NOVON 2(2): 124-128. 1992.

Notes on the Solanaceae of China and Neighboring Areas

William G. D'Arcy

Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A.

Zhang Zhi-yun

Institute of Botany, Academia Sinica, 141 Xizhimenwai Dajie, Beijing 100044, China

ABSTRACT. Nomenclatural changes are made for the Flora of China, affecting Anisodus, Archiphysalis, Leucophysalis, Physaliastrum, and Scopolia.


The nomenclature relating to the Asian genus Anisodus and its type species differs in various floras and revisions. The generic name and its type species are clarified below, and another species is transferred from Scopolia to Anisodus.

Since Anisodus was described in 1824, the taxonomic validity of its separation from Scopolia Jacq. (1764) has been disputed. Some workers have simply placed species of Anisodus in Scopolia (Dunal, 1852); others have recognized Anisodus as a section or series in Scopolia (Wettstein, 1895; Weinert, 1972); and others have considered it a distinct genus (Pascher, 1909; Chen & Chen, 1977; Wu & Chen, 1978; Sandina & Tarasevich, 1982). It will be treated as a distinct genus in the Flora of China (Zhang et al., in press).

Scopolia and Anisodus are members of the solanaceous tribe Hyoscyameae, characterized by capsular fruits that are partly or completely covered by the persistent fruiting calyx and strongly curved embryos contained in compressed seeds. Both genera include herbaceous plants with subactinomorphic flowers and five stamens situated at the level of the corolla mouth. However, in addition to grossly different overall appearance, the groups differ in the conspicuous details shown in Table 1.

Sandina & Tarasevich (1982) studied the pollen of Anisodus (Whitleya) and Scopolia and found differences, particularly in the structure of the pores and the pattern of the exine, that they considered to be of generic significance. Differences in the pollen of Scopolia (S. carniolica) (Punt & Monna-Brands, 1980) and Anisodus (A. carniolicoides (as S. carniolicoides), A. tanguticus (as A. mairei), and A. acutangulus (Zhang & Lu, 1984)) are contrasted in Table 1.

Anisodus and Scopolia are widespread in the Old World; their geography was reviewed by Weinert (1972) and by Lu & Zhang (1986). When separated, the two genera have distinct geographical ranges. Anisodus occurs in the Himalayan region from western Nepal to Yunnan, China, and northward from Assam along the central Yangtze--Mekong--Salwin watersheds (Hengduan Mountains) of China to nearly 39;dgN latitude. The two species of Scopolia show a disjunction between Japan, S. japonica Maxim., and eastern Europe, S. carniolica Jacq.

Weinert (1972) referred to an unpublished dissertation by Semenowa (1955), which we have not seen, but names he attributed to her in his publication were not effectively published by her nor validly published by him.

Nomenclature of Anisodus and A. luridus

Anisodus Link in Sprengel, Syst. Veg. 1: 699. 1825 [1823] fide Flora 9: 2, 495. 1826, late 1824 fide Stafleu & Cowan 5: 813]. Scopolia sect. Anisodus Dunal in DC., Prodr. 13(1): 555. 1852. TYPE: A. luridus Link ex Sprengel.

Whitleya Sweet, News of Lit. & Fashion 3: 108. 13 Aug. 1925; Brit. Flower Gard. 2: 125. 1 Oct. 1825 [dates fide Airy Shaw, J. Bot. 85: 192--193. 1937]. TYPE: Whitleya stramonifolia Sweet.

Anisodus was described by Link in Sprengel's edition of Caroli Linnaei ... Systema vegetabilium, edition 16, which actually appeared before its title page date of 1825. The original publication, which included the description of A. luridus, cited as a synonym Nicandra anomala Link, which was not validly described until the following year by Link and Otto (see below). The material on which Anisodus was based was grown in Berlin from seed obtained from England that was noted as being from Nepal. About a year later, the genus was again described under the name Whitleya from different material that also had come into cultivation in Europe from seed obtained in Nepal.

The plant known as Anisodus luridus was described four different times with three different epithets in publications that bear the frontispiece date 1824 or 1825; confusion about its correct name persists. The following reviews the history of naming this plant and presents the correct name.

Anisodus luridus Link in Sprengel, Syst. Veg. 1: 699. 1825 [late 1824 fide Stafleu & Cowan 5: 813]. Scopolia lurida (Link) Dunal in DC., Prodr. 13(1): 555. 1852. TYPE: same as Nicandra anomala.

