A synopsis of the genus Parrya (Brassicaceae)

Dmitry German

Parrya nauruaq Al-Shehbaz, JR Grant, R. Lipkin, DF Murray & CL Parker (Brassicaceae) is described from the Seward Peninsula of Alaska. It differs from the other North American species in Parrya R. Brown by its smaller flowers and fruits, and by its fewer seeds per fruit. A key to the four North American species of Parrya is provided.

Six taxa of the central Asian Cruciferae: Chorispora tianschanica Z. X. An, Draba denserosulata Pobed., Erysimum brevifolium Z. X. An, Malcolmia humilis Z. X. An, Solms-laubachia pamirica Z. X. An, and Yinshania albiflora Ma & Y. Z. Zhao var. gobica Z. X. An, are reduced to synonymy of Parrya beketovii Krassn., Draba lanceolata Royle, Erysimum cheiranthoides L. subsp. transiliense (M. Pop.) D. German, Strigosella brevipes (Bunge) Botsch., Chorispora macropoda Trautv. and Smelowskia micrantha (Botsch. & Vved.) Al-Shehbaz & S. I. Warwick, respectively.

KEW BULLETIN VOL. 68: 457 Y 475 (2013) DOI 10.1007/S12225-013-9450-4 ISSN: 0075-5974 (print) ISSN: 1874-933X (electronic) A synopsis of the genus Parrya (Brassicaceae) Ihsan A. Al-Shehbaz1 & Dmitry A. German2,3 Summary. Parrya is broadly circumscribed to include 42 species previously assigned to Achoriphragma, Neuroloma, and Pseudoclausia. An expanded generic description, synopsis of all taxa previously assigned to Parrya, and key to all species herein assigned to it are presented. Fourteen new combinations: Leiospora beketovii (Krasn.) D. A. German & Al-Shehbaz, L. saposhnikovii (A. N. Vassiljeva) D. A. German & Al-Shehbaz, Parrya glabra (Royle) D. A. German & Al-Shehbaz, P. gracillima (Popov ex Botsch. & Vved.) D. A. German & Al-Shehbaz, P. hispida (Regel) D. A. German & Al-Shehbaz, P. mollissima (Lipsky) D. A. German & Al-Shehbaz, P. olgae (Regel & Schmalh.) D. A. German & Al-Shehbaz, P. papillosa (Vassilcz.) D. A. German & Al-Shehbaz, P. pazijae (Pachom.) D. A. German & Al-Shehbaz, P. pjataevae (Pachom.) D. A. German & Al-Shehbaz, P. podlechii (Dvořák) D. A. German & AlShehbaz, P. sarawschanica (Regel & Schmalh.) D. A. German & Al-Shehbaz, P. tschimganica (Popov ex Botsch. & Vved.) D. A. German & Al-Shehbaz, and P. vvedenskyi (Pachom.) D. A. German & Al-Shehbaz] and two new names [P. junussovii D. A. German & Al-Shehbaz and P. lipskyi D. A. German & Al-Shehbaz] are proposed. Thirty-eight names are excluded from Parrya (tribe Chorisporeae) and assigned to various genera of the tribes Anchonieae, Arabideae, Boechereae, Camelineae, Erysimeae, Euclidieae, and Smelowskieae, thus reﬂecting the former broad and artiﬁcial circumscription of the genus. Seven taxa are reduced to synonymy with what follows them in parentheses: Neuroloma botschantzevii Pachom. (Parrya pinnatiﬁda Kar. & Kir.), N. grifﬁthii Botsch., N. kunawarense (Royle ex Regel) Botsch., and P. nudicaulis var. dasycarpa Regel (P. glabra), P. angrenica Botsch. & Vved. (P. albida Popov), P. korovinii A. N. Vassiljeva (P. pulvinata Popov), and P. simulatrix Nikitina (P. alba Nikitina). Lectotypes are designated for 19 names, including four partially typiﬁed before. Key Words. Achoriphragma, Central Asia, Cruciferae, Leiospora, Neuroloma, Pseudoclausia. Introduction The number of species assigned by various authors to Parrya R. Br. has varied substantially. The genus was recognised as monotypic (Botschantzev 1972; Soják 1982), but other authors recognised ten species (Schulz 1936), 22 spp. (Busch in Komarov 1939), 25 spp. (Zhou et al. 2001; Appel & Al-Shehbaz 2003), 30 spp. (Jafri 1973; Rollins 1993), 34 spp. (Al-Shehbaz et al. 2006), 40 spp. (Ovczinnikov & Junussov in Ovczinnikov 1978), or 43 spp. (Al-Shehbaz 2012). Based on this synopsis, Parrya is distributed primarily in central Asia, especially Kazakhstan (21 spp.), Kyrgyzstan (18 spp.), Uzbekistan (17 spp.), Tajikistan (13 spp.), Afghanistan (6 spp.), and China (5 spp.), with the major centre of diversity in Tian Shan and Pamir-Alai. Few species extend