Abstract. Staintoniella nepalensis, the generic type, five species of Aphragmus are presented. The generic limits of Aphragmus and related genera are discussed.

Keywords: Aphragmus, Brassicaceae, nomenclature,

As originally delimited by Hara (1974), the genus Staintoniella H. Hara included two species that have not been subjected to a subsequent critical study. The generic type, S. nepalensis H. Hara, is endemic to Nepal, whereas S. verticillata (Jeffrey & W. W. Smith) H. Hara is endemic to Xizang (Tibet) and Yunnan, China (Kuan, 1987).

The two species of Staintoniella are significantly different from each other in several important characters that support their placement in different genera. Plants of S. nepalensis are perennials with many branched, rhizomelike, slender caudices, persistent petiolar remains of basal leaves, leafless scapes, blue flowers, minute glandlike papillae on the inflorescence, basally unexpanded staminal filaments, and terete slender replums. By contrast, those of S. verticillata are biennial with fleshy, fusiform taproots crowned with oblong scales, deciduous basal leaves, leafy stems with the lowermost leaves whorled, white flowers, distinct trichomes, basally dilated filaments, and considerably flattened replums. Al-Shehbaz (2000a) transferred S. verticillata to Taphrospermum C. A. Meyer and showed that it is most closely related to T. lowndesii (H. Hara) Al-Shehbaz, a species that was initially described by Hara (1978) in the monotypic Glaribraya H. Hara.

Hara (1974) indicated (p. 196) that Staintoniella is "without a close ally" and suggested that "it somewhat resembles" Pegaeophyton Hayek & Handel -Mazzetti, a Himalayan genus of six species (Al-Shehbaz, 2000b). These genera are not closely related, and they can be separated readily by the presence of bracteate racemes in Staintoniella and solitary flowers from the axils of basal leaves in Pegaeophyton.

A closer examination of Staintoniella nepalensis clearly shows that it is very similar to Aphragmus oxycarpus (J. D. Hooker & Thomson) Jafri, especially in the perennial habit and in having petiolate basal leaves that form distinct rosettes, persistent and broadly expanded petiolar remains of previous years, leafless scapes, bracteate inflorescences, minutely puberulent upper parts, purple flowers, confluent and well-developed nectaries, adaxially puberulent pedicels, nonapiculate anthers, basally slender filaments, flattened, eseptate fruits, entire stigmas, obscurely veined valves, and well-developed styles. The flower color in A. oxycarpus ranges from white to purple. The two species can be separated only by compact caudex, smaller petals 3.5-5.0(-6.0) mm long, oblanceolate to spatulate or oblong to linear basal leaves, and septate fruits in A. oxycarpus, and slender, rhizomelike caudex, larger petals (6-)7-9 mm long, ovate to broadly ovate basal leaves, and eseptate fruits. The extensive similarities clearly support the assignment of the two species in one genus. Therefore, Staintoniella is reduced herein to synonymy of Aphragmus Andrzejowski ex de Candolle.

Although some authors (e.g., Schulz, 1924; Hedge, 1968; Yunussov, 1978; Hara, 1979) maintained Braya oxycarpa J. D. Hooker & Thomson in Brava Sternberg & Hoppe, the species is better accommodated in Aphragmus, as was done by Jafri (1956, 1973), An (1987), Hajra et al. (1993), and Czerepanov (1995). Aphragmus differs from Braya by having fully bracteate inflorescences, stems minutely puberulent with papillate trichomes less than 0.1 mm long, often filiform funicles as long as or longer than seeds, adaxially puberulent pedicels, and median nectaries confluent with laterals. By contrast, Braya has ebracteate inflorescence. (rarely lowermost flowers bracteate), stems or other parts with distinct trichomes to 1 mm long, thick funicles often shorter than seeds, pedicels pubescent all around, and median nectaries totally lacking. In my opinion, these differences are significant enough to support the maintenance of both genera.

Although Jafri (1957) confused the limits of Aphragmus with Lignariella Baehni, the two genera are quite distinct, and their relationships and boundaries were discussed by Al-Shehbaz et al. (2000).

Because of the lack of adequate material of Aphragmus eschscholtzianus Andrzejowski ex de Candolle and A. involucratus (Bunge) O. E. Schulz, the author preferred to provide the following synopsis of Aphragmus without descriptions and citation of specimens.

Aphragmus Andrzejowski ex de Candolle, Prodr. 1: 209. 1824.

Type species: Aphragmus eschscholtzianus Andrzejowski ex de Candolle.

Synonyms: Oreas Chamisso & Schlechtendal, Linnaea 1: 29. 1826. TYPE: O. involucrata Chamisso & Schlechtendal.