Physalis stramonifolia Wall. in Roxburgh, Fl. India. ed. Carey, 2: 242. 1824. [Mar.--Apr. 1824 fide Stafleu & Cowan 4: 957]. Scopolina stramonifolia (Wall.) Kuntze, Rev. Gen. Pl. 2: 452. 1891. Scopolia stramonifolia (Wall.) Shrestha, Bull. Dept. Med. Pl. Nepal 2: 81. 1969. Scopolia stemonifolia Chen Cheih & Chen Chung-lien, Acta Phytotax. Sin. 15(2): 61. 1977, orthographic variant. Scopolia stramonifolia (Wall.) N. P. Balakrishnan, Bull. Bot. Surv. India 22: 176. 1980 [1982] redundant combination. TYPE: cultivated, London, from seed obtained from Robert Henry Jenkinson, perhaps through Reginald Whitley, noted as being from Nepal, specimen not known. Nepal, herb. Wallich 2632 not seen, IDC microfiche 7394: 295.

Nicandra anomala Link & Otto, Icon. pl. select. 77. t. 35. July--Dec. 1825 [fide Stafleu & Cowan 3: 68]. TYPE: cultivated, Berlin, from seed obtained from England, noted as being from Nepal, specimen not known (lectotype, Link & Otto plate 35; same as Anisodus luridus). Scopolia anomala (Link & Otto) Airy Shaw, J. Bot. 75: 195. 1937.

Whitleya stramonifolia Sweet, News of Lit. & Fashion 3: 108. 13 Aug. 1825; Brit. Flower Gard. 2: 125. 1825 [1 Oct. 1825 date fide Airy Shaw, J. Bot. 85: 192--193. 1937]. Anisodus stramonifolius (Sweet) G. Don in Louden, Hort. Brit. 61. 1830. Based on Whitleya stramonifolia Sweet. Physalis stramonifera Chen Cheih & Chen Chung-lien, Acta Phytotax. Sin. 15(2): 61. 1977. orthographic variant. Anisodus stemonifolius Wu Cheng-yih & Chen Cheih, Fl. Reipubl. Popularis Sin. 67(1): 23. 1978, orthographic variant.

Anisodus fischerianus Pascher, Repert. Spec. Nov. Regni Veg. 7: 226. 1909. Anisodus luridus var. fischerianus (Pascher) C. Y. Wu & C. Chen, Acta Phytotax. Sin. 15(2): 62. 1977. TYPE: cultivated in Leningrad, seed from Sikkim--Tibet, specimen not seen.

Scopolia mairei Leveille, Bull. Geogr. Bot. 25: 37. 1915 [Ann;aaee 24 (4;dg S;aaerie)], fide Lauener, Notes Roy. Bot. Gard. Edinburgh 37: 146. 1978. Anisodus mairei (Leveille) C. Y. Wu & C. Chen, Acta Phytotax. Sin. 15(2): 64. 1977. TYPE: China. Yunnan (Yun-Nan): Ma-Kong, rocks, 2,800 m, E. E. Maire s.n. not seen.

The first description of a plant of Anisodus was of Physalis stramonifolia by Nathaniel Wallich in a compilation of his plant descriptions published by Carey early in 1824. The description of Anisodus luridus followed later in the year: late in the following year, the same species was described as Whitleya stramonifolia by Robert Sweet. One might suspect that Sweet's name was in some way based on that chosen by Wallich, but there is no evidence to substantiate this. Sweet noted that his plant was "nearest related" to Link's Anisodus, which was first cultivated in Berlin in 1821, but he did not mention Wallich's work: his description is in different terms and differs in some details from that of Wallich, for example, "peduncle short, densely villous" compared with Wallich's "peduncle ... sparingly villous."

Soon thereafter, George Don (1830) accepted Link's generic concept and transferred Sweet's name to Anisodus. Thus, the name A. stramonifolius is occupied by A. stramonifolius (Sweet) G. Don, preventing the transfer of the earlier Physalis stramonifolius Wall. into Anisodus and supplanting A. luridus Link.

We recognize Anisodus as a genus, but for those who place Anisodus in Scopolia, the correct name in Scopolia is S. stramonifolia (Wall.) Shrestha.

We saw no material of Anisodus fischerianus and place it in synonomy based on the original description, which notes a funnelform-campanulate flowering calyx. Wu & Chen (1978), in making the combination A. luridus var. fischerianus, noted leaves with 1--3 pairs of irregular coarse teeth and purple eyes inside the base of the corolla tube, but apparently they did not see the type. Pascher suspected this species to be a hybrid of A. luridus and A. tanguticus. Its provenance from Xizang (Tibet), China, and Sikkim, India, is within the range of A. lurida. Anisodus luridus is recorded from Sikkim, Bhutan, Nepal, and in China from Sichuan, Xizang, and northwestern Yunnan, occurring from 3,000 to 4,450 m.

