A SYNOPSIS OF SMELOWSKIA (BRASSICACEAE) IHSAN A. AL-SHEHBAZ1 AND SUZANNE I. WARWICK 2 Abstract. On the basis of extensive molecular data and comparative morphological studies, the genera Ermania, Gorodkovia, Hedinia, Hediniopsis, Redowskia, Sinosophiopsis, and Sophiopsis, which form a mono- phyletic group with Smelowskia, are herein united with Smelowskia, a recently conserved generic name. Seven new combinations are proposed in Smelowskia: S. bartholomewii, S. czukotica, S. furcata, S. heishuiensis, S. jacutica, S. micrantha, and S. sophiifolia. A key to the 25 species of Smelowskia, as well as a detailed descrip- tion of the expanded genus, major synonymies, and distributions are given. Keywords: Brassicaceae, Ermania, Gorodkovia, Hedinia, Hediniopsis, Redowskia, Sinosophiopsis, Smelowskia, Sophiopsis. Generic delimitation is one of the most fre- The ITS data indicate that Smelowskia, quently encountered problems in the systemat- Ermania, Gorodkovia, Hedinia, Redowskia, ics of the Brassicaceae (Al-Shehbaz, 1984; Sinosophiopsis, and Sophiopsis form a poly- Appel and Al-Shehbaz, 2003), and Smelowskia tomy supported by 90% bootstrap values. By C. A. Mey. is no exception. The genus is dis- contrast, the trnL sequence data place tributed in central and northeastern Asia, west- Smelowskia, Ermania, Gorodkovia, and ern Canada, and neighboring United States, and Redowskia in a polytomous clade supported by the number of species assigned to it has ranged 75% bootstrap values, whereas the genera from 5 to 14 (Schulz, 1924, 1936; Drury and Hedinia, Sinosophiopsis, and Sophiopsis were Rollins, 1952; Botschantzev, 1968; Velichkin, rather weakly separated (54% bootstrap) from 1974, 1979; Rollins, 1993; Mulligan, 2001; that polytomy. All seven genera are weakly dis- Zhou et al., 2001). tinguished by character combinations, and the Schulz (1936) and Appel and Al-Shehbaz interested reader should consult Warwick et al. (2003) considered Smelowskia to be closely (2004) for details on comparative morphology related to the Asian genera Ermania Cham. ex and critical evaluation of individual genera. On Botsch. (1 sp.), Gorodkovia Botsch. & Karav. the basis of morphological, molecular, and (1 sp.), Hedinia Ostenf. (3 spp.), Hediniopsis cytological data and because of the lack of ade- Botsch. & V. V. Petrovsky (1 sp.), Redowskia quate morphological grounds to separate gen- Cham. & Schltdl. (1 sp.), Sinosophiopsis Al- era in this complex, Warwick et al. (2004) Shehbaz (3 spp.), and Sophiopsis O. E. Schulz concluded that the most practical taxonomic (4 spp.). Except for Hediniopsis, these genera account is to reduce the seven genera to one. were subjected to extensive molecular phyloge- Redowskia, the earliest published generic netic and morphological studies by Warwick et name in the complex (see below), is a poorly al. (2004). known genus, whereas Smelowskia, the second Warwick et al. (2004) compared sequence oldest, is well known to most Asian and North data from the nuclear internal transcribed ribo- American botanists. In order to unite all seven somal DNA (ITS) and the chloroplast trnL genera of the complex, one was faced with intron to study the phylogeny of Smelowskia either recognizing Redowskia or conserving and the above genera. Data of both markers Smelowkia. The latter course of action was clearly show that Smelowskia is polyphyletic. taken in a proposal by Al-Shehbaz (2003), and We are grateful to Richard K. Brummitt and Nicholas J. Turland for advice on nomenclatural matters. We wish to thank Tatyana Shulkina for translating some of the Russian literature, and Dmitry Geltman, Vladimir Dorofeyev, Olga Cherneyva, Alica Grabovskaya, and Tatyana Krestovskaya for their help in locating the types of various taxa at LE. We thank the direc- tors and curators of the herbaria cited for allowing us to examine the type collections. Last but not least, we are indebted to Gustavo A. Romero-González for his editorial advice. 1 Missouri Botanical Garden, 2345 Tower Grove, St. Louis, Missouri 63110, U.S.A. E-mail: [email protected] 2 Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Central Experimental Farm, Ottawa, Ontario K1A 0C6, Canada. E-mail: [email protected] Harvard Papers in Botany, Vol. 11, No. 1, 2006, pp. 91–99. © President and Fellows of Harvard College, 2006. 