Staintoniella H. Hara, J. Jap. Bot. 49: 196. 1974. TYPE: S. nepalensis H. Hara.

Herbs perennial, with thick caudex covered with petiolar remains of previous years, sometime with slender rhizomes. Trichomes eglandular, simple or shortly stalked-forked, less than 0.1 mm. Stems erect to ascending, branched basally, often minutely puberulent. Basal leaves petiolate, rosulate, simple, entire. Stem leaves petiolate or sessile, base cuneate to attenuate, not auriculate. Racemes few to several flowered, bracteate throughout, slightly elongated or not in fruit. Fruiting pedicels slender, erect, ascending, divaricate, sometimes recurved, puberulent adaxially. Sepals oblong, base of inner pair not saccate. Petals white, pink, or purple; blade broadly obovate to spatulate, apex obtuse or rounded; claw subequaling sepals. Stamens 6, slightly tetradynamous; filaments dilated or not at base; anthers not apiculate at apex. Nectar glands confluent and subtending bases of stamens. Fruit dehiscent silicles or siliques, ovate, elliptic, lanceolate, rarely linear, latiseptate; valves with an obscure or distinct midvein, smooth; replum flattened basally; septum complete or perforate and membranous, or absent; style obsolete, or short, rarely to 2 mm; stigma capitate, entire. Seeds uniseriate or biseriate, wingless, oblong to ovoid, plump, on filiform funicles often longer than seeds; seed coat minutely reticulate, not mucilaginous when wetted; cotyledons incumbent.

Five species: Central Asia, Himalayas, and North America.

KEY TO THE SPECIES OF APHRAGMUS

1a. Plants apparently annual, with slender long rhizomes; fruit linear; fruiting pedicels often recurved; petioles slender to base; seeds uniseriate … 1. A. obscurus

1b. Plants distinctly perennial, with compact or rhizomelike caudex; fruit elliptic to lanceolate; fruiting pedicels straight; petioles expanded into persistent base; seeds biseriate … 2

2a. Plants with many-branched. rhizomelike caudex covered with distinct internodes separating whorls of petiolar remains of successive growing seasons; petals (6-)7-9 mm long … 5. A nepalensis

2b. Plants with compact caudex not differentiated into distinct internodes; petals 2.5-5.0(-6.0) … 3

3a. Septum lacking; plants of Alaska and Canada … 2. A. eschscholtzianus

3b. Septum complete or perforate; plants of Asia … 4

4a. Infructescence capitate; bracts ovate; Russia, Mongolia … 3. A. involucratus

4b. Infructescence short racemose: bracts linear to lanceolate or oblong; Himalayas ... 4. A. oxycarpus

1. Aphragmus obscurus (Dunn) O. E. Schulz, Repert. Sp. Nov. Regni Veg. 31: 330. 1933.

Basionym: Draba obscura Dunn, Bull. Misc. Inform. Kew 1924: 383. 1924. TYPE: KASHMIR. Sonamarg, 12,000-13,000 ft, September 1917, R. R. & I. D. Stewart 3547 (Holotype: K).

Synonyms: Lignariella obscura (Dunn) Jafri, Candollea 16: 134. 1957.

Aphragmus himalaicus O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 9: 1058. 1927. TYPE: KASHMIR. Sadul, Kagan, Hazra, 15 August 1897, Inayat 21124a (Lectotype, here designated: CAL, n. v.; Isolectotype: B).

Distribution: endemic to Kashmir.

2. Aphragmus eschscholtzianus Andrzejowski ex de Candolle, Prodr. l: 210. 1824. TYPE: ALEUTIAN ISLANDS, 1817, Eschscholz s.n. (Holotype: LE).

Synonyms: Brava eschscholtziana (Andrzejowski ex de Candolle) Bentham & J. D. Hooker ex S. Watson, Bibl. Ind. North Am. Bot. 1: 51. 1878; Eutrema eschscholtziana (Andrzejowski ex de Candolle) Robinson, Syn. Fl. N. Amer. 1: 135. 1895; Hesperis eschcholtzianus (Andrzejowski ex de Candolle) Kuntze, Revis. Gen. Pl. 2: 935. 1891.

Oreas involucrata Chamisso & Schlechtendal, Linnaea 1: 30. 1826. TYPE: UNITED STATES. "Summis montibus alpinis insulae Unalaschca inter lapides acervatos" (Holotype: ?HAL, n. v.).

Distribution: Alaska and Yukon.

