HARVARD PAPERS IN BOTANY 3(1): 79-94. 1998.
Ihsan A. Al-Shehbaz, Yang Guang,2 Lu Lian-li, and Cheo Tai-yen3
2Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
3Herbarium, Jiangsu Institute of Botany, P.O. Box 1435, Nanjing, Jiangsu, People's Republic of China.
Abstract. The genera Cochleariella and Hilliella are reduced to synonymy of Yinshania, a genus considered to be endemic to China. The following new combinations are proposed: Yinshania acutangula ssp. microcarpa, Y. acutangula ssp. wilsonii, Y. yixianensis, Y. rivulorum, Y. sinuata, Y. sinuata ssp. qianwuensis, Y. lichuanensis, Y. rupicola, Y. rupicola ssp. shuangpaiensis, and Y. hunanensis. Descriptions and distributions are provided, and keys to the species and subspecies are given.
There has been considerable controversy with regard to the generic identity of some of the Chinese-endemic Brassicaceae (Cruciferae) that were originally placed by Schulz (1923, 1929, 1936) in Cochlearia L. sect. Hilliella O. E. Schulz. The number of taxa recognized varied between 10 and 25 species in one to four genera. A compilation of the work of Zhang (1985, 1986, 1987a, 1987b, 1996a, 1997) and Zhang and Cai (1989) shows that the 25 species and five varieties recognized in this complex were assigned to the genera Cochleariella Y. H. Zhang & R. Vogt (1 species), Hilliella (O. E. Schulz) Y. H. Zhang & H. W. Li (16 species), and Yinshania Ma & Y. Z. Zhao (9 species), whereas Ying et al. (1993) accepted 1 species in Cochleariella, 13 in Hilliella, and 9 in Yinshania. By contrast, Kuan (1987) placed 13 species in Cochlearia, Lu (1991, 1993) treated 9 in Cochlearia and 1 in Cochleariopsis Y. H. Zhang (=Cochleariella), and Zhao (1992) recognized 14 species in Yinshania and 1 in Cochlearia. The lack of agreement among these authors and the need to treat the complex for the forthcoming account of the Brassicaceae for the Flora of China necessitated a critical evaluation of the taxa to determine the number of genera and species involved. As shown below, the endemic Chinese plants of this complex of four genera (Cochlearia, Cochleariella, Hilliella, and Yinshania) belong to one genus hereafter called Yinshania.
A critical comparison of the Chinese Yinshania with the Eurasian and North American Cochlearia reveals that the genera have very little in common. Species of Cochlearia are distributed in the arctic, subarctic, and northern latitudes of the northern temperate region, and they most commonly grow in coastal habitats, mountain streamsides, sandy beaches, seashore rock crevices, gravel bars, alpine habitats, saltmarshes, saline meadows, and calcareous screes (Pobedimova 1969, 1970; Rollins, 1993; Jackson and Akeroyd, 1993). Cochlearia consists of usually glabrous and glaucous small plants with ± fleshy, simple, entire to dentate leaves, erect petals and stamens, inflated but somewhat angustiseptate (flattened at a right angle to the septum) fruits, prominently 1-veined valves, often biseriate tuberculate seeds, and accumbent or rarely incumbent cotyledons. In contrast, Yinshania is endemic to China, and the plants grow from near sea level to 1,600(--3,000) m on mountain slopes, rocky crevices, shady moist areas, dense forests, roadsides, streamsides, shady river banks, limestone areas, and wet valleys. Yinshania differs from Cochlearia in being robust, usually pubescent plants, with nonfleshy, pinnatisect or trifoliolate to pinnately compound leaves (Y. sinuata has large simple or rarely trifoliolate leaves), widely spreading sepals, petals, and stamens, terete to latiseptate (flattened parallel to septum) or rarely slightly angustiseptate fruits, inconspicuously veined valves, uniseriate or rarely biseriate and reticulate or sometimes papillate seeds, and incumbent or rarely accumbent cotyledons.
The monotypic Cochleariella Y. H. Zhang & R. Vogt was originally described as Cochleariopsis Y. H. Zhang (Zhang, 1985; Zhang and Cai, 1989), a later homonym of Cochleariopsis of Löve and Löve (1976). It rests primarily on the presence of coarse, inflated, thin-walled papillae on the fruit valves, as compared with their absence in species of Yinshania. The density of papillae varies within and among populations and ranges from being abundant to sparse or completely lacking. Plants exhibiting the various extremes of this variation were described as independent taxa. For example, those with fruit valves densely covered with large papillae are recognized as C. zhejiangensis (Y. H. Zhang) Y. H. Zhang & R. Vogt, whereas those with glabrous fruit valves are called Y. warburgii (O. E. Schulz) Y. Z. Zhao, and plants with sparse, slender valve papillae are treated as Y. fumarioides (Dunn) Y. Z. Zhao. Fruits of the Y. warburgii holotype are immature, but they show a clear similarity to immature fruits of C. zhejiangensis and Y. fumarioides in the developmental stages of papillae. The types of these three "species" are basically indistinguishable in other morphological characters. Lu (1993) was correct in reducing C. zhejiangensis to synonymy of Y. warburgii, but she maintained the species in Cochleariopsis and did not associate it with Y. fumarioides. In our opinion, Cochleariella does not merit recognition.
Cochlearia L. sect. Hilliella was raised to the generic rank by Zhang and Li (Zhang, 1986), and it was said to differ from Yinshania in having compound instead of pinnatisect leaves, simple instead of forked trichomes, eseptate instead of septate fruits, and papillate instead of reticulate seeds. Only 2 of the 11 Yinshania species recognized by Zhang (1996a) have branched trichomes, and reticulate seeds occur in both Hilliella and Yinshania (Zhang, 1996a; Zhao, 1991). As for the presence versus absence of septa, this feature is not useful at the generic rank, and septate and eseptate fruits occur within numerous genera of the Brassicaceae. Some of the species described or maintained by Zhang (1986, 1987b) in Hilliella have fruits with complete septa. Finally, both compound and pinnatisect leaves are found within Yinshania, and as delimited by Zhang (1986), Hilliella included species with simple or compound leaves. The genus Cardamine L. includes species with simple, pinnatisect, trifoliolate, or pinnately compound leaves. As shown by Al-Shehbaz and Yang (1998), two of the eight species recognized in Hilliella by Zhang (1986) are members of the genus Cardamine. We fully agree with Zhao (1992) and Lu (1991) that the alleged differences between Hilliella and Yinshania are inconsistent and that a single genus should be recognized.
As delimited here, Yinshania consists of 13 species and four subspecies. The genus is endemic to China and is distributed in 17 of the 28 provinces and autonomous regions of China. These include Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hebei, Hubei, Hunan, Jiangxi, Nei Mongol, Shaanxi, Sichuan, Taiwan, Xizang, Yunnan, and Zhejiang.
INFRAGENERIC CLASSIFICATION OF YINSHANIA
Zhang (1987a) divided Yinshania (excluding Hillliella, which she recognized as a distinct genus), into two sections solely on the basis of differences in fruit shape. Section Microcarpa Y. H. Zhang was characterized by having subglobose to broadly ovoid fruits 1.0--2.2 mm long, whereas sect. Yinshania was said to have oblong-ovoid, oblong, to lanceolate-ellipsoid fruits 1.5--4.5 mm long. However, these alleged differences represent continuous variation, and they do not justify the division of the genus into sections. As shown below, in Y. acutangula and Y. zayuensis, fruit shape sometimes varies on the same plant, depending on the number of ovules that mature into seeds. Zhang (1987a) divided sect. Yinshania into two series: ser. Henryanae Y. H. Zhang (infructescence rachis flexuous, trichomes simple or branched, dense, leaves 3- to 5-foliolate or pinnatisect to pinnatipartite) and ser. Yinshania (infructescence rachis straight, trichomes simple or branched, sparse or absent, leaves pinnatisect to pinnatipartite). We believe that these differences are also unreliable, and in Y. zayuensis the infructescence rachis can be straight or flexuous.
Zhao (1992) maintained Y. sinuata as a species of Cochlearia and divided Yinshania into two sections: sect. Yinshania (leaves pinnatisect or pinnatipartite) and sect. Hilliella (leaves all or lowermost compound). He followed Zhang (1987a) in using fruit shape to divide sect. Yinshania into ser. Microcapae and ser. Yinshaniae. Zhao divided sect. Hilliella into four series on the basis of differences in leaflet shape and the presence versus absence or type of trichomes. However, these subdivisions are artificial, and his monotypic series Alatipes included a species of Cardamine (see Al-Shehbaz and Yang, 1998).
In our opinion, the infrageneric subdivisions above are neither practical nor useful, and they do not reflect the phylogenetic relationships in Yinshania. Unless the infrageneric subdivisions meet these criteria, there is no need for them in the Brassicaceae, especially in small genera.
TAXONOMIC TREATMENT OF YINSHANIA
Yinshania Y. C. Ma & Y. Z. Zhao, Acta Phytotax. Sin. 17(3): 113. 1979.
Synonyms: Cochleariella Y. H. Zhang & R. Vogt, Acta Bot. Bor.-Occid. Sin. 9(4): 224. 1989. Cochleariopsis Y. H. Zhang, Acta Bot. Yunnan. 7(2): 143. 1985; not Löve & Löve, Bot. Notiser 128: 513. 1976. TYPE: C. zhejiangensis (Y. H. Zhang) Y. H. Zhang & R. Vogt.
Hilliella (O. E. Schulz) Y. H. Zhang & H. W. Li, Acta Bot. Yunnan. 8: 401. 1986; Cochlearia L. sect. Hilliella O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem, 8: 544. 1923. TYPE: H. fumarioides (Dunn) Y. H. Zhang & H. W. Li.
