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STUBENDORFFIA AND WINKLERA BELONG TO THE EXPANDED LEPIDIUM (BRASSICACEAE)

Published online by Cambridge University Press:  14 June 2011

I. A. Al-Shehbaz  and
K. Mummenhoff
I. A. Al-Shehbaz
Affiliation:
Missouri Botanical Garden, PO Box 299, St Louis, MO 63166-0299, USA. E-mail: Ihsan.Al-Shehbaz@mobot.org
K. Mummenhoff
Affiliation:
Department of Botany, Faculty of Biology/Chemistry, University of Osnabrück, Barbarastrasse 11, 49069 Osnabrück, Germany.

Abstract

The genus Stubendorffia (Brassicaceae) is distinguished from Lepidium solely by the dehiscent vs. indehiscent angustiseptate fruits. By contrast, Winklera is separated from Lepidium by a combination of perennial habit, pinnatisect leaves, yellow flowers, and wingless fruits, characters all of which occur individually and in various combinations within Lepidium. Extensive molecular studies strongly show that Winklera and polyphyletic Stubendorffia are nested within the earlier-published Lepidium and, therefore, the three genera are herein formally united. The new name Lepidium pavlovii and 10 new combinations, L. afghanicum, L. apterum, L. botschantzevii, L. curvinervium, L. lipskyi, L. olgae, L. orientalis, L. patrinoides, L. pterocarpum, and L. silaifolium, are proposed. Lepidium apterum is lectotypified. A complete generic synonymy of Lepidium and an expanded generic description are presented.

Keywords

Brassicaceaegeneric synonymyLepidiumnew combinationsnew nameStubendorffiatypificationWinklera

Information

Type
Articles
Information
Edinburgh Journal of Botany , Volume 68 , Issue 2 , July 2011 , pp. 165 - 171
Copyright
Copyright © Trustees of the Royal Botanic Garden Edinburgh 2011

Introduction

Lepidium L. is a cosmopolitan genus distributed on all continents except Antarctica (Al-Shehbaz, Reference Al-Shehbaz1986). It is easily distinguished from other genera of the Brassicaceae by the angustiseptate, often dehiscent fruits, one (rarely two) subapical ovules/seeds per locule, and (when present) simple trichomes. Nearly 50% of the species have flowers with two stamens, a feature not found elsewhere in the family. The three species previously assigned to Cardaria Desv. and now in Lepidium sometimes have two ovules per locule, and two of these species, L. chalepense L. and L. appelianum Al-Shehbaz, have inflated instead of angustiseptate fruits. For further discussion on these exceptions, see Al-Shehbaz et al. (Reference Al-Shehbaz, Mummenhoff and Appel2002).

Based on extensive molecular studies (e.g. Mummenhoff, Reference Mummenhoff1995; Mummenhoff et al., Reference Mummenhoff, Kuhnt, Koch and Zunk1995, Reference Mummenhoff, Brüggemann and Bowman2001; Bowman et al., Reference Bowman, Brüggemann, Lee and Mummenhoff1999; Brüggemann, Reference Brüggemann2000) and detailed morphological studies (Appel & Al-Shehbaz, Reference Appel, Al-Shehbaz, Kubitzki and Bayer2003), the limits of Lepidium were expanded by Al-Shehbaz et al. (Reference Al-Shehbaz, Mummenhoff and Appel2002) and Al-Shehbaz (Reference Al-Shehbaz2003) to include all species of Cardaria, Coronopus Zinn, and Stroganowia Kar. & Kir. More recent molecular studies (e.g. Franzke et al., Reference Franzke, German, Al-Shehbaz and Mummenhoff2009; German et al., Reference German, Friesen, Neuffer, Al-Shehbaz and Hurka2009; Mummenhoff et al., Reference Mummenhoff, Polster, Mühlhausen and Theissen2009; Couvreur et al., Reference Couvreur, Franzke, Al-Shehbaz, Bakker, Koch and Mummenhoff2010) further confirmed the above taxonomic conclusions and clearly showed that Winklera Regel (two species) and Stubendorffia Schrenk ex Fisch. (nine species) are nested within Lepidium. These 11 species are transferred herein to Lepidium. As a result, the genus is further expanded to include about 250 species, compared to earlier estimates of 175 species (Al-Shehbaz, Reference Al-Shehbaz1986) and 220 species (Appel & Al-Shehbaz, Reference Appel, Al-Shehbaz, Kubitzki and Bayer2003). Stubendorffia subdidyma N.Busch was transferred earlier to Isatis L. by Avetisian (Reference Avetisian1982), a position with which we agree based on floral and fruit morphology.

