young fruit.

Species E. kachinensi Prain similis, sed foliis margine intergris, sessile vel subsessile, capsula 4 alis-angulis, floris viridis differt; proxima E. nitidi Bentham and E. oblongifolii Loesener & Rehder, ab illa et hac capsula 4 valde alis-angulis et apice emaginatis differt.

Shrubs to 3 m tall; stems and branches terete, nearly 4-angled when young, twigs slender, thin. Leaves sessile or subsessile, chartaceous, elliptic, 5-9 cm long, 2.5-3.5 cm wide, acuminate or attenuate at both ends, margins entire, apex caudate, lateral veins not visible. Cymes with several flowers. Flowers light green, 4-merous, 3-4 mm in diam. Capsules ca. 1.5 cm in diam., ca. 2 cm long, apex emarginate, strongly angled, angles 4, wing-like. Seeds 2 in each locule, round or nearly globose covered by orange aril.

Distribution: China, Guangdong, Hainan.

Additional specimens examined: China: Hainan: Baoting Xian, Tai Ping Gang, F. C. How 72158, 72833 (MO).

Etymology: named for Ching-yung Cheng of Beijing Medical University, who has made a lifelong study of the Celastraceae of China and who interested me originally in this family.

This species is similar to E. kachinensis Prain, but differs from it by the sessile or subsessile leaves with entire margins, the capsule with four large wing-like angles, and the green flowers. It is also similar to E. nitida Bentham and E. oblongifolia Loesener & Rehder in leaf shape, but the fruit differs totally in its 4 wing-like angles and emarginate apex.

Proxima E. oblongifilii Loesener & Rehder et E. nitidi Bentham, ab illa et hac capsula 5 lobes, apice emarginatis differt; species E. chengi J. S. Ma similis, sed foliis chartaceois, capsula 5 lobes differt.

Shrubs ca. 2-3 m tall; stems and branches terete, twigs nearly sturdy. Leaves elliptic, 7-8 cm long, 2-3 cm wide, cuneate or attenuate at both ends, margin nearly entire but with obscure crenulations, lateral veins obscure; petiole 3-4 mm long, sturdy. Flowers not seen. Fruit solitary, yellow (from label data); capsule subglobose, 7-8 mm in diam., clearly 5-angled, emarginate or indented at apex. Seeds 2 in each locule, black or dark purple, partly covered by orange aril.

Distribution: China, endemic to Guangxi.

Etymology: Merrill apparently realized this species to be undescribed, but although he used the name E. dolichopa on herbarium specimens, he never published a description.

This species is characterized by its globose fruit with 5 angles and emarginate or indented apex and leaves with remote, obscure crenulations on the margin. It is similar to E. oblongifolia Loesener & Rehder and E. nitida Bentham in the shape and size of the leaves, but the leaves are chartaceous, and the fruit with 5 lobes is totally different.

Species E. salicifolii Loesener similis, sed in foliis magnis, lanceolatis, pedunculis usque ad 10-15 cm longis, floris magnis, usque ad 8 cm diam., capsula magna, 5 loba, in 5 valvis in adultis differt.

Shrubs to 4 m tall; stems and branches terete, sometimes young twigs 4-angled. Leaves subsessile or petioles to 10 mm long, blade lanceolate or sometimes oblong-elliptic or elliptic-lanceolate, 12-16 cm long, 3.5-4.5 cm wide, margin nearly entire, sometimes with small, obscure crenulations; lateral veins forming network before reaching margin. Cymes several flowered, peduncle 10-15 cm long; flowers 5-merous, sepals semi rounded, petals dark pink to purple or purplish-brown, ca. 8 mm in diam., margin ciliate. Capsules nearly globose, but clearly 5-angled, apex emarginate, opening by 5 valves at maturity. Seeds ovoid, 2 in each locule, dark purple, lower part covered by orange aril.

Distribution: China (Guangxi, Yunnan) and Vietnam.