Herbs annual or perennial with elongated or rarely tuberous rhizomes, glabrous or sparsely to densely hairy; trichomes simple, furcate, or bifurcate. Stems erect or decumbent, terete to conspicuously angled and striate, usually hollow. Leaves petiolate, pinnately lobed, trifoliolate, or pinnately compound, rarely simple. Inflorescences corymbose racemes, elongated in fruit, ebracteate or rarely lower half bracteate; rachis straight or rarely flexuous. Sepals oblong, spreading, not saccate at base, margins hyaline. Petals white or rarely pinkish, spreading, not differentiated into distinct claw and blade, base cuneate, apex rounded. Lateral nectar glands in pairs, median ones absent. Filaments white, spreading; anthers ovate to oblong. Fruit dehiscent, globose, ovoid, oblong, or elliptic, terete to somewhat latiseptate, rarely slightly angustiseptate; valves convex, veinless or with obscure midvein, glabrous or papillate or pubescent. Style distinct; stigma entire. Seeds several or rarely few or 1 in each locule, 1- or 2-seriate, slightly flattened, reticulate to minutely papillate; cotyledons incumbent or rarely accumbent.
TYPE: Yinshania albiflora Y. C. Ma & Y. Z. Zhao (=Y. acutangula [O. E. Schulz] Y. H. Zhang). When describing Yinshania, Ma and Zhao (1979) recognized a single species, Y. albiflora. Although Zhang (1987a, 1989) reduced Y. albiflora to the synonymy of Y. acutangula and thus effectively indicated that the latter is the generic type, it is rather strange that Zhao (1992) still recognizes Y. albiflora as the valid generic type and lists the much earlier published C. acutangula (Schulz, 1929) and Y. acutangula (Zhang, 1987a) as synonyms.
KEY TO THE SPECIES OF YINSHANIA
1a. Most or all leaves simple, entire, repand, or sinuate, rarely lowermost few trifoliolate ... Y. sinuata
1b. Leaves compound or pinnately lobed 2
2a. Flowers bracteate along entire length or only on lower half of inflorescence.
3a. Plants annual; ovary ca. 8-ovuled, cylindric, papillate ... Y. hui
3b. Plants perennial; ovary ca. 4-ovuled, obovate-ellipsoid, glabrous ... Y. hunanensis
2b. Flowers ebracteate ... 4
4a. Trichomes bifurcate ... 5
5a. Leaves and stems canescent, densely villous with trichomes 0.25--0.5(--0.6) mm long ... Y. furcatipilosa
5b. Leaves and stems not canescent, puberulent with trichomes 0.05--0.08(--0.1) mm long ... Y. zayuensis
4b. Trichomes simple or absent ... 6
6a. Infructescence rachis conspicuously flexuous ... 7
7a. Leaves canescent, densely villous, margin of lower leaves deeply crenate; seeds 0.7--0.9 mm long, (2--)5--10 per locule ... Y. henryi
7b. Leaves not canescent, glabrous to sparsely pubescent, margin of lower leaves not deeply crenate; seeds 1.5--2.0 mm long, 1 (or 2) per locule ... Y. yixianensis
6b. Infructescence rachis straight ... 8
8a. Leaflets of middle and lower leaves less than 1 cm wide ... 9
9a. Leaves glabrous, lower ones trifoliolate; fruit papillate ... Y. fumarioides
9b. Leaves usually pilose, rarely glabrous, lower ones pinnate, with 2--5 pairs of lateral leaflets; fruit glabrous or rarely pilose ... Y. acutangula
8b. Leaflets of middle and lower leaves more than 1 cm wide ... 10
10a. Lower leaves with 5--9 leaflets; fruit conspicuously compressed; plants tuberous perennials ...Y. rupicola
10b. Lower leaves with 3 (or 5) leaflets; fruit plump, not compressed; plants annuals with slender roots ... 11
11a. Leaflet apex acuminate-caudate; fruit appressed to rachis; plants puberulent with trichomes to 0.1 mm long ... Y. lichuanensis
11b. Leaflet apex obtuse to acute, rarely subacuminate; fruit widely spreading to reflexed; plants glabrous throughout or rarely glabrescent ... 12
12a. Fruit ovate to obovate, 2- or 3-seeded; style 0.5--0.8 mm; pedicel (1.5--)2.5--4.0 mm long ... Y. paradoxa
12b. Fruit narrowly oblong, 7- to 10-seeded; style (1.0--)1.5--2.0 mm; pedicel 6--10 mm long ... Y. rivulorum
Yinshania acutangula (O. E. Schulz) Y. H. Zhang, Acta Phytotax. Sin. 25: 217. 1987.
Annuals, (8--)30--60(--100) cm tall. Stems slender, distinctly angled to subterete, branched, sparsely to densely covered with simple trichomes (0.1--)0.3--0.4(--0.5) mm long, rarely glabrescent or glabrous. Basal leaves and lowermost stem leaves pinnate, with 2--5 pairs of lateral leaflets, often withered at fruiting, pilose with subappressed simple straight trichomes; petiole 0.5--2.0 cm long; leaflets membranous, oblong to ovate-lanceolate, 5--20 x 2--9 mm, sparsely pubescent or rarely glabrous, base cuneate, margins deeply serrate to entire, apex obtuse, with a minute callose mucro; upper-stem leaves similar to lower ones but progressively shorter upward. Racemes terminal and lateral, many flowered, elongated in fruit. Fruiting pedicel 4--7 mm long, slender, straight, divaricate or slightly recurved, glabrous or pilose. Sepals ovate, 1.0--1.5 mm long. Petals white, obovate, 2.0--2.5 x ca. 0.7 mm. Filaments ca. 1 mm long. Fruit oblong-linear, oblong, to narrowly ovoid or globose, 1.5--4.0 x 1.0--1.5 mm, not compressed, sessile; valves membranous, papillate or pilose. Style 0.5--0.8 mm long. Seeds (1--)2--10(--12) per locule, brown, oblong-ovate, 0.6--1.0(--1.1) x 0.4--0.5(--0.6) mm, reticulate.
From the other members of Yinshania, Y. acutangula is readily distinguished by having angled stems with subappressed, simple straight trichomes, ebracteate inflorescences with straight rachis, and pinnate leaves with leaflets less than 1 cm wide. It is quite variable in the degree of development of stem angles, size of leaf divisions, density of flowers on the inflorescence branches, fruit shape and indumentum, and number of seeds per locule. The variation, however, is rather continuous, and only the following three subspecies can be consistently recognized on the basis of the characters in the key below.
KEY TO THE SUBSPECIES OF Y. ACUTANGULA
1a. Fruit oblong to oblong-linear, rarely narrowly ovoid; seeds (5--)7--10(--12) per locule, 0.6--0.8 mm long ... ssp. acutangula
1b. Fruit globose, rarely globose-ovoid; seeds 1 or 2 (or 3) per locule, 0.8--1.0(--1.1) mm long ... 2
2a. Fruit pilose; stems obscurely angled to subterete ... ssp. microcarpa
2b. Fruit minutely papillose to glabrous; stems distinctly angled ... ssp. wilsonii
Yinshania acutangula ssp. acutangula
Basionym: Cochlearia acutangula O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 10: 554. 1929. TYPE: CHINA. Shaanxi [as Nord-Shensi]: Miao-nan-san, Pao-ki-seen, October 1898, G. Giraldi 2423 (holotype, B).
Synonyms: Yinshania albiflora Y. C. Ma & Y. Z. Zhao, Acta Phytotax. Sin. 17(3): 114. 1979. TYPE: CHINA. Nei Mongol, Daqing shan, Y. Z. Zhao 155 (holotype, HIMC). Yinshania acutangula (O. E. Schulz) Y. H. Zhang var. albiflora (Y. C. Ma & Y. Z. Zhao) Y. H. Zhang, Acta Phytotax. Sin. 25: 217. 1987.
Yinshania wenxianensis Y. H. Zhang, Acta Phytotax. Sin. 25: 215. 1987. TYPE: CHINA. Gansu: Wen Xian, Tielou, 1400 m, 7 November 1958, Y. Q. He 989 (holotype, PE!; isotype, PE!).
Yinshania wenxianensis Y. H. Zhang var. songpanensis Y. H. Zhang, Acta Phytotax. Sin. 25: 212. 1987. TYPE: CHINA. Sichuan: Songpan, Erdaoqiao, 1800 m, 22 October 1937, K. T. Fu 2177 (holotype, PE!; isotype, WUG).
Stems distinctly angled. Fruit oblong to oblong-linear, rarely narrowly ovoid, glabrous or minutely papillose. Seeds (5--)7--10(--12) per locule, 0.6--0.8 mm long.
Habitat and phenology: Mountain slopes, meadows, rock crevices, valleys, fields, among bushes, roadsides; 900--3000 m. Flowering July to August, fruiting August through October.
Distribution: Gansu, Hebei, Nei Mongol, Shaanxi, Sichuan.
Additional specimens examined: CHINA. Gansu: Ma-yu Shan, Yuchou Xian, Hou 6121 (PE); Lanzhou, Pongchen Zucha Gou, Hou 5242 (PE); Daqing Shan, Zhaoer Gou, 30 August 1974, Yin s.n. (PE). HEBEI: Wu-An County, He 21252 (PE); same county, 12 September 1979, Hu s.n. (PE). NEI MONGOL: Sartchy, David 2889 (P).
Zhao (1992) has correctly recognized Y. wenxianensis and Y. acutangula as conspecific, but he wrongly adopted the much-later-published Y. albiflora for the combined species.
The alleged differences listed by Zhang (1987a) to distinguish between the two varieties of Y. wenxianensis (i.e., distribution and density of indumentum, infructescence length, and trichomes length) show a great deal of overlap, and the distinction is unrealistic. Furthermore, the slight differences in fruit length and seed number between the two varieties of Y. wenxianensis on the one hand and var. acutangula on the other are unreliable, and therefore they are reduced here to synonymy of var. acutangula.
Yinshania acutangula ssp. microcarpa (K. C. Kuan) Al-Shehbaz, G. Yang, L. L. Lu, & T. Y. Cheo, comb. et stat. nov.