their ranges into Pakistan (2 spp.), and Turkmenistan, Iran, India, and Bhutan (1 sp. each). P. nudicaulis (L.) Regel is distributed in Russia (Siberia and Far East extending to eastern part of Arctic Europe) and northern North America (Alaska, northern Canada), but earlier works (e.g., Rechinger 1968; Jafri 1973; Zhou et al. 2001) reported its occurrence in the Himalayas based on an entirely different species (German et al. 2011). Three other species are restricted to North America, including P. arctica R. Br. (Canada: Northwest Territories, Nunavut, Yukon), P. nauruaq Al-Shehbaz et al. (USA: Alaska), and P. rydbergii Botsch. (USA: Utah, Wyoming). The last species was treated by Rollins (1993) as a synonym of P. nudicaulis, but the two species are substantially different (Al-Shehbaz 2010). P. nauruaq is unique in the genus in having silicles instead of siliques (Al-Shehbaz et al. 2007). Although Parrya included a very heterogeneous assemblage of more than 80 species currently assigned to eight tribes, molecular data (German et al. 2011) do not support treatment of the genus as monotypic, as advanced by Botschantzev (1972). The latter author transferred 35 species of Parrya to Accepted for publication 1 May 2013. Published online 22 June 2013 1 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, USAR e-mail: ihsan.al-shehbaz@mobot.org 2 South-Siberian Botanical Garden, Altai State UnivR, Lenina Street 61, 656049 Barnaul, RussiaR 3 Present Address: Department of Biodiversity and Plant Systematics, Im Neuenheimer Feld 345, Centre for Organismal Studies (COS) Heidelberg, Heidelberg University, D-69120 Heidelberg, GermanyR e-mail: oreoloma@rambler.ru © The Board of Trustees of the Royal Botanic Gardens, Kew, 2013 458 Neuroloma Andrz. ex DC. and placed many others in several smaller genera. In our opinion, the characters used by Botschantzev (1972) to separate Parrya from Neuroloma (e.g. ascending and non-saccate vs erect and saccate sepals, fruit valves with a prominent vs obscure midvein and readily vs tardily detached from replum, short vs long style) are unreliable, and the overall morphology supports uniting both genera. Soják (1982) interpreted Neuroloma as a later homonym of Nevroloma Raf. (Poaceae) and proposed Achoriphragma Soják to which he transferred all Neuroloma species sensu Botschantzev (1972) and Pachomova (in Vvedensky 1974). Based solely on morphology, Neuroloma and Achoriphragma were reduced by Appel & Al-Shehbaz (2003) to synonymy with Parrya, a position that was later supported by molecular data (German et al. 2011). Interestingly, the generic type P. arctica, which was treated as the only true species of the genus Parrya by Botschantzev (1972) and Soják (1982), was nested in the molecular study above within the P. nudicaulis subclade and was genetically very slightly different from it. Botschantzev (1972) designated the latter species as the lectotype of Neuroloma, a genus he placed in the tribe Matthioleae while assigning Parrya to the tribe Arabideae. Curiously, Hultén (1971: 42) suspected P. arctica to be conspeciﬁc with P. nudicaulis and representing just a ‘high-arctic extreme’ of it. These are just a few examples illustrating the complexity of Parrya taxonomy and highlighting the need for comprehensive treatment using both morphological and molecular data. Parrya was divided by Ovczinnikov & Junussov (in Ovczinnikov 1978) into three sections, of which sect. Neuroloma was recognised by Vassiljeva (1974) as a genus that she divided into three sections and eight series. However, those infrageneric subdivisions of Parrya were based on artiﬁcially delimited characters, and molecular studies (German et al. 2011) do not support their delimitation or the subdivision of the genus into infrageneric groups. Pseudoclausia was said to differ from Parrya by being densely hirsute or pilose (vs glabrous or rarely pilose), annuals, biennials, or short-lived perennials (vs long-lived cespitose