Anisodus carniolicoides (C. Y. Wu & C. Chen) D'Arcy & Zhang Zhi-yun, comb. nov. Basionym: Scopolia carniolicoides C. Y. Wu & C. Chen, Acta Phytotax. Sin. 15(2): 59. 1977. TYPE: China. Yunnan: Deqen Xian, Baima Mountain (Pei-ma-shan), T. T. Yu 8773 (holotype, HY). Paratypes: K. M. Fang 392, T. T. Yu 9451, 11396, 13692 (all A).

Scopolia carniolicoides C. Y. Wu & C. Chen var. dentata C. Y. Wu & C. Chen, syn. nov. Acta Phytotax. Sin. 15(2): 60. 1977. TYPE: China. Sichuan: Mu-li Xian, T. T. Yu 6291 (holotype, HY not seen; isotype, A). Paratype: T. T. Yu 7180 (A).

Following our review of material from the full geographical range of the tribe Hyoscyameae---Europe to Japan and the former U.S.S.R. to India---we conclude that Scopolia carniolicoides should be placed in Anisodus and not with the European and Japanese species that have been called Scopolia. Variety dentata is based on specimens with toothed leaves, but such leaves sometimes occur in other specimens that are otherwise not different from more typical plants. It should be noted that during the period when Wu and Chen described these taxa, difficulties beyond their control prevented consultation of collections from places outside of China.

Lycianthes neesiana (Nees) D'Arcy & Zhang Zhi-yun, comb. nov. Basionym: Solanum neesianum Wall. ex Nees, Trans. Linn. Soc. London 17: 42. 1837. Solanum subtruncatum Wall. ex Dunal in DC., Prodr. 13(1): 180. 1852. Lycianthes subtruncata (Wall. ex Dunal) Bitter, Abh. Naturwiss. Vereine Bremen 25: 478. 1919 [1920]. TYPE: Wallich. Cat. supp. 2620, year 1828 or later?

Lycianthes neesiana was treated as Solanum subtruncatum in the handwritten catalog of specimens collected by Nathaniel Wallich that was prepared by Wallich and George Bentham, and as Solanum neesianum in the supplement to the catalog. The catalog and supplement are in the handwritten lists without descriptions, and they do not meet requirements for effective, much less valid, publication. Up to the present, this species has gone under the name Solanum subtruncatum, probably because this name was used first in the 1828 catalog; Solanum neesianum first appeared in the supplement, presumably a later publication. The names were not validly published until later, the first being Solanum neesianum by Nees in 1837, who attributed the name to Wallich. The name Solanum subtruncatum was not validated until 1852 by Dunal and is considered to be a synonym of L. neesiana.

This species has a calyx with an entire, truncate margin and 1--10 teeth arising below the apex, a feature placing it in Lycianthes rather than Solanum.


Solanaceae subtribe Physalidinae Miers includes about a dozen genera characterized by having accrescent calyces, longitudinally dehiscent anthers, and mostly rotate corollas. The subtribe is centered in northern Mexico (Chamaesaracha, Jaltomata, Margaranthus, Physalis, Quincula), but it is also represented in temperate Asia (Physaliastrum, Physalis) and elsewhere. Generic lines have been unclear, leading to diverse nomenclature. A series of Asian species that has been variously known in Archiphysalis, Chamaesaracha, Leucophysalis, Physaliastrum, and Physalis appear to belong to a single genus, which should be called Physaliastrum.

Physaliastrum was described by Makino (1914) with two Japanese species, P. echinatum (Yatabe) Makino and P. savatieri (Makino) Makino, which were transferred from the American genus, Chamaesaracha. In a revision of Chamaesaracha, Averett (1973) concurred in excluding these species from Chamaesaracha. Kuang & Lu (1965) revised Physaliastrum, recognizing seven species, all in Asia. Kuang (1966) described the genus Archiphysalis, basing it on one element from Japan and another from western China. These species had been known as Physalis, Physaliastrum, or Chamaesaracha. Averett (1977), extending his studies on Chamaesaracha, reviewed a series of the Chinese species hitherto placed in Physaliastrum and concluded that they and other enigmatic North American species are congeneric with Leucophysalis; he had transferred these species into Leucophysalis earlier (Averett, 1970, 1973). Following Averett's view, Grierson & Long (1978) described a new variety in Leucophysalis from Bhutan.