Basionym: Cochlearia microcarpa K. C. Kuan, Bull. Bot. Lab. N. E. Forest. Inst., Harbin 1980(8): 40. 1980. TYPE: CHINA. Sichuan: Gulin, 1100 m, 25 August 1976, Gulin Expedition 943 (holotype, PE!; isotype, SM).
Synonym: Yinshania microcarpa (K. C. Kuan) Y. H. Zhang, Acta Phytotax. Sin. 25: 211. 1987.
Stems obscurely angled to subterete. Fruit globose, pilose. Seeds 1 or 2 (or 3) per locule, 0.8--1.0(--1.1) mm long.
Habitat and phenology: dry areas by rocks; ca. 1100 m. Fruiting August.
Distribution: Sichuan. Known only from the type collection.
The presence of pilose instead of minutely papillose fruits in this subspecies is significant and might justify its recognition as an independent species. However, all other characters of the plant readily fall within the variation range of Y. acutangula ssp. wilsonii, and glabrous and pubescent fruits are found in other species, such as Y. henryi. Both Zhang (1987a) and Zhao (1992) recognize Y. microcarpa as a species distinct from Y. acutangula, whereas Lu (1991) did not include it in her account. In fact, Zhang (1987a) placed Y. acutangula and Y. microcarpa in independent sections separated mainly on the basis of differences in fruit shape.
Yinshania acutangula ssp. wilsonii (O. E. Schulz) Al-Shehbaz, G. Yang, L. L. Lu, & T. Y. Cheo, comb. et stat. nov.
Basionym: Cochlearia henryi (Oliver) O. E. Schulz var. wilsonii O. E. Schulz, Repert. Spec. Nov. Regni Veg. 38: 108. 1935. TYPE: CHINA. Sichuan: 4500 m, E. H. Wilson 3210 (holotype, BM!; isotypes, A!, K!, P!).
Synonym: Yinshania qianningensis Y. H. Zhang, Acta Phytotax. Sin. 25: 212. 1987. TYPE: CHINA. Sichuan: Qianning, 3000 m, 14 August 1974, collector unknown (holotype, PE!; isotype, CDBI).
Yinshania qianningensis Y. H. Zhang var. brachybotrys Y. H. Zhang, Acta Phytotax. Sin. 25: 212. 1987. TYPE: CHINA. Gansu: Wen Xian, Caoheba, 1600 m, 10 July 1973, J. X. Yang 3373 (holotype, PE!; isotypes, IBSC!, WUG).
Stems distinctly angled. Fruit globose, rarely globose-ovoid, minutely papillose to glabrous. Seeds 1 or 2 (or 3) per locule, 0.8--1.0(--1.1) mm long.
Habitat and phenology: valleys, roadsides; 1400--3000 m. Flowering July to August, fruiting August through October.
Distribution: Gansu, Sichuan.
Additional specimen examined: CHINA. Sichuan: without a locality, Wilson 3210a (A, P).
Although Schulz (1935) cited two collections (Wilson 3210 and Wilson 3210a) in his original description of C. henryi var. wilsonii, the single specimen at BM (Wilson 3210), which Schulz cited and which is annotated in Schulz's handwriting, should be considered the type. Furthermore, Schulz gave detailed descriptions of the fruit and seeds that were observed in Wilson 3210. Plants of Wilson 3210a have flower buds and no fruits.
It is difficult to understand why Schulz (1935) described this taxon as a variety of C. henryi (here Yinshania) instead of his C. acutangula. Yinshania henryi is readily distinguished from the three subspecies of Y. acutangula in having distinctly flexuous infructescence axes and canescent leaves densely with trichomes (0.5--)0.6--1.0 mm long, whereas Y. acutangula has straight infructescence axes and green leaves glabrous or sparsely with trichomes (0.1--)0.3--0.4(--0.5) mm long.
The presence of short (vs. long) inflorescence branches is the sole feature that distinguishes var. barchybotrys from var. qianningensis. The type collection of var. wilsonii above includes both inflorescence types. The degree of elongation of the lateral inflorescence branches does not appear to be taxonomically useful.
Zhao (1992) recognized Y. qianningensis as an independent species, but the overall morphology clearly supports the recognition of a single taxon here treated as ssp. acutangula.
Yinshania yixianensis (Y. H. Zhang) Al-Shehbaz, G. Yang, L. L. Lu, & T. Y. Cheo, comb. nov.
Basionym: Hilliella yixianensis Y. H. Zhang, Acta Phytotax. Sin. 33: 94. 1995. TYPE: CHINA. Anhui: Yixian County, Hedongcun, 250 m, July 1991, S. X. Qian, s. n. (holotype, PE).
Annuals or biennials. Stems erect, 40--90 cm tall, striate, glabrous or rarely pilose with simple trichomes. Basal to median stem leaves compound, with 2 or 3 pairs of lateral leaflets; petiole 0.5--1.5 cm long; leaflets thin papery, ovate to elliptic, 1--4 cm long, 0.8--1.7 cm wide, hispidulous, base obtuse to cuneate, margin obtusely dentate with the teeth mucronate, apex acute; petiolule to 5 mm; uppermost stem leaves smaller, with fewer teeth and lateral leaflets. Inflorescences terminal and lateral racemes, elongated considerably in fruit; rachis conspicuously flexuous, slender. Fruiting pedicels slender, 5--6 mm long. Sepals elliptic, ca. 1.5 x 0.5 mm, spreading. Petals white, spreading, ovate, ca. 2.5 x 1.5 mm. Fruit subglobose-obovoid, ca. 3.5 x 2.5 mm; valves papillose. Seeds 1 (or 2) per locule, ovoid, 1.5--2.0 mm long.
Habitat and phenology: Roadsides; ca. 250 m. Fruiting in July.
Distribution: Anhui. Known only from the type collection, which was grown from seeds.
Although we have not seen the type of this interesting species, the detailed protologue and photograph clearly support its close relationship to Y. acutangula. Zhang (1995a) compared and associated Y. yixianensis (as Hilliella) with H. paradoxa, but the latter has trifoliolate instead of pinnate leaves. The flexuous lateral branches of the inflorescence of Y. yixianensis are a remarkable feature that readily distinguishes it from most species of the genus. Except for the unrelated Y. henryi and Y. furcatipilosa, both of which have canescent leaves, conspicuously flexuous infrutescences are not found elsewhere in the genus.
Yinshania zayuensis Y. H. Zhang, Acta Phytotax. Sin. 25: 214. 1987. TYPE: CHINA. Xizang (Tibet) [as Sikang]: Chawalong [as Tsa-wa-rung], 2800 m, August 1935, C. W. Wang 65236 (holotype, PE!; isotypes, A!, IBSC!, PE!, WUG).
Synonyms: Yinshania exiensis Y. H. Zhang, Acta Bot. Yunnan. 15: 363. 1993. Syn. nov. TYPE: CHINA. Hubei: Shennongjia, Xinhua, Longkou, 1000 m, 2 July 1976, Shennongjia Expedition 20597 (holotype, HIB; listed as WUBI).
Yinshania ganluoensis Y. H. Zhang, Acta Phytotax. Sin. 25: 211. 1987. Syn. nov. TYPE: CHINA. Sichuan: Ganluo, Suxiong, 800 m, 26 June 1979, collector unknown 556 (holotype, MS; isotype, MS).
Annuals, (14--)30--60 cm tall. Stems erect, simple or branched, puberulent with forked and bifurcate trichomes 0.05--0.08(--0.1) mm long, rarely glabrous above. Leaves pinnatisect; petiole 2--7 mm; leaf blade oblong, 0.6--3.0 x 0.4--1.5(--2.3) cm, puberulent with minute sturdy simple and forked trichomes, rarely glabrescent; terminal lobe oblong, 3--15 x 1--5 mm, base cuneate, margins serrate to subentire, apex acute; lateral lobes 1--3 pairs, 1.5--13.0 x 0.5--3.0 mm. Inflorescences paniculate; rachis straight or rarely flexuous. Pedicel divaricate to ascending, slender, straight, 5--10 mm long, pubescent or glabrous. Sepals oblong, 1.0--1.6 x 0.5--0.7 mm, white at margins and apex. Petals white or rarely pinkish, 1.5--2.0 x 0.7--1.0 mm. Filaments 1.1--1.2 mm long; anthers oblong, ca. 0.2 mm long. Fruit oblong to ellipsoid or ovoid, plump, (2.0--)2.5--4.0 x 0.9--1.6 mm; valves with distinct midvein and reticulate veins, glabrous. Style 0.1--0.5 mm long. Seeds deep brown, ovate, 0.6--0.9 x 0.4--0.6 mm, reticulate.
Habitat and phenology: woods; 2600--3000 m. Flowering and fruiting July through August.
Distribution: Hubei, Sichuan, Xizang, Yunnan.
Additional specimens examined: CHINA. Hubei: Shennong-Jia, August 1976, C. Y. Yang, s.n. (PE). Yunnan: Deiqin Xian, Chuan Jiang, Jinshu, Zihou Shan, K. M. Feng 6329 (KUN, PE); Deiqin Xian, Yunlin, Gongshei, Xilang, Yang 8717 (KUN); Dokerla, A-tun-tze, Wang 65136 (A, KUN, PE); Zhixi, Yegue, Molidayren, Baiguolu, Yang 6972 (KUN); Mekong Valley, 28°12' N, Forrest 13955 (P).
Although we have been unable to examine the types of Y. exiensis and Y. ganluoensis, the descriptions and illustration provided by Zhang (1987a, 1993, 1996a) clearly show that these "species" are indistinguishable from Y. zayuensis. The presence of minutely puberulent furcate trichomes on the leaves and stems is not found in any other species of Yinshania. The three "species" are very similar in all other characters, except for slight differences in fruit shape. Zhang (1993) placed Y. zayuensis and Y. ganluoensis in two different sections of Yinshania primarily on the basis of fruit shape (subglobose to broadly ovoid in sect. Microcarpa vs. ovoid to oblong in sect. Yinshania). However, it is abundantly clear from the morphology of the two plants that they are conspecific and that the division of the genus into sections on the basis of slight differences in fruit shape is unwarranted.