perennials or subshrubs) with usually crisped (vs non-crisped) petals and subterete-quadrangular (vs latiseptate) fruits. However, none of these differences are consistently present in either genus, and German et al. (2011) clearly showed that Pseudoclausia is nested within Parrya. The goals of the present treatment are to provide a comprehensive account of all names previously placed in Achoriphragma, Neuroloma, Parrya and Pseudoclausia, as well as to verify their types and current generic and tribal assignments. A detailed description of the © The Board of Trustees of the Royal Botanic Gardens, Kew, 2013 KEW BULLETIN VOL. 68(3) expanded Parrya and keys to its currently recognised 42 species are presented. Parrya R. Br. (Brown 1823: 10, 195). Type: Parrya arctica R. Br. Achoriphragma Soják (1982: 105). Based on Cardamine nudicaulis L. = Parrya nudicualis (L.) Regel. Neuroloma Andrz. ex DC. (de Candolle: 1824: 156), non Neuroloma Cardot (1911: 50), nom. illegit., nec Nevroloma Raf. (Raﬁnesque 1819: 106). Based on Cardamine nudicaulis L. = Parrya nudicualis (L.) Regel. Pseudoclausia Popov (1955: 18). Based on Chorispora hispida Regel = Parrya hispida (Regel) D. A. German & Al-Shehbaz. Herbs, annual, biennial or perennial with or without well-developed caudices, sometimes scapose, rarely subshrubs. Trichomes simple or absent. Multicellular glands present or absent. Stems erect to ascending, simple or branched, leafy or leaﬂess. Basal leaves petiolate, rosulate, simple, entire, dentate, or pinnately lobed; cauline leaves petiolate or sessile, not auriculate, entire or dentate, rarely pinnatiﬁd, often absent. Racemes corymbose, few to many ﬂowered, lax, ebracteate or rarely lowermost ﬂowers bracteate, elongated considerably in fruit; rachis straight; fruiting pedicels erect to ascending or divaricate, slender or thickened, persistent. Sepals linear or oblong, erect, free, caducous, unequal, glabrous, glandular or not, base of lateral pair usually saccate. Petals purple, pink, lavender, white, or rarely brownish, erect with ﬂaring blade, much longer than sepals; blade obovate to oblanceolate or rarely linear, apex rounded or emarginate; claw subequalling or longer than sepals, glabrous, unappendaged. Stamens 6, strongly tetradynamous or abaxial pair longest; ﬁlaments dilated or not at base, wingless, unappendaged, glabrous, free; anthers oblong or linear, obtuse or rarely apiculate at apex. Nectar glands 2, lateral, annular to semiannular; median glands absent. Ovules 10 – 50 per ovary, placentation parietal. Fruit dehiscent capsular siliques or rarely silicles, linear, oblong, or lanceolate, strongly latiseptate or rarely subterete or 4angled, not inﬂated, sessile or subsessile, persistently attached to pedicel, unsegmented; valves leathery, with a prominent midvein and obscure or distinct lateral and marginal veins, glabrous or glandular, not keeled, smooth or torulose, wingless, unappendaged; gynophore absent or rarely to 1 mm; replum ﬂattened, visible; septum complete, membranous or thickened, translucent or opaque, veinless or with a midvein; style usually distinct, (0.2 –) 0.5 − 7 (− 10) mm, stout or slender, persistent, glabrous; stigma conical to cylindrical, 2-lobed, lobes prominent, connate, decurrent, unappendaged. Seeds uniseriate, winged, suborbicular to oblong or ovate, strongly ﬂattened; seed coat A SYNOPSIS OF THE GENUS PARRYA (BRASSICACEAE) smooth, not mucilaginous when wetted; cotyledons accumbent. x = 7. 459 DISTRIBUTION. North America, Asia (W China, C Asia, Himalayas, Russian Far East, Siberia), E Arctic Europe. Key to Parrya species 1. Annuals, biennials or slightly cespitose perennials; fruits subterete to quadrangular, rarely distinctly ﬂattened, (1 –) 1.5 – 2.5 (− 3) mm wide; fruit valves often thickened, ± corky; stems usually leafy, simple or branched . . . . . . 