3. Aphragmus involucratus (Bunge) O. E. Schulz in Engler, Pflanzenreich IV. 105(Heft 86): 198. 1924.

Basionym: Platypetalum involucratum Bunge, Verzeichn. Pfl. Altai 77. 1836. TYPE: [RUSSIA]. Altai to Chuya River, Bunge s.n. (Holotype: LE).

Orobium involucratum (Bunge) Bunge, Del. Sem. Hort. Dorpat. 8. 1841; Brava involucrata (Bunge) Ledebour, Fl. Ross. 1: 194. 1842; Hesperis involucrata (Bunge) Kuntze, Revis. Gen. Pl. 2: 935. 1891.

Distribution: Mongolia, Russia (Altai, Siberia).

4. Aphragmus oxycarpus (J. D. Hooker & Thomson) Jafri, Notes Roy. Bot. Gard. Edinburgh 22: 96. 1956.

Basionym: Brava oxycarpa J. D. Hooker & Thomson, J. Linn. Soc., Bot. 5: 169. 1861. TYPE: W. TIBET. Piti prope Lara, 12,000-13,000 ft, T. Thomson s.n. (Holotype: K).

Synonyms: Brava rubicunda Franchet, Bull. Soc. Bot. Fr. 33: 403. 1889. TYPE: CHINA. Yunnan, Mt. Tsang-chan supra Tali, 16 June 1884, Delavay 80 (Holotype: P; Isotype: US).

Aphragmus stewartii O. E. Schulz, Repert. Sp. Nov. Regni Veg. 31: 330. 1933. TYPE: KASHMIR. Mt. Kolahoi, 16,000 ft, August 1927, R. R. Stewart 9393 (Holotype: B; Isotype: NY).

Aphragmus tibeticus O. E. Schulz, Repert. Sp. Nov. Regni Veg. 12: 387. 1922. TYPE: TIBET. Tatsien Lu-Dawo, an Bachlaufen in der Nahe von Schneeflecken der westlichen Parallelkette des Dshara 6stlich Dschungku, 4700 m, 26 June 1924, W. Limpricht 1818a (Holotype: WRSL, n. v.; Isotype: WU).

Eutrema przewalskii Maximowicz, Fl. Tangut. 1: 68. 1889. TYPE: TIBET. 14,000 ft., 7 July? 1884, Przewalski s.n. (Holotype: LE).

Aphragmus oxycarpus var. microcarpa Z. X. An, Bull. Bot. Res., Harbin 1(1 & 2): 102. 1891. TYPE: CHINA. Xinjiang, Tian Shan, Urumqi-Nanshan, June 1965, collector? 6501037 (Holotype: KUN).

Brava foliosa Pampanini, Bull. Soc. Bot. Ital. 1926: 40. 1926. TYPE: KASHMIR. Karakorum, valley above Sciaiok, 4830 m, 12 July 1914, Danielli & Marinelli s.n. (Holotype: FI).

Brava oxycarpa f. glaber Vassilchenko in Komarov, Fl. URSS 8: 74. 1939; Aphragmus oxycarpus var. glaber (Vassilchenko) Z. X. An in T. Y. Cheo, Fl. Reipubl. Popularis Sin. 33: 422. 1987.

Brava oxycarpa var. stenocarpa O. E. Schulz, Notizbl. Bot. Gart. BerlinDahlem 9: 1068. 1927; Aphragmus oxycarpus var. stenocarpus (O. E. Schulz) G. C. Das in B. D. Sharma & N. P. Balakrishnan, Fl. India 2: 226. 1993. TYPE: INDIA. Uttar Pradesh, Tihri Garhwal, Dudu Gadh unter Srikanta, 5000-5300 m, 9 August 1883, J. F. Duthie 898 (Holotype: CAL, n. v.).

Lignariella duthiei Naqshi, J. Econ. Tax. Bot. 3: 976. 1982. TYPE: PAKISTAN. Mt. Kolhoi, above Pahalgam, A. R. Naqshi 5064 (Holotype: KASH, n. v.).

Distribution: Afghanistan, Bhutan, China, India, Kashmir, Nepal, Pakistan, Sikkim, Tajikistan.

5. Aphragmus nepalensis (H. Hara) Al-Shehbaz, comb. nov.

Basionym: Staintoniella nepalensis H. Hara, J. Jap. Bot. 49:196. 1974.

TYPE: W NEPAL. Dolpo, Tarap, sandy shale 29°2' N 83° 11' E, 16,500 ft, 3 July 1963, Stainton 4394 (Holotype: BM; Isotype: E).

Distribution: endemic to Nepal.

ACKNOWLEDGMENTS

I am grateful to the directors and curators of the herbaria cited.

LITERATURE CITED

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