Yinshania zayuensis was misidentified and illustrated in Flora Reipublicae Popularis Sinicae (An, 1987) as Camelina yunnanensis W. W. Smith, but as indicated below, the latter is a synonym of Rorippa globosa (Turczaninow) Thellung.
One collection, Forrest 13955, was listed as Sisymbrium sonnei Robinson (Notes Roy. Bot. Gard. Edinburgh 17: 31. 1929), and the label carries the name Mekongia yunnanensis W. W. Smith. However, neither of these two names has been validly published.
Yinshania fumarioides (Dunn) Y. Z. Zhao, Acta Sci. Nat. Univ. Intramongol. 23: 568. 1992.
Basionym: Cochlearia fumarioides Dunn, J. Linn. Soc., Bot. 38: 355. 1908. TYPE: CHINA. Fujian: near Buong Kang, S. T. Dunn 2360 (holotype, HK; isotypes, GH!, K!).
Synonyms: Hilliella fumarioides (Dunn) Y. H. Zhang & H. W. Li, Acta Bot. Yunnan. 8: 403. 1986.
Cochlearia warburgii O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 8: 545. 1923. TYPE: CHINA. Zhejiang [as Tsche Kiang]: Ningpo, May 1887, Warburg 6340 (holotype, B!). Cochleariopsis warburgii (O. E. Schulz) L. L. Lu, Acta Phytotax. Sin. 31: 287. 1993; Hilliella warburgii (O. E. Schulz) Y. H. Zhang & H. W. Li, Acta Bot. Yunnan. 8: 405. 1986; Y. warburgii (O. E. Schulz) Y. Z. Zhao, Acta Sci. Nat. Univ. Intramongol. 23: 568. 1992.
Cochleariopsis zhejiangensis Y. H. Zhang, Acta Bot. Yunnan. 7(2): 144. 1985. TYPE: CHINA. Zhejiang: Suichang, Jiulong Shan, 720 m, C. Ling 3154 (holotype, ZJMA [as ZMDB]; isotype, HZU [as HUH]). Cochleariella zhejiangensis (Y. H. Zhang) Y. H. Zhang & R. Vogt, Acta Bot. Bor.-Occid. Sin. 9: 224. 1989; Yinshania zhejiangensis (Y. H. Zhang) Y. Z. Zhao, Acta Sci. Nat. Univ. Intramongol. 23: 569. 1992.
Hilliella warburgii (O. E. Schulz) Y. H. Zhang & H. W. Li var. albiflora S. X. Qian, Bull. Bot. Res., Harbin 10(4): 63. 1990. TYPE: CHINA. Zhejiang: Tiantai Xian, Tiantaishan, Huading, c. 950 m [as cm], 24 May 1987, S. X. Qian 10002 (holotype, Medical Research Institute, Shanghai Railway Medical College).
Annuals, (5--)10--20(--35) cm tall, glabrous throughout except for papillate fruit valves. Stems slender, branched below or above. Basal and lowermost stem leaves 3-foliolate; petiole 0.2--3.0 cm long, not winged; terminal leaflet blade ovate or suborbicular, 0.7--1.5(--2.0) x 0.4--1.0 cm, base cuneate to subcordate, margin lobed, without apiculate callosities, apex truncate to emarginate; lateral leaflet blade obliquely ovate, 2.3--11.0 x 1.2--8.0 mm, margin lobed to crenate; uppermost leaves simple, gradually reduced upward. Racemes several- to many-flowered, ebracteate; rachis slender, only somewhat flexuous. Pedicel slender, horizontal or slightly reflexed, (3--)6--10 mm long in fruit, straight or slightly recurved, glabrous. Sepals oblong, 0.9--1.2 x 0.5--0.6 mm, glabrous. Petals white, obovate, 1.8--2.1 x 0.9--1.3 mm, base short clawed, apex rounded. Filaments 0.7--0.9(--1.3) mm long; anthers oblong-ovate, 0.2--0.4 mm long. Fruit suborbicular, 1.5--2.5 mm in diam., plump; valves densely papillate, appearing minutely papillate or subglabrous when young. Style 0.2--0.5 mm long. Seed 1 per locule, brown, broadly ovate to ovate-oblong, 1.2--1.5 x 0.8--1.1 mm, minutely papillate.
Habitat and phenology: Wet slopes, rocky cliffs, forests, along streams; 400--1000 m. Flowering and fruiting May through August.
Distribution: Fujian, Zhejiang.
Additional specimens examined: CHINA. Zhejiang: Dongyang Co., 25 July 1972, Zhong. s.n., (PE); Putuo, Taohua district, 17 May 1972, no collector name 6137 (PE); Longquan, Maoshan, S. Y. Zhang 2823 (PE); near Siachu, R. C. Ching 1674 (US); Rueyen, no collector name 4271 (PE); Leqin, Yan Dang Shan, Hangzhou Botanic Garden 2376 (MO); Shuechan, Xiaxi Ken, Deng Wei 82108 (NAS). Province not known: Henshan, Hajacava 744 (TI).
Fruit indumentum is the most variable feature in Y. fumarioides, and this variation was the principal reason for recognizing three species variously placed in one to four genera. As shown above, Cochleariella is indistinguishable from Yinshania in every feature, and Zhao (1992) was the first to point that out. However, he recognized Y. fumarioides, Y. warburgii, and Y. zhejiangensis as independent species. Lu (1993), on the other hand, was the first to reduce the last species to synonymy of Y. warburgii, but she maintained Cochleariella as independent from the remaining species that she placed in Cochlearia. Furthermore, Lu (1991) retained Y. fumarioides in Cochlearia. By contrast, Zhang (1995b) maintained Y. zhejiangensis in the monotypic Cochleariella, and she recognized Y. warburgii and Y. fumarioides in Hilliella (1987a, 1995b). The failure of these authors to associate Y. warburgii and Y. zhejiangensis with the earlier-published Y. fumarioides probably resulted from a failure to examine the type material of the last species.
Yinshania henryi (Oliver) Y. H. Zhang, Acta Phytotax. Sin. 25: 213. 1987.
Basionym: Nasturtium henryi Oliver, Hooker's Icon. Pl. 18: t. 1719. 1887; Cochlearia henryi (Oliver) O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 8: 546. 1923. TYPE: CHINA: [Hubei]: Ichang, A. Henry 2899 (holotype, K!; isotypes, K!, TI!).
Synonym: Nasturtium kouytchense H. Léveillé, Bull. Soc. Agr. Sci. Art. Sarthe 31: 321. 1904. Syn. nov. TYPE: CHINA. Guizhou [as Kouy-Tchéou]: environ de Gan-pin, 20 September 1897, L. Martin & E. Bodinier 1804 (holotype, E!; isotype, P!).
Annuals, 15--35(--50) cm tall. Stems branched, terete. Leaves 3- to 5(--9)-foliolate, canescent, usually densely pubescent with straight simple trichomes (0.5--)0.6--1.0 mm long; petiole 1.5--6.0 cm long; terminal leaflet blade broadly ovate to suborbicular, rarely narrowly ovate, 1.5--2.5 x 0.8--2.8 cm, base slightly cordate to subtruncate, margins 3-lobed, lobes deeply crenate, ending in a small callose mucro, apex rounded; lateral leaflets petiolate to sessile; leaflet blade 6--10 mm long, base attenuate. Racemes ebracteate, elongated in fruit; rachis slender, slightly to strongly flexuous, at least apically. Pedicel slender, straight, spreading, 5--10(--15) mm long in fruit, pubescent or rarely glabrous. Sepals oblong, 1.5--2.0 x 0.7--1.2 mm, pubescent outside. Petals white, obovate, 2.0--3.5 x 1.0--1.5 mm, base short clawed, apex rounded. Filaments 1.3--2.0 mm long; anthers ovate, ca. 0.3 mm long. Fruit narrowly oblong to oblong-lanceolate or sublinear, (2.5--)3.0--5.0 x 1.0--1.2 cm, plump; valves navicular, pubescent or sometimes glabrous, thickened at margin; septum complete or perforate. Style 0.3--0.7(--1.2) mm. Seeds (2--)5--10 in each locule, reticulate, brown, ovate, 0.7--0.9 x 0.5--0.7 mm.
Habitat and phenology: mountain slopes, rocky areas, valleys; 800--1200 m. Flowering and fruiting June through September.
Distribution: Guizhou, Hubei, Sichuan, Yunnan.
Additional specimens examined: CHINA. Guizhou: cave, Ganping, Esquirol 4348 (K). Hubei: Ichang, Henry 4125 (B, E, GH, K, NY, P, US); western Hubei, Wilson 1960 (K, NY, W); Canqianhe Gou, Li 9409 (PE); Fang County, Jiang & Tao 383 (E, MO). Sichuan: Shanhue, Honghuo Gou, Xong & Zhou 91832 (IBSC, PE); Tchen-kéou-tin, Farges 1045 (US). Yunnan: Yunnan-sen district, Cavalerie 690 (E, K).
In his original description of Nasturtium henryi, Oliver (1887) cited a single collection (Henry 2899) and indicated that "this singular plant, which for the present may be left in Nasturtium, may prove ultimately better disposed of in Lepidineae as a distinct generic type." Another collection (Henry 4125, see below), which was made in 1888 presumably from the same general area of the type locality, has been wrongly annotated by various botanists as a syntype, but this collection was not cited in the original publication and therefore has nothing to do with the type collection. It is interesting that Oliver's suggestion that N. henryi belongs to a distinct genus was overlooked until Ma and Zhao (1979) described Yinshania more than a century later.
Lu (1991) did not recognize Y. henryi nor list it in the synonymy of any species. The species can easily be distinguished from other members of Yinshania in having canescent, deeply crenate leaves with simple trichomes (0.5--)0.6--1.0 mm long and flexuous infructescences.