After studying a range of material for preparation of the Solanaceae treatment for the Flora of China and for a study of Leucophysalis grandiflora (D'Arcy et al., 1990; D'Arcy & Keating, in prep.), the type species of Leucophysalis, we are able to separate the Asian from the American groups at the generic level on the basis of aspects of their fruiting calyces. In some of the Asian species the walls of the fruiting calyx are elaborated by emergences that give the entire living fruit a bristly appearance. This is illustrated for Physaliastrum japonicum by Yasaka (1983: 80, figs. 4, 5) and by Yoshisuke (1985: 199). When dried, the emergences are sometimes difficult to see, resembling flattened, near-hyaline scales or giving the calyx wall a muricate or roughened appearance. In two other cases, which had been segregated in the genus Archiphysalis, the calyx modification is somewhat different. In one of these, Physaliastrum sinense, the ribs are greatly thickened. In the other, Physaliastrum chamaesarachoides (Makino) Makino, illustrated by Kuang & Lu (1978: 51, plate 14, fig. 7, as Archiphysalis kwangsiensis Kuang), the calyx ribs are thickened and bumpy, suggestive of incipient teeth. Such emergences or elaborations are lacking on American species of Leucophysalis and related groups, e.g., Physalis, Chamaesaracha.

The separation of Physaliastrum from Physalis rests largely on the elaboration of the calyx in Physaliastrum. Flowering material of most species of Physaliastrum appears to be scarce in herbaria and was not available for our study. However, the literature notes that in most species of Physaliastrum the corolla is lobed, while in most species of Physalis---all those from the Americas---the corolla is apically subentire. (Although Waterfall's (1958, 1967) revision of the North American species included some species with lobed corollas, these have since been removed to other genera.) Physalis alkekengi, perhaps the only species of Physalis native to the Old World, does have shallowly lobed corolla lobes, but it has no sign of calyx elaborations typical of Physaliastrum.

One of the main characters that was used to justify separation of Archiphysalis is the degree to which the berry fills the fruiting calyx. In the species that were left in Physaliastrum, the berry fills the fruiting calyx, which is appressed to the berry wall. In Archiphysalis, the fruiting calyx greatly exceeds the berry in diameter and length, and the berry is free within the bladdery calyx. In Physalis, the fruit is usually like that of Archiphysalis, but sometimes it is like that in Physaliastrum. For example, in Physalis philadelphica, which is widely marketed for food in North America and Mexico, within a single lot of fruits, some will have the berry filling the calyx and tightly appressed to it, and others will have a small berry that is free from the bladdery calyx. This variability seems to be related to degree of overall development of the fruit: larger berries tend to fill the calyx, while younger or "poorer" fruits tend to have loose calyces. Thus, the degree to which the berry fills the calyx varies within a single population or species, and because the species in Physaliastrum and Archiphysalis are closely related to Physalis, we do not think this is an appropriate character for generic separation of Archiphysalis from Physaliastrum.

In Physaliastrum (Archiphysalis) chamaesarachoides, there are no emergences on the fruiting calyx walls, but the greatly thickened calyx ribs have erect teeth, presenting a somewhat bristly appearance. In Physaliastrum (Archiphysalis) sinense, the calyx ribs are thickened, resembling those of P. chamaesarachoides, but there are no teeth. This species is more like Physalis than any of the others in Physaliastrum, but differs in the thickened calyx ribs, which we have not seen in any species of Physalis or in any species of Physalidinae from North America. We consider the Asian species to be distinct from the American species and to form the single genus Physaliastrum on the basis of the elaborations of the fruiting calyx.

The differences noted between the species from Archiphysalis and those previously part of Physaliastrum may warrant recognition of the group at the sectional level or perhaps as a subgenus.

Physaliastrum yunnanense subsp. bhutanicum (Grierson & Long) D'Arcy & Zhang Zhi-yun, comb. nov. Basionym: Leucophysalis yunnanensis subsp. bhutanica Grierson & Long, Notes Roy. Bot. Gard. Edinburgh 36: 141. 1978. TYPE: Bhutan: Rinchu-Kancham, 5,000 ft., Cooper 3943 (holotype, E not seen; isotype, BM not seen).

Although no material of this taxon was seen, the description by Grierson and Long notes "the accrescent acutely muricate calyx," which is characteristic of the Asian Physaliastrum and not the American Leucophysalis.

Physaliastrum sinense (Hemsley) D'Arcy & Zhang Zhi-yun, comb. nov. Basionym: Chamaesaracha sinensis Hemsley, J. Linn. Soc., Bot. 26: 174. 1890. Archiphysalis sinensis (Hemsley) Kuang, Acta Phytotax. Sin. 11(1): 62, pl. 8, 9, figs. 1--6. 1966. Physalis sinensis (Hemsley) Averett, Ann. Missouri Bot. Gard. 57: 380. 1970. TYPE: China. Hubei (Hupeh): Yichang (Ichang) Xian and immediate neighborhood, Henry 2902 (holotype, K not seen).

Acknowledgment. We are grateful to Armen L. Takhtajan, V.L. Komarov Botanical Institute, Leningrad, for assistance relating to the unpublished dissertation of Semenowa (1955).

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