Additional specimens examined: China: Yunnan: Xichou Xian, Nan Chang Xiang, S. Z. Wang 17 (KUN), 357 (KUN, PE); Malipo Xian, Sze Tai Po, K. M. Feng 13866 (A, KUN). Vietnam: Tonkin, NE Mon Cay, Pac Si and vicinity, W. T. Tsang 26916 (A).

This is a very unusual species with long cymes and lanceolate leaves; the leaves are similar to those of E. salicifolia Loesener in size, but differ in their oblong-lanceolate shape. It also differs in the petals with ciliate margins and the large, round capsule, which is emarginate at the apex.

Species E. tsoi Merr. similis, sed in foliis ellipticis; capsula circa 6-7 cm diam., paucis angulatis, apice brevis apiculo differt.

Trees to 25 m tall; stems and branches terete, greenish-brown; twigs sometimes 4-angled, brownish-green. Leaves elliptic, 8-14 cm long, 3-6 cm wide, acuminate at both ends; margins crenate or serrate, lateral veins 8-11 pairs, disappearing before reaching margin; petiole ca. 1 cm long. Cymes to 10 cm long. Flowers not seen. Capsules nearly globose or ovoid-globose, brown, clearly 4-angled, 6-7 mm broad, ca. 8 mm long, brown, slightly apiculate at apex. Seeds 2 in each locule, immature, covered by orange aril.

Distribution: China, endemic to Sichuan.

Etymology: Named in honor of Dr. Shiu Ying Hu of the Arnold Arboretum, who collected the type specimen and who has spent her lifetime studying the Chinese flora.

This species is similar to E. tsoi Merr. in fruit, but differs from the latter by its elliptic leaves. The medium to small sized nearly globose fruit, ca. 6-7 mm in diam., with a very short apiculate apex, is similar to species of Sect. Ilicifolia, but is clearly 4-angled, and the color is deep brown, which is seldom found in other species of Sect. Ilicifolia.

Species E. hamiltoniani Wallich similis, sed in foliis crassis coriaceis, ellipticis vel rotundis-ellipticis, basi et apice rotundis, margine integris and obscuris crenulatis differt.

Trees to 10 m tall, dbh ca. 30 cm. Stems and branches terete. Leaves thick coriaceous, elliptic to broadly elliptic, 9-10 cm long, 4-6 cm wide, base nearly round, margin entire to finely crenulate, apex round or nearly round; lateral veins 6-9 pairs, disappearing before reaching margin; petiole 3-6 mm long, sturdy. Cymes ca. 8 cm long. Flowers not seen. Fruit obpyramidal, ca. 1.5 cm in diam., 8 mm long, clearly 4-angled or sometimes tetragonal. Seeds 2 in each locule, triangular, dark brown, covered by orange aril.

Distribution: China, endemic to Yunnan.

Additional specimen examined: China: Yunnan: Pingbian Xian, 9 Jul 1934, H. T. Tsai 62517 (A).

This species is similar to E. hamiltoniana Wallich in fruit shape and in size, but differs from it by the thick, elliptic to broadly elliptic, coriaceous leaves rounded at both ends and the entire to obscurely crenulate margins. It is an unusual species in the Section.

Species E. macrocarpi Gamble ex Oliver similis, sed in foliis intergris, apice apiculo differt; species E. nitidi Bentham et E. oblongifolii Loesener & Rehder similis, sed illa et hac capsula 4 angulatis, obovoideis, magnis usque ad 14 mm long et 10 mm diam. differt.

Shrubs 5-6 m tall; stems and branches terete. Leaves elliptic, coriaceous, 4-10 cm long, 2-3 cm wide, acuminate or acute at base, margins entire, slightly caudate at apex; lateral veins 5-7 pairs, disappearing before reaching margin; petiole 3-4 mm long. Flowers not seen. Capsule solitary, axillary, obovoid, 4-angled, ca. 1 cm broad, ca. 14 mm long, base attenuate, apex apiculate; pedicel ca. 1 cm long, opening at maturity. Seeds 2 in each locule, globose, dark brown, covered by orange aril.