Yinshania henryi is most closely related to Y. furcatipilosa, which it resembles in having flexuous infructescences, canescent leaves, and long trichomes ca. 1 mm long. The latter has bifurcate trichomes, whereas Y. henryi has simple ones. The differences in leaf shape and petal length listed by Ying et al. (1993) to distinguish between Y. henryi and Y. fumarioides are unreliable, and both species have mostly ovate leaflets and petals over 3 mm long.
Yinshania furcatipilosa (K. C. Kuan) Y. H. Zhang, Acta Phytotax. Sin. 25: 214. 1987.
Basionym: Cochlearia furcatopilosa K. C. Kuan, Bull. Bot. Lab. N. E. Forest. Inst., Harbin 1980(8): 41. 1980. TYPE: CHINA. Hubei: Shennong-Jia, 5 July 1976, 1500 m, Shennong-Jia Expedition 21332 (holotype, PE!; isotype, PE!).
Annuals, 10--20(--30) cm tall, canescent, densely covered with forked trichomes. Stems erect, branched, terete. Basal and lower stem leaves 3- to 5-foliolate, canescent, densely pubescent with bifurcate trichomes 0.25--0.5(--0.6) mm long; petiole 1.0--3.0 (--4.5) cm long; terminal leaflet blade ovate to rarely oblong or subreniform, 0.6--1.2 x 0.4--1.5 cm, base subtruncate to rounded, margins lobed to crenate, not ending in callose apiculae, apex rounded; lateral leaflet blade suborbicular to ovate, 3--6 x 2--7 mm, margins crenate to entire. Uppermost leaves simple, pubescent; petiole 1--6 mm long; leaf blade oblong, 3--10 x 1.5--4.5 mm, base rounded, margins entire, apex obtuse. Inflorescences paniculate, ebracteate; rachis distinctly flexuous. Pedicel divaricate, straight, slender, (3--)5--9(--12) mm in fruit, glabrous or pubescent at base. Sepals oblong, 1.2--1.6 x 0.8--0.9 mm, glabrous, white at margins and apex. Petals white, obovate, 3.0--3.3 x 1.6--2.0 mm, base short clawed, apex rounded. Filaments 1.0--1.6 mm long; anthers oblong-ovate, ca. 0.3 mm long. Fruit ovate or obovate, 1.7--2.2 x 0.8--1.1 mm; valves without veins, glabrous; septum perforate. Style 0.4--0.5 mm long. Seeds brown, oblong, 0.7--0.8 x 0.3--0.4 mm, finely papillate, narrowly margined.
Habitat and phenology: mountain slopes, rocky areas, roadsides; 800--1600 m. Flowering and fruiting June through July.
Additional specimens examined: CHINA. Hubei: Shengrong-Jia, Dayiewu, Taoyuan, Houzhi 2598 (PE); Shennong-Jia; Shennong-Jia Expedition 21828 (PE).
This is the most distinctive species of the genus because of its canescent leaves densely covered with bifurcate trichomes 0.25--0.5(--0.6) mm long and its distinctly flexuous inflorescences. It is most closely related to Y. henryi, from which it is readily distinguished in having furcate instead of simple trichomes.
Yinshania rivulorum (Dunn) Al-Shehbaz, G. Yang, L. L. Lu, & T. Y. Cheo, comb. nov.
Basionym: Nasturtium rivulorum Dunn, J. Linn. Soc., Bot. 38: 354. 1908. TYPE: CHINA. Fujian [as Fokien]: Fong Kong Tze, near Foochow, along the mossy banks of rivulets in shady woods, Dunn 2354 (holotype, HK; isotypes, GH!, K!).
Synonyms: Cochlearia rivulorum (Dunn) O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 8: 546. 1923; Hilliella rivulorum (Dunn) Y. H. Zhang & H. W. Li, Acta Bot. Yunnan. 8: 404. 1986.
Cochlearia formosana Hayata, J. Coll. Sci. Imp. Univ. Tokyo 30(1): 32. 1911. TYPE: CHINA. Taiwan: Shinko, Remogansha, 27 February 1908, Hayata 1390 (holotype, TI!). Hilliella formosana (Hayata) Y. H. Zhang & H. W. Li, Acta Bot. Yunnan. 8: 403. 1986; Yinshania formosana (Hayata) Y. Z. Zhao, Acta Sci. Nat. Univ. Intramongol. 23: 567. 1992.
Hilliella alatipes (Handel-Mazzetti) Y. H. Zhang & H. W. Li var. micarntha Y. H. Zhang, Acta Bot. Yunnan. 8: 403. 1986. TYPE: CHINA. Hunan: Jianhua, 22 April 1973, Hunan Institute of Chinese Medicine 7300069 (holotype, IBSC!; as SCBI).
Annuals, 8--50 cm tall, glabrous throughout. Basal and lowermost stem leaves 3-foliolate, rarely 1 basal leaf simple; petiole 4--10 cm long, gradually shorter upward, not winged; leaflet blade ovate to oblong or rarely lanceolate, 1--5 x 0.8--3.0 cm, papery, base cordate to rounded or cuneate, margin repand or rarely sinuate-toothed, with conspicuous callose mucros terminating veins, apex obtuse or acute; petiolule 2--12 mm long, longer on terminal leaflet. Upper leaves similar to lower ones, becoming simple at the inflorescence. Racemes lax, ebracteate; rachis straight. Pedicel 6--10 mm long, slender, straight, reflexed. Sepals 1--2 mm long. Petals white, obovate, 1.5--2.5 mm long, minutely clawed. Filaments white, 1--2 mm long; anthers ca. 0.2 mm long. Fruit oblong, 3--6 x ca. 1.5 mm, papillate when immature; septum absent. Style 0.5--0.8 mm long. Seeds 7--10, oblong, 0.8--1.0 x ca. 0.5 mm.
Habitat and phenology: shady river banks, mountain slopes; 300--800 m. Flowering and fruiting April through December.
Distribution: Fujian, Hunan, Taiwan.
Specimens examined: CHINA. Taiwan: Urai, Fairiu 534 (TI); same area, 30 March 1977, Togashi s.n. (TI); Taipei Co., Wulai, Lin 496 (TAI), Kao 7166 (TAI, TI), Wang 9208 (HAST), Li 442 (TNM); Taipei City, Wulai, en route to Hsiao-yi, Wang 9601 (HAST); Taipei City, Yangmingshan National Park, Luchiaokensi Preserve, Liao, Lee, & Hsu 235 (HAST); Taipei Co., Wuali, Wa-wa-ku, 2 March 1987, Liao s.n. (TAI); Taihohusyu, Bunzangun, Rahua, Suzuki ST-19075 (PE, TAI); Mt. Nuiohei, 10 April 1927, Sasaki s.n. (TNS); Prov. Taihoku, Urai-Rarasan, Ohwi 806 (TNS); Urai-sya, Taihoju, 1 April 1918, Sasaki s.n. (TNS); Taipen Shan, 25 April 1930, Sasaki s.n. (TNS).
Yinshania rivulorum is the only species of the genus that extends outside mainland China into Taiwan, where it is known as Cochlearia formosana. Zhao (1992) confused the nomenclature of the species and reduced it (as C. rivulorum) to synonymy of the later published Y. formosana (see the synonymy above).
Zhang (1986) recognized three varieties of Hilliella alatipes, each of which clearly represents a distinct species. As shown by Al-Shehbaz and Yang (1998), two of these varieties belong to two remarkably different species of Cardamine, and Cochlearia (or Hilliella) alatipes is a synonym of Cardamine fragariifolia O. E. Schulz. The third of Zhang's varieties, var. micrantha, is definitely a synonym of Y. rivulorum, a species also recognized by Zhang (1986). Zhao (1992) treated all of Zhang's three varieties of H. alatipes, along with Y. lichuanensis, as synonyms of Y. alatipes. Yinshania lichuanensis is a very distinct species that differs from Y. rivulorum (here including only one of Zhang's three varieties of H. alatipes) in being puberulent plants with acuminate-caudate leaf apices and fruit appressed to rachis. In contrast, Y. rivulorum has usually glabrous plants with obtuse to acute leaf apices and widely spreading to reflexed fruit.
Yinshania rivulorum appears to be very rare in Fujian and Hunan, as evidenced from the few type collections above. It is much more widespread in Taiwan. The type of H. alatipes var. micrantha differs from the rest of the collections here treated as Y. rivulorum in having slightly longer (6.0 vs. 3.0--4.5 mm) fruits and acute instead of obtuse or subacute leaf apex. Because only the type of var. micrantha has been examined, it is unclear whether or not this variety deserves recognition.
Yinshania sinuata (K. C. Kuan) Al-Shehbaz, G. Yang, L. L. Lu, & T. Y. Cheo, comb. nov.
Basionym: Cochlearia sinuata K. C. Kuan, Bull. Bot. Lab. N. E. Forest. Inst., Harbin 1980(8): 39. 1980. TYPE: China, Jiangxi, Suichuan, 31 March 1970, L. K. Dai 150 (holotype, PE!).
Synonyms: Hilliella sinuata (K. C. Kuan) Y. H. Zhang & H. W. Li, Acta Bot. Yunnan. 8: 404. 1986.
Annuals, (8--)15--30 cm tall, glabrous throughout. Stems erect to subdecumbent, branched. Petiole 0.5--5.0 cm long; leaf blade simple or rarely lowermost trifoliolate, ovate, 1.5--6.0 x 1--4 cm, base cordate to subtruncate, margin sinuate to repand, with apiculate callosities at vein tips, apex obtuse. Racemes ebracteate; rachis straight. Flower parts spreading. Pedicel slender, spreading in flower, distinctly reflexed and (5--)8--12 mm long in fruit, straight, glabrous. Sepals oblong-ovate, 1.7--2.5 x 0.8--1.1 mm, glabrous, narrowly membranous at margins and apex. Petals white, obovate, 2.5--3.0 x 1.2--1.7 mm, base short clawed, apex rounded. Filaments 1.5--2.0 mm long; anthers 0.45--0.55 mm long. Fruit oblong, 6--12 x 1.7--2.3 mm, glabrous. Style 0.8--1.5 mm long. Seeds (5--)7--14(--21), brown, ovate, 0.8--1.3 mm long.