92 HARVARD PAPERS IN BOTANY Vol. 11, No. 1 the proposal was approved by the general com- S. bifurcata, S. calycina, S. inopinata, S. parry- mittee during the nomenclatural sessions of the oides) are central and eastern Asian, and 1 (S. Vienna Botanical Congress, on the basis of the porsildii) grows in both continents. Their data recommendation by the Committee for also showed that the North American varieties Spermatophyta (Brummitt, 2005). of S. calycina sensu Drury and Rollins (1952) As delimited by Drury and Rollins (1952), and Rollins (1993) represent distinct species. Botschantzev (1968), Velichkin (1974, 1979), The generic names Acroschizocarpus Rollins (1993), and Mulligan (2001), the Gombócz, Chrysanthemopsis Rech. f., and generic boundaries of Smelowskia can easily Melanidion Greene are kept as synonyms of accommodate Redowskia. The only differences Smelowskia, following Drury and Rollins between the two are the presence in (1952) and Hedge (1968). In Warwick et al. Smelowskia of entire stigmas and uniseriate (2004), S. holmgrenii Rollins fell with seeds and in Redowskia of slightly lobed stig- Polyctenium Greene in a well-supported clade mas and biseriate seeds (see Warwick et al., separate from Smelowskia, and the authors con- 2004). These features are insufficient for the cur with its recent transfer to a new genus recognition of the two genera as distinct, and Nevada N. H. Holmgren (Holmgren, 2004). there are many other genera of Brassicaceae The present paper deals with nomenclatural (e.g., Draba L., Rorippa Scop., Tropidocarpum adjustments based on the conclusions above, Hook., Vella L.) that exhibit similar or even and it provides an enumeration of all species, greater fruit diversity (Appel and Al-Shehbaz, main homotypic synonymies, distributions, a 2003). key to species, and a description of the Warwick et al. (2004) recognized 12 species expanded Smelowskia. For additional syn- in Smelowskia, 6 of which (S. americana, onymies, the reader should consult S. borealis, S. johnsonii, S. media, S. ovalis, Botschantzev (1968), Velichkin (1974, 1979), S. pyriformis) are North American, 5 (S. alba, Rollins (1993), and Mulligan (2001). TAXONOMY Smelowskia C. A. Mey. in Ledeb., Icon. Pl. 2: Hediniopsis Botsch. & V. V. Petrovsky, 17. 1830, nom. cons. TYPE: S. cinerea C. A. Bot. Zhurn. (Moscow & Leningrad) 71: Mey., nom. illeg. (= S. alba (Pall.) Regel; based 1548. 1986. TYPE: H. czukotica Botsch. on Sisymbrium album Pall.). & V. V. Petrovsky. Synonyms: Acroschizocarpus Gombócz, Bot. Melanidion Greene, Ottawa Naturalist Közlem. 37: 1. 1940. TYPE: A. kolianus 25: 146. 1912. TYPE: M. boreale Gombócz. Greene. Chrysanthemopsis Rech. f., Phyton Redowskia Cham. & Schltdl., Linnaea 1: (Horn) 3: 51. 1951. TYPE: C. koelzii 32. 1826. TYPE: R. sophiifolia Cham. & Rech. f. Schltdl. Ermania Cham. ex Botsch., Bot. Zhurn. Sinosophiopsis Al-Shehbaz, Novon 10: (Moscow & Leningrad) 41: 730. 1956. 340. 2000. TYPE: S. bartholomewii Al- TYPE: E. parryoides (Cham.) Botsch. Shehbaz. (based on Draba parryoides Cham.). For Sophiopsis O. E. Schulz in A. Engler, a discussion on the validation of Ermania Pflanzenreich IV. 105(Heft 86): 346. by Botschantzev (1956), see Al-Shehbaz 1924. LECTOTYPE (here designated): S. (2001). sisymbrioides (Regel & Herder) O. E. Gorodkovia Botsch. & Karav., Bot. Schulz. Mater. Gerb. Bot. Inst. Komarova Akad. Annual or perennial herbs, sometimes pulv- Nauk SSSR 19: 109. 1959. TYPE: G. inate and with well-developed, often branched jacutica Botsch. & Karav. caudex covered with persistent petioles of previ- ous years. Trichomes dendritic, sometimes Hedinia Ostenf. in Hedin, S. Tibet 6(3), mixed with fine, long, simple and forked stalked Bot.: 76. 1922. TYPE: H. tibetica ones. Stems erect, ascending, or decumbent, sin- (Thomson) Ostenf. (based on Hutchinsia gle to many from caudex, simple or branched tibetica Thomson). apically. Basal leaves herbaceous, petiolate, 2006 AL-SHEHBAZ AND WARWICK, SMELOWSKIA (BRASSICACEAE) 93 rosulate, 1- or 2-pinnatisect, sometimes flabel- apex. Nectar glands confluent and subtending late, rarely entire, often densely pubescent. bases of all stamens; median nectaries present or Cauline leaves short petiolate or sessile, not rarely absent. Ovules 4–30 per ovary. Fruit auriculate, entire to pinnatisect, rarely absent. dehiscent silique or silicle, linear, lanceolate, Racemes many flowered, dense or lax, oblong, ovoid, obovoid, ellipsoid, fusiform, or ebracteate or bracteate, often elongated consid- suborbicular, terete, slightly 4-angled, or erably in fruit; rachis straight; fruiting pedicels strongly flattened and latiseptate (flattened par- slender, suberect, ascending, divaricate, to allel to the septum) or angustiseptate (flattened recurved. Sepals ovate to oblong, free, decidu- at a right angle to the septum), not inflated, ses- ous or persistent, ascending to spreading, equal, sile or rarely short stipitate; valves papery, with base of inner pair not saccate, margin membra- an obscure or prominent midvein, often nous. Petals white, creamy white, pink, purple, glabrous, keeled or not, smooth, wingless, unap- or rarely yellow, erect at base with flaring blade, pendaged; replum rounded; septum complete