Distribution: China, endemic to Yunnan.

Additional specimen examined: China: Wenshan Xian, Lao Jun Shan, K. M. Feng 11214 (A).

Etymology: Named in honor of Professor Cheng-yi Wu, emeritus director of the Kunming Institute of Botany and the Yunnan Branch, Chinese Academy of Sciences, and who has made significant contributions to our understanding of the flora and biogeography of China.

This species is similar to E. macrocarpa Gamble ex Oliver in fruit, but the leaf margins are entire, and the fruit apex is apiculate. It is also similar to E. nitida Bentham and E. oblongifolia Loesener & Rehder in leaf characteristics, but the fruit are different in being large, obovoid and 4-angled; they open at maturity as in E. macrocarpa Gamble ex Oliver.

Species E. wrayi King similis, sed in fructis parvis, foliis multo parvis, coriceis differt.

Shrubs ca. 2 m tall; stems terete, brown, twig sturdy, brown, 4-angled when young. Leaves thick coriaceous, elliptic or broadly-elliptic, 4-5 cm long, 2-3 cm wide, base rounded, upper 2/3 of margin serrate or crenulate, apex nearly round or acuminate; lateral veins obscure; petiole ca. 3 mm long, sturdy. Flowers not seen. Fruit solitary, pedicel ca. 1 cm long; capsule obturnate, brownish red, ca. 1 cm broad., 8 mm long, 4-angled, opening by 4 valves at maturity. Seeds 2 in each locule, subglobose, dark brown, covered by orange aril.

Distribution: China, endemic to Yunnan.

Additional specimens examined: China: Yunnan: Xichou Xian, Fa Dou, Tai Yang Shan, 1500 m, Q. A. Wu 8027 (KUN); C. W. Wang 86087 (KUN, PE).

This is an unusual species with small, thick, coriaceous, elliptic to broadly elliptic, mostly serrate or crenulate leaves with the lower part entire nearly to the base. The 4-angled fruit is shorter than wide. The species is similar to E. wrayi King in capsule features, but differs from it by being a small shrub with much smaller, coriaceous leaves.

Acknowledgments:

This work, included in the program of the Flora of China, was supported by the National Natural Science Foundation of China, the Arnold Arboretum of Harvard University, and the Missouri Botanical Garden, to whom my sincere thanks are due. I thank the curators and keepers of the herbaria of A, B, BM, CDBI, GH, IBK, IBSC, K, KUN, LBG, MO, PE, PEM, SM, SYS and SZ who permitted me to use their collections or who kindly sent me specimens on loan. I especially thank Dr. Anthony Brach and Emily Wood for their assistance and help in the Harvard University Herbaria, Dr. Gustavo A. Romero of the Oakes Ames Orchid Herbarium of the Harvard University Herbaria for help in preparation of the plates and the editorial work on this paper, Mr. Li Xuedong of the Kunming Institute of Botany, and Mrs. Ban Qin of the Institute of Botany, Beijing, for valuable help at their institutions, Dr. Ihsan Al-Shehbaz at the Missouri Botanical Garden for kindly sending me all the specimens of the genus for this study. My special thanks are due Dr. David E. Boufford of the Harvard University Herbaria, not only for his valuable help and longtime encouragement of my research work when I was in Beijing during the past ten years, but also for arranging and assisting in my research visit to the Harvard University Herbaria in 1995-1997. Without this visit the work could not have been done. Finally I would like to acknowledge my great indebtedness to my Ph. D supervisor, Professor Ching-Yung Cheng, of Beijing Medical University, who has worked on the Celastraceae in the flora of China for about 30 years; some of this work is based on her manuscript for Flora Reipublicae Popularis Sinicae (in Chinese), however, the opinions expressed in this paper are my own and may not reflect her views precisely.