In superficial examination, Y. sinuata looks very much like some forms of Eutrema. However, the two genera are unrelated. Yinshania species have predominantly compound or deeply pinnatifid basal leaves not deeply cordate basally; nontorulose fruits; spreading sepals, petals, and stamens; and petals about the same size as sepals and not differentiated into a claw and blade. In contrast, Eutrema always has simple basal leaves with cordate base, predominantly torulose fruits, erect floral parts, and petals distinctly longer than sepals and always differentiated into blade and claw. In our opinion, Yinshania appears to be more closely related to Cardamine than to Eutrema.
KEY TO THE SUBSPECIES OF Y. SINUATA
1a. Lowermost leaves always simple; seeds 0.8--0.9 mm long ... ssp. sinuata
1b. At least some of the lowermost leaves trifoliolate; seeds 1.2--1.3 mm long ... ssp. qianwuensis
Yinshania sinuata ssp. sinuata
Lowermost leaves always simple. Fruit 5--10 mm long. Seeds 0.8--0.9 mm long.
Habitat and phenology: mountain slopes, rocky crevices, shady moist sand, dense forest; near sea level to 200 m. Flowering in March, fruiting in May.
Distribution: Guangdong, Hunan, Jiangxi.
Specimens examined: CHINA. Guangdong: Kweiyang, Mandse-schan, Mell 532 (WU); Man Chi Shan, Shek Pik Ha Village, Jen Hua district, Tsang 26144 (A, NAS). Hunan: Xingning County, Bajiao village, Luo 2094 (PE). JIANGXI: Jian County, Chen 354 (PE); Jian County, Futian, Da-ken, Qingshua Jian, Dai 240 (PE). Zhejiang: Chunan County, Qiuyuan, Hong 966 (MO),
The original publication gave Dai 150 as the holotype, but the type in PE bears Dai 159. All of the other information given in the original publication matches Dai 159, and there is no Brassicaceae specimen at PE that bears Dai 150. Therefore, we believe that the collection number in the original publication was a typographical error.
Yinshania sinuata ssp. qianwuensis (Y. H. Zhang) Al-Shehbaz, G. Yang, L. L. Lu, & T. Y. Cheo, comb. et stat. nov.
Basionym: Hilliella sinuata (Kuan) Y. H. Zhang & H. W. Li var. qianwuensis Y. H. Zhang, Acta Bot. Yunnan. 8: 405. 1986. TYPE: CHINA. Jiangxi: Qianwu, 700 m, 8 May 1958, Q. M. Hu & Q. H. Li 1761 (holotype, LBG (as HLG); isotype, PE!).
At least some of the lowermost leaves trifoliolate. Fruit 6--13 mm long. Seeds 1.2--1.3 mm long.
Habitat and phenology: In forests; 200--700 m. Fruiting in May.
Additional specimens examined: CHINA. Jiangxi: Xun Wu Zhizhu, Hu 1701 (KUN).
Both subspecies of Y. sinuata are poorly collected, and the apparently sufficient differences between the two may not remain with the gathering of more extensive material. Zhao (1992) reduced this subspecies to synonymy of Y. rivulorum (as Y. formosana), but it is evident that the two belong to different species. Yinshania rivulorum is characterized by having all stem leaves trifoliolate, petals 1.5--2.5 mm long, anthers ca. 0.2 mm long, and oblong papillate fruits 6--12 x 1.7--2.3 mm. In contrast, Y. sinuata has all leaves simple or only the lowermost trifoliate, and the petals are 0.8--1.1 mm, anthers 0.45--0.55 mm, and ovate glabrous fruits 4--6 x ca. 1.5 mm.
According to Zhang (1986), the type of ssp. qianwuensis was collected on 8 May 1958 at 700 m. The other collection of this taxon (Hu 1701), which was annotated by Zhang as the isotype, was collected on 6 May 1958 at 200 m and therefore does not belong to the type collection.
Yinshania paradoxa (Hance) Y. Z. Zhao, Acta Sci. Nat. Univ. Intramongol. 23: 567. 1992.
Basionym: Cardamine paradoxa Hance, J. Bot. 6: 111. 1868. TYPE: CHINA. Guangdong [as prov. Cantonensis]: Kai-kunshek, 27 May 1867, Sampson 13844 (holotype, K!; isotypes, GH!, K!, LE!, P!, W!).
Synonyms: Cochlearia paradoxa (Hance) O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 8: 546. 1923. Hilliella paradoxa (Hance) Y. H. Zhang & H. W. Li, Acta Bot. Yunnan. 8: 404. 1986.
Annuals, 30--70 cm tall, glabrous throughout or rarely puberulent with minute simple trichomes to 0.06 mm long. Stems erect, angled, simple or branched above. Basal and lowermost stem leaves 3- or very rarely 5-foliolate; petiole 4--8 cm long; leaflet blade ovate to oblong or lanceolate, 1.5--5.0(--6.0) x 1--2(--3) cm, base subtruncate to cuneate or obtuse to oblique, margin crenate or repand and with mucronate callosities terminating veins, apex obtuse to acute and mucronate; petiolule 0.2--2.0 cm long, longer on terminal leaflets. Racemes terminal and lateral, ebracteate; rachis slender, straight or subflexuous apically. Pedicel (1.5--)2.5--4.0 mm long in fruit, straight, widely spreading to reflexed. Sepals oblong, 1.5--2.0 mm long. Petals white, obovate, 2.0--2.5 mm long, apex rounded; claw to 1 mm long. Filaments white, 1.5--2.0 mm long; anthers oblong, ca. 0.4 mm long. Fruit ovoid to ellipsoid, 3--4 x 2.0--2.5 mm; valves glabrous; septum absent. Style 1--2 mm long. Seeds 2 or 3, oblong, 1.5--1.8 x ca. 1 mm.
Habitat and phenology: mountain slopes, valleys, roadsides; 300--1000 m. Flowering April to November, fruiting May to December.
Distribution: Guangdong, Guangxi, Hubei, Sichuan.
Specimens examined: CHINA. Guangdong: Xing Hua, Tianzhu Yei, Shi 12525 (IBSC). Guangxi: Dayao Shan County, Pin Zhu Xiang, Li 400198 (IBSC). Hubei: without locality, Henry 4199 (GH). Sichuan: Bei Yei, Xiong et al. 1773 (PE); Chongqing, Xiong et al. 3226 (PE); Chongqing, Low 481 (PE).
Yinshania paradoxa is most closely related to Y. rivulorum, from which it is readily separated by its ovate to obovate 2- or 3-seeded fruits, styles 0.5--0.8 mm, fruiting pedicels (1.5--)2.5--4.0 mm, and seeds 1.5--1.8 x ca. 1 mm. In contrast, Y. rivulorum has narrowly oblong 7- to 10-seeded fruits, styles (1.0--)1.5--2.0 mm, fruiting pedicels 6--10 mm, and seeds 0.8--1.0 x ca. 0.5 mm.
Yinshania lichuanensis (Y. H. Zhang) Al-Shehbaz, G. Yang, L. L. Lu, & T. Y. Cheo, comb. nov.
Basionym: Hilliella lichuanensis Y. H. Zhang, Acta Bot. Yunnan. 9: 158. 1987. TYPE: China, Jiangxi, Lichuan, 7 June 1958, M. X. Nie & S. S. Lai 2831 (holotype, PE!; isotypes, IBSC!, KUN!, LBG [as HLG], SHM).
Synonyms: Cochlearia lichuanensis (Y. H. Zhang) L. L. Lu; Bull. Nanjing Bot. Gard. Mem. Sun Yat Sen 1991: 17. 1991.
Hilliella changhuaensis Y. H. Zhang, Acta Bot. Yunnan. 9: 155. 1987. TYPE: China, Zhejiang, Linan, Changhua, 17 June 1957, X. Y. He 23624 (holotype, NAS!; isotype, PE!). Cochlearia changhuaensis (Y. H. Zhang) L. L. Lu, Bull. Nanjing Bot. Gard. Mem. Sun Yat Sen 1991: 17. 1991.
Lilliella longistyla Y. H. Zhang, Acta Bot. Yunnan. 9: 153. 1987. TYPE: CHINA. Zhejiang: Longquan, 3 June 1919, H. H. Hu, s.n. (holotype, ZM [as ZJM]). Cochlearia longistyla (Y. H. Zhang) L. L. Lu, Bull. Nanjing Bot. Gard. Mem. Sun Yat en 1991: 18. 1991.
Hilliella guangdongensis Y. H. Zhang, Acta Bot. Yunnan. 9: 157. 1987. TYPE: CHINA. Guangdong: Renhua, 900--1150 m, 29 August 11958, L. Deng 7298 (holotype, PE!: isotype, IBSC [as SCBI]).
Annuals to perennials, (40--)60--110 cm tall, puberulent with simple trichomes 0.03--0.08(--0.1) mm long. Stems erect, sulcate. Basal and lowermost stem leaves 3- to 5-foliolate; petiole 2.5--7.0 cm long, usually narrowly winged, grooved and sparsely to densely puberulent adaxially; leaflet blade lanceolate, (3--)6--14 x (0.8--)2.0--5.0 cm, terminal ones usually larger, base cuneate to rounded or oblique, margin serrate to irregularly dentate, mucronate, apex acuminate-caudate, abaxially puberulent, adaxially sparsely puberulent at least on veins; petiolule of lateral leaflets obsolete to 5 mm. Uppermost leaves simple or rarely 3-foliolate, similar in morphology to leaflets of lower leaves. Racemes terminal and lateral, elongated considerably in fruit. Fruiting pedicel straight, slender, 2--5 mm long, erect-ascending, subappressed to rachis. Sepals oblong, 2.0--2.5 mm long. Petals white, spatulate, 2.5--3.0(--3.5) x ca. 1 mm, base clawed. Filaments white, 2.0--2.5 mm long; anthers oblong, 0.5--0.7 mm long. Fruit obovoid to oblong or ellipsoid, 3--5 x 1.5--2.0 mm; valves thick; gynophore obsolete to 0.5 mm long; septum absent. Style 0.5--1.5(--2) mm long. Seeds 1--3, brown to blackish brown, oblong, 1.5--2.5 x 1.0--1.4 mm, reticulate.