The extensive molecular studies above, and others (e.g. Mummenhoff et al., Reference Mummenhoff, Bowman, Linder, Friesen and Franzke2004), amply demonstrated that Coronopus, Stroganowia, and Stubendorffia are polyphyletic and each evolved from various lineages within Lepidium. Despite the overwhelming molecular evidence, some authors (e.g. Dorofeyev et al., Reference Dorofeyev, Korotyaev and Gultekin2004; Holmgren, Reference Holmgren, Holmgren, Holmgren and Cronquist2005; Dorofeyev, Reference Dorofeyev2006; Karpova, Reference Karpova2006; Karpova & Grigoreva, Reference Karpova and Grigoreva2007) still maintain a polyphyletic Stroganowia. In agreement with Rollins (Reference Rollins1982) and Holmgren (Reference Holmgren, Holmgren, Holmgren and Cronquist2005), the western North American Stroganowia tiehmii Rollins (now L. tiehmii (Rollins) Al-Shehbaz) evolved from Central Asian ancestors (Mummenhoff et al., Reference Mummenhoff, Polster, Mühlhausen and Theissen2009).

A comprehensive account of the generic synonyms of Lepidium, a detailed description of the expanded genus, and nomenclatural adjustments resulting from the merging of Stubendorffia and Winklera with Lepidium are presented below.

Expanded Generic Description and Synonymy

  • Lepidium L., Sp. Pl. 2: 643 (1753). – Type: Lepidium latifolium L. (lecto designated by Britton & Brown, Reference Britton and Brown1913: 164).

  • Carara Medik., Pflanzen-Gatt. 34 (1792). – Type: not designated.

  • Cardaria Desv., J. Bot. Agric. 3: 163 (1815). – Type: Cardaria draba (L.) Desv. (= Lepidium draba L.).

  • Cardamon (DC.) Fourr., Ann. Soc. Linn. Lyon ser. 2, 16: 338 (1868). – Type: Cardamon sativum (L.) Fourr. (= Lepidium sativum L.).

  • Coronopus Zinn, Cat. Pl. Hort. Gott. 325 (1757), nom. cons., non Coronopus Mill., Gard. Dict. Abr. ed. 4 (1754), nom. rej. – Type: Coronopus ruellii All. = Cochlearia coronopus L. (= Lepidium coronopus (L.) Al-Shehbaz).

  • Cotyliscus Desv., J. Bot. Agric. 3: 164 (1815). – Type: Cotyliscus niloticus (Delile) Desv. = Cochlearia nilotica Delile (= Lepidium niloticum (Delile) Sieber ex Steud.).

  • Hymenophysa C.A.Mey. ex Ledeb., Icon. Pl. Nov. 2: 20 (1830). – Type: Hymenophysa pubescens C.A.Mey. (= Lepidium appelianum Al-Shehbaz).

  • Kandis Adans., Fam. 2: 422 (1763). – Type: Kandis perfoliata (L.) Kerguélen (= Lepidium perfoliatum L.).

  • Lepicochlea Rojas Acosta, Bull. Acad. Int. Geogr. Bot. 28: 165 (1918). – Type: Lepicochlea americana Rojas Acosta (= Lepidium didymum L.).

  • Monoploca Bunge in Lehm., Pl. Preiss. 1: 259 (1845). – Type: not designated.

  • Nasturtiolum Medik., Pflanzen-Gatt. 82 (1792), non Nasturtiolum Gray, Nat. Arr. Brit. Pl. 2: 692 (1792). – Type: Nasturtiolum castratum Medik., nom. illeg. (= Lepidium didymum L.).