Habitat and phenology: limestone areas, wet valleys, shady woods, mountain slopes, roadsides, streamsides; 300--1200 m. Flowering May through June, fruiting June through August.
Distribution: Anhui, Fujian, Guangdong, Jiangxi, Zhejiang.
Additional specimens examined: CHINA. Anhui: Huan Shan, Fu Xi, Shao 810105 (PE). Fujian: Shuen Chan, Tian Ping, Ban Shan, Li 4454 (PE), Li 4516 (PE). Guangdong: Ren Hua County, Chang Jiang Xiang, Fan-Wu Shan, Deng 7298 (IBSC, MO). Jiangxi: Jian County, Futian, anonymous 227 (PE); Xing Zi County, Lou 82103 (NAS); Wu Ning, Ruo Peng Chang Shu, Lai 2825 (KUN, PE); Li Chuan, Lou 87301 (NAS). ZHEJIANG: Chang Hua, Lon Tan Zhi Keng, Chiu 449 (MO); Xi Hu, Ji Guan Shan, Li & Cheng 889 (PE); Chan Hua, Jou Pu, Deng 4773 (PE).
An examination of a large number of specimens of this complex reveals that H. lichuanensis, H. longistyla, H. guangdongensis, and H. changhuaensis are conspecific. Zhang (1987b) separated these primarily on the basis of differences in the density of indumentum on the inflorescence, length of style, and size and shape of the fruit. However, all of these characters show continuous variation, and it is impossible to subdivide this complex into distinct entities. Zhang (1987b) separated H. lichuanensis from the other three species above primarily on the basis of its having hairy versus glabrous sepals and inflorescence. When used alone in the Brassicaceae, the density or even presence versus absence of indumentum often are among the most unreliable features for recognizing taxa. Zhang separated H. longistyla from the other three species on the basis of its having long styles ca. 1 mm in flower and ca. 2 mm in fruit. However, the other three species (H. lichuanensis, H. guangdongensis, and H. changhuaensis) were said to have styles 1.0--1.5 mm long. Zhao (1992) was the first to correctly place these four species under one name, but he wrongly listed them as synonyms of Y. alatipes, a species clearly shown by Al-Shehbaz and Yang (1998) to be the same as Cardamine fragariifolia. In the absence of mature fruits, Cardamine can be easily separated from Yinshania because of its having erect rather than spreading petals and stamens and a broad rather than narrow placental region. Three of Zhang's (1987b) species of Hilliella (H. lichuanensis, H. changhuaensis, and H. longistyla), along with Y. alatipes, were recognized as distinct species of Cochlearia by Lu (1991), but as indicated above, these Chinese species are unrelated to Cochlearia.
Yinshania rupicola (D. C. Zhang & J. Z. Shao) Al-Shehbaz, G. Yang, L. L. Lu, & T. Y. Cheo, comb. nov.
Basionym: Cochlearia rupicola D. C. Zhang & J. Z. Shao, Acta Phytotax. Sin. 24: 404. 1986. TYPE: CHINA. Anhui: Shitai, Kuniujiang, Qimencha, 1150 m, 29 June 1983, J. Z. Shao 835008 (holotype, ANUB; isotype, NAS!).
Synonym: Hilliella rupicola (D. C. Zhang & J. Z. Shao) Y. H. Zhang, J. Wuhan Bot. Res. 8: 322. 1990.
Perennials, 30--100 cm tall, with a thick tuberous rhizome 2--4 cm wide, glabrous throughout. Stems 1 or few from rhizome, erect, sulcate, branched above. Basal and lowermost stem leaves compound, 5- to 9-foliolate; petiole (3--)8--15(--20) cm long; leaflets broadly oblong to ovate or lanceolate, (2--)4--7(--10) x (1--)2--3(--4) cm, base oblique to cuneate or subtruncate, margin coarsely serrate with a distinct mucro at tooth apex, apex obtusely acuminate with a distinct mucro; petiolule 1--5(--20) mm long. Upper stem leaves 3- or 5-foliolate, becoming gradually smaller and narrower upward. Racemes terminal and lateral, many flowered; lowermost flowers sometimes bracteate. Fruiting pedicel filiform, recurved to reflexed 4--10 mm long. Sepals oblong, 1.5--2.0 mm long, whitish at margin. Petals white, oblong to obovate, 2--4 x 1--2 mm long, apex rounded; claw to 1 mm long. Filaments white, 1.0--1.5 mm long; anthers ovate, ca. 0.5 mm long. Fruit suborbicular-obovate to narrowly elliptic, conspicuously flattened parallel to replum, 2.5--6.0 x 2--3(--4) mm; valves obscurely veined; septum sometimes absent; gynophore ca. 0.5 mm long. Style (0.5--)1.0--2.0 mm long. Seeds 2 or 3, brown, ovate oblong, flat, 1.2--1.8 x 0.6--1.5 mm.
Key to the subspecies of Yinshania rupicola
1a. Fruit narrowly elliptic, 5--6 x 2.0--2.5 mm ... ssp. rupicola
1b. Fruit suborbicular-obovate, 2.5--4.0 x 2.5--4.0 mm ... ssp. shuangpaiensis
Yinshania rupicola ssp. rupicola
Petals white. Fruit narrowly elliptic, 5--6 x 2.0--2.5 mm. Flowering in May, fruiting in June.
Habitat and phenology: forests, rocky crevices in wet shady places; 1000--1200 m. Flowering in May, fruiting in July.
Known only from the type collection.
Yinshania rupicola ssp. shuangpaiensis (Z. Y. Li) Al-Shehbaz, G. Yang, L. L. Lu, & T. Y. Cheo, comb. et stat. nov.
Basionym: Hilliella shuangpaiensis Z. Y. Li, Acta Bot. Yunnan. 10: 117. 1988. TYPE: CHINA. Hunan: Shuangpai, Ma Jiang, Tong-zhi Shan, 1,000 m, 22 September 1986, C. L. Liao 1439 (holotype, HUTM, as HNMI).
Synonyms: Hilliella xiangguiensis, Y. H. Zhang, Acta Bot. Yunnan. 19: 139. 1997. TYPE: CHINA. Hunnan: Suining, Huangshuangxiang, Chibancun, Laolongta, 1700 m, 17 July 1985, C. Z. Yuan 319 (holotype, HUTM).
Petals white to lilac. Fruit suborbicular-obovate, 2.5--3.0 x ca. 3 mm.
Habitat and phenology: wet places, valleys, forests, along streams; 700--1000 m. Flowering May to September, fruiting June through October.
Distribution: Fujian, Guangxi, Hunan, Sichuan.
Additional specimens examined: CHINA. Fujian: Chong An County, Shan Gong, Wuling Shan Plant Expedition 2233 (IBSC). Guangxi: Chuen Yuen, Chung 82026(A). Hunan: Heng Shan, Nan Yue, Guan Ji Shi, Chen 3357 (MO, PE); Lin County, anonymous 660507 (PE). Sichuan: no locality, cultivated from seeds, Murata & Mizuno s.n. (KUN).
The original description of Cochlearia rupicola indicated that the plant is annual, but the illustration and the isotype above clearly show that the plant is perennial. Subspecies rupicola is only known from the type collection, and nothing can be said about its overall variability. It is remarkably similar to ssp. shuangpaiensis in all other morphological characters, including leaf shape, flower size, rhizomes, lack of septum, length of style, flattening of fruit, and presence of a short gynophore, and the only basic difference between the two is the fruit shape and size given in the key above. Therefore, it is more practical to treat the two taxa under the same species.
The strongly recurved, slender fruiting pedicels, the flattened suborbicular fruit, and oblong-lanceolate, trifoliolate, uppermost stem leaves are quite characteristic of this subspecies. Although we have not examined the holotype, the excellent illustration and detailed description of H. xiangguiensis clearly show that the taxon is indistinguishable from the plants treated here as ssp. shuangpaiensis.
Yinshania hunanensis (Y. H. Zhang) Al-Shehbaz, G. Yang, L. L. Lu, & T. Y. Cheo, comb. nov.
Basionym: Hilliella hunanensis Y. H. Zhang, Acta Bot. Yunnan. 9: 160. 1987. TYPE: CHINA. Hunan: Hengshan, Guangjisi, 750 m, 17 May 1963, L. H. Liu 15702 (holotype, PE!; isotype, KUN!).
Perennials, glabrous throughout. Rhizomes thick, to 5 mm in diam. Stems slender, branched from base, ascending. Basal and lowermost stem leaves 5-foliolate; petiole 5.0--8.5 cm long; leaflets thin, broadly ovate to suborbicular, (0.5--)1.0--2.5(--3.5) x (5.0--)1.0--2.0(--2.5) cm, margin shallowly 5- to 7-lobed to repand, veins ending in a distinct mucro; terminal leaflet cuneate to subcordate at base, lateral leaflets oblique at base; petiolule (2--)5--10(--30) mm long. Middle and upper stem leaves gradually reduced in size, 3-foliolate. Inflorescences corymbose raceme, lax, considerably elongated in fruit, bracteate along lower half; bracts simple or lowermost 3-foliolate, considerably reduced in size upward. Fruiting pedicel slender, often recurved, 0.8--1.5 cm long. Sepals 1.5--1.7 mm long. Petals white, 2.0--2.5 x ca. 1.5 mm. Filaments ca. 1 mm long; anthers ca. 0.3 mm long. Fruit broadly elliptic to suborbicular, (3--)4--6 x 3--4(--5) mm, flattened; valves glabrous, inconspicuously veined; gynophore ca. 0.5 mm long. Style (0.5--)1.0--2.0(--3.0) mm long. Seeds 2 or 3, oblong, 1.5--2.0 x 0.7--1.2 mm, minutely papillose.