  • Neolepia W.A.Weber, Phytologia 67: 427 (1989). – Type: Neolepia campestris (L.) W.A.Weber (based on Thlaspi campestre L.) (= Lepidium campestre (L.) W.T.Aiton).

  • Papuzilla Ridl., Trans. Linn. Soc. London, Bot. ser. 2, 9: 17 (1916). – Type: Papuzilla minutiflora Ridl. (= Lepidium minutiflorum (Ridl.) Hewson).

  • Physolepidium Schrenk in Fisch. & C.A.Mey., Enum. Pl. Nov. 1: 97 (1841). – Type: Physolepidium repens Schrenk (= Lepidium chalepense L.).

  • Senebiera DC., Bull. Sci. Soc. Philom. Paris 1: 172 (1798). – Type: not designated.

  • Sprengeria Greene, Leafl. Bot. Observ. Crit. 1: 198 (1905). – Type: Sprengeria flava (Torr.) Greene (= Lepidium flavum Torr.).

  • Stroganowia Kar. & Kir., Bull. Soc. Imp. Naturalistes Moscou 14: 386 (1841). – Type: Stroganowia sagittata Kar. & Kir. (lecto designated by Botschantsev, Reference Botschantsev1984: 73) (= Lepidium sagittatum (Kar. & Kir.) Al-Shehbaz).

  • Stubendorffia Schrenk ex Fisch., Index Seminum Hortus Petrop. 9: Suppl. 20 (1844), syn. nov. – Type: Stubendorffia orientalis Schrenk ex Fisch. (= Lepidium orientale (Schrenk) Al-Shehbaz & Mummenhoff).

  • Uranodactylus Gilli, Feddes Repert. Sp. Nov. Regni Veg. 61: 209 (1959). – Type: Uranodactylus afghanicus Gilli (= Lepidium silaifolium (Hook.f. & Thomson) Al-Shehbaz & Mummenhoff).

  • Winklera Regel, Trudy Imp. S.-Peterburgsk. Bot. Sada 9: 617 (1886), non Winkleria Rchb., Deutsch. Bot. Herbarienbuch 236 (1841), syn. nov. – Type: Winklera patrinoides Regel (= Lepidium patrinoides (Regel) Al-Shehbaz & Mummenhoff).

Herbs, annual, biennial, or perennial with caudex, sometimes subshrubs, rarely shrubs or lianas. Trichomes absent or simple. Multicellular glands absent. Stems erect or ascending, sometimes creeping, simple or branched basally and/or apically. Basal leaves rosulate or not, simple, entire or variously toothed or divided and 1–3-pinnatisect; cauline leaves petiolate or sessile, base cuneate, attenuate, auriculate, sagittate, or amplexicaul, margin entire, dentate, or dissected. Racemes often many flowered, ebracteate, corymbose, elongated or not in fruit; fruiting pedicels terete, flattened, or winged, erect to divaricate, persistent. Sepals ovate or oblong, rarely orbicular, free, caducous or persistent, erect to spreading, equal, base of lateral pair not saccate. Petals white, yellow, pink, or purple, erect or spreading, longer to shorter than sepals, sometimes rudimentary or absent; blade obovate, spatulate, oblong, oblanceolate, orbicular, linear, or filiform, apex obtuse, rounded, or emarginated; claw absent or distinct and shorter than sepals, glabrous, unappendaged, entire. Stamens 2 or 4 and equal in length, lateral or median, or 6 and usually tetradynamous; filaments wingless, unappendaged, glabrous, free; anthers ovate or oblong, not apiculate. Nectar glands 2, 4, or 6, distinct; median glands often present. Ovules 2(–4) per ovary; placentation apical. Fruit dehiscent silicles, rarely indehiscent and samaroid or schizocarpic and breaking into two 1-seeded, nutletlike halves, oblong, ovate, obovate, cordate, obcordate, elliptic, orbicular, didymous, ovoid, obovoid, or globose, strongly angustiseptate or 4-angled, rarely inflated and terete, sessile, unsegmented; valves papery, leathery, or woody, veinless or prominently veined, glabrous or pubescent, strongly keeled or rarely rounded, apically winged or wingless, unappendaged, readily releasing or enclosing seed; gynophore absent; replum rounded, visible; septum complete or perforated, not veined; style absent, obsolete, or distinct, included or exserted from apical notch of fruit; stigma capitate, entire or rarely 2-lobed. Seeds 1(or 2) per locule, winged, margined, or wingless, oblong or obovate, plump or flattened; seed coat smooth, minutely reticulate, or papillate, copiously or rarely not mucilaginous when wetted; cotyledons incumbent, rarely accumbent or diplecolobal.