Habitat and phenology: rocky areas, valleys, streamsides, dense forests; 500--1600 m. Flowering May through June, fruiting June through August.
Distribution: Guangxi, Hunan, Jiangxi.
Additional specimens examined: CHINA. Guangxi: Lonshen, Dadiziang, Huoya-tang, Guangfu Forest Investigation Team 187 (IBSC, MO, PE). Hunan: Mongshan, Shangshiken, Huang 112168 (MO); Hengshan, Nanyue, Guangjishi, Cheng 3466 (IBSC, MO). Jiangxi: Lushan, Yulaofen, Wulaodong, Guan 74571 (PE); Lushan, Wulaofeng, Wang 87010 (NAS), Deng et al. 82101 (NAS), Guan et al. L-140 (KUN).
Zhao (1992) reduced Y. hunanensis to synonymy of Y. huii. Although we have not seen material of the latter, the protologue shows that it is quite different from Y. hunanensis, and the two species can be readily separated by characters listed under Y. huii.
Yinshania huii (O. E. Schulz) Y. Z. Zhao, Acta Sci. Nat. Univ. Intramongol. 23: 567. 1992.
Basionym: Cochlearia huii O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 8: 546. 1923. TYPE: CHINA. Jiangxi [as Kiangsi]: Anfu, Wookung San, 1500 m, April 1921, H. H. Hu 731 and H. H. Hu 752 (syntypes, B). Hilliella huii (O. E. Schulz) Y. H. Zhang & H. W. Li, Acta Bot. Yunnan. 8: 404. 1986.
Annuals, glabrous throughout, 15--20 cm tall. Stems branched, flexuous. Basal leaves trifoliolate; petiole ca. 3 cm; terminal leaflet ovate to subcordate, 1--2 cm long, unequally coarsely crenate, apex emarginate and mucronulate, petiolule long; lateral leaflets smaller and with shorter petiolules, obliquely ovate; middle stem leaves ca. 1 cm long, trifoliolate, with ovate terminal leaflet and much smaller lateral leaflets; uppermost leaves simple. Inflorescences lax racemes, 8- to 10-flowered, bracteate to apex. Lowermost pedicels to 1.5 cm long. Sepals oblong-ovate, ca. 2 mm long. Petals white to pink, ca. 3 mm long, ovate, cuneate at base, rounded at apex. Stamens 2.0--2.5 cm long; anthers oblong, ca. 0.5 mm long. Pistil narrowly cylindric; ovary 8-ovuled. Style ca. 0.7 mm long. Immature fruits ellipsoid, ca. 1.5 mm long; valves minutely papillate.
Habitat and phenology: Known only from two syntypes above. Both are flowering material collected in April at 1500 m.
Yinshania huii is most closely related to Y. hunanensis, and they are easily separated from the remaining species of Yinshania on the basis of their bracteate inflorescences. The syntypes of Y. huii have not been seen. From the original description, C. huii is an annual, with narrowly cylindric pistils, 8-ovuled locules, and papillate fruit valves. Yinshania hunanesis is a perennial, with ovate pistils, 1- or 2-ovuled locules, and glabrous fruit valves. Yinshania huii was recognized by Lu (1991) as Cochlearia, but its relationship to Y. hunanensis was not indicated, and the latter species was not mentioned
Cochlearia alatipes Handel-Mazzetti, Symb. Sin. 7: 370. 1931; Hilliella alatipes (Handel-Mazzetti) Y. H. Zhang & H. W. Li, Acta Bot. Yunnan. 8: 402. 1986; Yinshania alatipes (Handel-Mazzetti) Y. Z. Zhao, Acta Sci. Nat. Univ. Intramongol. 23: 568. 1992. TYPE: China, SW Hunan, in monte Yün-schan prope urbem Wukang, 1000 m, 12 June 1918, Handel-Mazzetti 12097 (holotype, WU!; isotypes, E!, W!, WU!) = Cardamine fragariifolia O. E. Schulz, Bot. Jahrb. Syst. 32: 446. 1903 (see Al-Shehbaz & Yang, 1998).
Hilliella alatipes (Handel-Mazzetti) Y. H. Zhang & H. W. Li var. macranthus Y. H. Zhang, Acta Bot. Yunnan. 8: 403. 1986. TYPE: CHINA. Yunnan: Malipo, Guan-gaw, 1000 m, 14 February 1940, C. W. Wang 86836 (holotype, KUN!; isotype, IBSC!) = Cardamine cheotaiyienii Al-Shehbaz & G. Yang, Harvard Papers Bot. 3(1): 73-78. 1998.
Yinshania yunnanensis (W. W. Smith) Y. Z. Zhao, Acta Sci. Nat. Univ. Intramongol. 23: 565. 1992. Camelina yunnanensis W. W. Smith, Notes Roy. Bot. Gard. Edinburgh 11: 202. 1919. TYPE: CHINA. Yunnan: Sung-kwei valley, May 1927, G. Forrest 13776 (holotype, E!). As shown by Zhang (1996b), the species is a synonym of Rorippa globosa (Turczaninow) Thellung.
We are grateful to Paul Berry for his critical review of the manuscript. We thank Henry Noltie for his help in providing information about some collections at the Royal Botanic Garden, Edinburgh. We thank the directors and curators of A, B, BM, E, GH, HAST, IBSC, K, KUN, LE, MO, NAS, NY, P, PE, TAI, TI, TNS, US, W, and WU for allowing our study of the cited specimens.
Al-Shehbaz, I. A., and G. Yang. 1998. Notes on Chinese Cardamine. Harvard Papers Bot.
An, Z. X. 1987. Sisymbrieae. In T. Y. Cheo, ed., Crusiferae. Fl. Reipubl. Popularis Sin. 33: 396--453. Science Press, Beijing.
Jackson, P. S. W., and J. R. Akeroyd. 1993. Cochlearia. In T. G. Tutin et al., eds., Fl. Europaea, 2d ed. 1: 378--380. Cambridge Univ. Press, Cambridge.
Kuan, K. C. 1987. Lepidieae. In T. Y. Cheo, ed., Crusiferae. Fl. Reipubl. Popularis Sin. 33: 44--109. Science Press, Beijing.
Löve, A., and D. Löve. 1976. Nomenclatural notes on Arctic plants. Bot. Notiser 128: 497--523.
Lu [as Lou], L. L. 1991. The genus of Cochlearia (Cruciferae) in southeastern China. Bull. Nanjing Bot. Gard. 1991: 15--22.
------. 1993. The identification and revision of the type of Cochleariopsis Y. H. Zhang (Cruciferae). Acta Phytotax. Sin. 31: 286--287.
Ma, Y. C., and Y. Z. Zhao. 1979. Yinshania, a new genus of Chinese Cruciferae. Acta Phytotax. Sin. 17: 113--114.
Oliver, D. 1887. Nasturtium henryi. Icon. Pl. 18: pl. 1719.
Pobedimova, E. 1969. Revisio generis Cochlearia L. 1. Novit. Syst. Pl. Vasc. 6: 67--106.
------. 1970. Revisio generis Cochlearia L. 2. Novit. Syst. Pl. Vasc. 7: 167--195.
Rollins, R. C. 1993. The Cruciferae of Continental North America. Stanford University Press, Stanford.
Schulz, O. E. 1923. Eine neue Sektion der Gattung Cochlearia L. Notizbl. Bot. Gart. Berlin-Dahlem 8: 544--546.
------. 1929. Asiatische Cruciferen verschiedener Herkunft. Notizbl. Bot. Gart. Berlin-Dahlem 10: 554--557.
------. 1935. Neue Cruciferen Arten. II. Repert. Sp. Nov. Regni Veg. 38: 108--109.
------. 1936. Cruciferae. In A. Engler and K. Prantl, eds., Nat. Pflanzenfam., 2d ed. 17B: 227--658. Verlag von Wilhelm Engelmann, Leipzig.
Ying, T. S., Y. L. Zhang, and D. E. Boufford. 1993. The Endemic Genera of Seed Plants of China. Science Press, Beijing.
Zhang, Y. H. 1985. Cochleariopsis---A new genus of Chinese Cruciferae. Acta Bot. Yunnan. 7: 143--145.
------. 1986. Hilliella, a new genus of Cruciferae. Acta Bot. Yunnan. 8: 397--406.
------. 1987a. A revision of genus Yinshania (Cruciferae). Acta Phytotax. Sin. 25: 204--219.
------. 1987b. Five new species of the Hilliella (Cruciferae). Acta Bot. Yunnan. 9: 152--161.
-------. 1989. On the identity of Yinshania albiflora. Acta Phytotax. Sin. 28: 74--75.
------. 1993. A new species of Yinshania with a discussion on the evolution and origin of the genus. Acta Bot. Yunnan. 15: 364--368.
------. 1995a. A new species of Hilliella (Cruciferae) from Anhui. Acta Phytotax. Sin. 33: 94--96.
------. 1995b. On classification of Hilliella warburgii and Cochleariella zhejiangensis. Acta Phytotax. Sin. 33: 175--178.
------. 1996a. A study on the genus Yinshania. Bull. Bot. Res., Harbin 16: 445--454.
------. 1996b. A note of Yinshania zayuensis and Camelina yunnanensis. Acta Phytotax. Sin. 34: 87.
------. 1997. A new species of Hilliella from Hunan and Guangxi. Acta Bot. Yunnan. 19: 139--140.
------, and J. J. Cai. 1989. Observation on the genera Yinshania, Hilliella, Cochleariella, and Cochlearia (Cruciferae) by SEM. Acta Bot. Bor.-Occ. Sin. 9: 224--231.
Zhao, Y. Z. 1992. A taxological revision on Cochlearia, Yinshania, Hilliella, and Cochleariella in China. Acta Sci. Nat. Univ. Intramongol. 23: 561--573.