New Combinations and New Name

  • Lepidium afghanicum (Rech.f. & Köie) Al-Shehbaz & Mummenhoff, comb. nov. – Stubendorffia afghanica Rech.f. & Köie, Anz. Österr. Akad. Wiss., Mat.-Naturwiss. 91: 60 (1954), non Lepidium affghanum Boiss., Fl. Orient. 1: 358 (1867). – Winklera afghanica (Rech.f. & Köie) Hedge, Fl. Iranica 57: 74 (1968). – Type: Afghanistan, Farakulum, 3200 m, 21 vii 1948, M.E. Köie 3007 (holo C; iso W!).

Distribution. Afghanistan.

  • Lepidium apterum (Lipsky) Al-Shehbaz & Mummenhoff, comb. nov. – Stubendorffia aptera Lipsky, Trudy Imp. S.-Peterburgsk. Bot. Sada 18: 17 (1900). – Type: Tajikistan/Uzbekistan, Hissar Range, Sari Socho Pass, 7000 ft, 11 vi 1897, V.I. Lipsky 239 (lecto LE!, annotated by V.I. Dorofeyev and formally designated here; isolecto LE!).

Distribution. Tajikistan, Uzbekistan.

  • Lepidium botschantzevii (R.Vinograd.) Al-Shehbaz & Mummenhoff, comb. nov. – Stubendorffia botschantzevii R.Vinograd., Bot. Mater. Gerb. Inst. Bot. Akad. Nauk Uzbeksk. S.S.R. 19: 7 (1974), non Lepidium botschantsevianum Al-Shehbaz, Novon 12: 8 (2002). – Type: Uzbekistan, Pamir Alaj, Alaisky Mt., slope of Yorgan River, Kara Zhora, 18 vi 1963, Aigarova & Ubuneeva s.n. (holo LE!).

Distribution. Uzbekistan.

  • Lepidium curvinervium (Botsch. & Vved.) Al-Shehbaz & Mummenhoff, comb. nov. – Stubendorffia curvinervia Botsch. & Vved., Not. Syst. Herb. Inst. Bot. Acad. Sci. Uzbekistan 13: 14 (1952). – Type: Uzbekistan, Shakhimardan, Shaliang, 1650 m, 26 vi 1949, Shafeev s.n. (holo TASH).

Distribution. Uzbekistan.

  • Lepidium lipskyi (N.Busch) Al-Shehbaz & Mummenhoff, comb. nov. – Stubendorffia lipskyi N.Busch, Trudy Bot. Muz. Imp. Akad. Nauk 10: 125 (1913). – Type: Kazakhstan, Tian Shan Mts., valley west of River Karakol, 23 vi 1908, R. Roshevitz 1359 (holo LE!).

Distribution. Kazakhstan, Uzbekistan.

  • Lepidium olgae (R.Vinograd.) Al-Shehbaz & Mummenhoff, comb. nov. – Stubendorffia olgae R.Vinograd., Bot. Mater. Gerb. Inst. Bot. Akad. Nauk Uzbeksk. S.S.R. 19: 8 (1974). – Type: Uzbekistan, Pamir Alaj, Mt. Nuratau, 15 v 1941, Momotov s.n. (holo TASH).

Distribution. Uzbekistan.

  • Lepidium orientale (Schrenk) Al-Shehbaz & Mummenhoff, comb. nov. – Stubendorffia orientalis Schrenk ex Fisch., Index Seminum Hortus Petrop. 9: Suppl. 21 (1844). – Type: Kazakhstan, Khantau Mts., steppes near Dala-Ghainar stream, A.G. Schrenk s.n. (holo LE!).

Distribution. Kazakhstan, Tajikistan, Uzbekistan.

  • Lepidium patrinoides (Regel) Al-Shehbaz & Mummenhoff, comb. nov. – Winklera patrinoides Regel, Trudy Imp. S.-Peterburgsk. Bot. Sada 9: 617 (1886). – Uranodactylus patrinoides (Regel) Gilli, Feddes Repert. Sp. Nov. Regni Veg. 66: 194 (1962). – Type: Tajikistan? (as Turkestan), Bukhara, Regel s.n. (lecto LE!, designated by Jafri, Reference Jafri, Nasir and Ali1973: 66).

  • Winklera patrinoides Regel subsp. chitralica Jafri, Fl. W. Pakistan 55: 66 (1973). – Type: Pakistan, Chitral, Shah Jinali, 3700 m, 28 vi 1958, Bowes Lyon 1024 (holo BM!).

Distribution. Pakistan, Tajikistan.

  • Lepidium pavlovii Al-Shehbaz & Mummenhoff, nom. nov. – Stroganowia gracilis Pavlov, Bot. Zhurn. (Moscow & Leningrad) 18: 365 (1933), non Lepidium gracile (Chodat & Hassl.) Boelcke, Parodiana 4: 36 (1986). – Stubendorffia gracilis (Pavlov) Botsch. & Vved., Bot. Mater. Gerb. Bot. Inst. Uzbekistansk. Fil. Akad. Nauk S.S.S.R. 3: 20 (1941). – Type: Kazakhstan, Karatau Mt., slopes facing Dschalanasch-ata, near Karnak, 21 v 1931, N. Graz-Gusseva s.n. (holo MW).

Distribution. Kazakhstan.

The species is re-named in honour of its discoverer Nikolai Vasilievich Pavlov (1893–1971).

  • Lepidium pterocarpum (Botsch. & Vved.) Al-Shehbaz & Mummenhoff, comb. nov. – Stubendorffia pterocarpa Botsch. & Vved., Bot. Mater. Gerb. Bot. Inst. Uzbekistansk. Fil. Akad. Nauk S.S.S.R. 3: 20 (1941). – Type: Uzbekistan, W Tian Shan, Valley of river Shatkal, upper banks of river Sandalash, basin of river Ajik-Tash, 2650 m, 14 viii 1938, Piyataeva & Momotov 727 (holo TASH).

Distribution. Uzbekistan.

  • Lepidium silaifolium (Hook.f. & Thomson) Al-Shehbaz & Mummenhoff, comb. nov. – Heldreichia silaifolia Hook.f. & Thomson, J. Linn. Soc., Bot. 5: 176 (1861). – Winklera silaifolia (Hook.f. & Thomson) Korsh., Bull. Acad. Petersb. ser. V, 9: 419 (1898). – Uranodactylis silaifolius (Hook.f. & Thomson) Gilli, Feddes Repert. Sp. Nov. Regni Veg. 66: 194 (1962). – Type: Afghanistan, Siahsung, Griffith s.n. (holo K!).

  • Uranodactylus afghanicus Gilli, Feddes Repert. Sp. Nov. Regni Veg. 61: 209 (1959). – Type: Afghanistan, Band-e-Amir, 2950 m, 29 viii 1951, A. Gilli [1161-b] (holo W).

  • Lepidium hindukushense Kitam., Acta Phytotax. Geobot. 19: 97 (1963). – Type: Pakistan, Hindukush, between Diwangar and Ghizar, 4200 m, 10 viii 1957, K. Ogino 409 (holo KTO).

Distribution. Afghanistan, Pakistan, Tajikistan.

Acknowledgements

We thank the curators and directors of the herbaria cited, and Olga Chernyva (LE) for sending digital images of the type collection of Stubendorffia aptera.

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