2 1. Long-lived, often distinctly cespitose perennials or subshrubs; fruits always latiseptate, (2 –) 2.5 – 6 (− 8) mm wide; fruit valves not thickened; stems leaﬂess (rarely few leaved in P. pulvinata), simple . . . . . . . . . . . . . . . . . . . . . . .15 2. Densely to moderately pilose annuals, biennials or short-lived perennials; cauline leaves (1 –) 2 − 17, at least lowermost ones as large as basal leaves; stems often branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Glabrescent perennials; cauline leaves absent or 1 − 3, much smaller than basal leaves; stems simple or rarely few branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 3. Annuals 3 – 10 cm tall; fruits densely hirsute with papillate trichomes distinctly ﬂattened at base; fruiting pedicels 2 − 4 mm; glands minute, barely visible . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23. P. papillosa 3. Biennials or short-lived perennials (8 –) 10 – 120 cm tall; fruits glabrous to sparsely or rarely densely hirsute with trichomes not ﬂattened at base; fruiting pedicels (2.5 −) 4 − 22 mm; glands, if present, large and distinct . . . . . 4 4. Petal blade oblanceolate to obovate, 2 − 5 (− 8) mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Petal blade linear to linear-oblanceolate, 1 − 2 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 5. Fruiting pedicels subhorizontal-reﬂexed; petals brownish; fruits incurved . . . . . . . . . . . . . . 11. P. junussovii 5. Fruiting pedicels suberect to ascending or divaricate; petals purple or lavender; fruits usually straight . . . . . . . . 6 6. Ovules/seeds 40 − 62 per ovary/fruit; anthers 4 − 7 mm; petals 1.7 − 2.5 cm . . . . . . . . . . . .18. P. mollissima 6. Ovules/seeds 60 − 92 per ovary/fruit; anthers 2.5 − 4.5 mm; petals 1.1 − 1.8 cm . . . . . . . . . . . . . . . . . . . . 7 7. Fruits 1 − 1.5 mm wide; stigma 1 − 1.5 mm; seeds 1.3 − 2 × 1 − 1.3 mm . . . . . . . . . . . . . . . . 9. P. gracillima 7. Fruits 1.5 − 2.5 mm wide; stigma 2 − 4 mm; seeds 1.7 − 3 × 1.2 − 1.7 mm . . . . . . . . . . . . . . . . . 14. P. lipskyi 8. Fruiting pedicels and sepals densely pilose with spreading to retrorse straight trichomes, sepal trichomes often crisped; petals sharply bicoloured, with blackish violet distal part of claw and proximal part of blade and white distal part of blade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22. P. olgae 8. Fruiting pedicels glabrous or moderately to rarely densely hirsute or pilose with spreading trichomes, sepal trichomes straight; petals uniformly coloured, if bicoloured then not sharply so and never blackish . . . . 9 9. Sepals, fruiting pedicels, and fruits eglandular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42. P. vvedenskyi 9. Sepals, fruiting pedicels, and fruits sparsely to densely glandular, rarely fruits eglandular . . . . . . . . . . . . .10 10. Fruiting pedicels 7 – 14 mm; cauline leaves 6 − 17; petal claw 7 − 13 mm . . . . . . . . . . . . . . . . 10. P. hispida 10. Fruiting pedicels (2.5 –) 4 − 8 (− 10) mm; cauline leaves 1 − 6; petal claw 6 − 8 mm . . . . . . . . . . . . . . . . .11 11. Basal leaves dentate to sinuate; petals purple, 12 − 15 mm; fruits quadrangular, 1.8 − 2.1 mm wide; fruiting pedicels almost as thick as fruits; stigma 2 − 3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . 33. P. sarawschanica 11. Basal leaves pinnatiﬁd to lyrate; petals lavender, 9 − 11 mm; fruits ﬂattened-subterete, 2 − 2.5 mm wide; fruiting pedicels distinctly narrower than fruits; stigma 1 − 1.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . .40. P. tschimganica 12. Fruiting pedicels slender, 2 − 4.5 cm; S Uzbekistan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27. P. pjataevae 12. Fruiting pedicels stout, 0.5 – 2 cm; S Kazakhstan and N Kyrgyzstan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13. Petals purple, broadly obovate, 2 − 2.5 cm × 6 − 10 mm; fruits torulose, 2.5 – 3 wide; sepals with few subapical trichomes; plants 30 – 60 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38. P. subsiliquosa 13. Petals white to lavender, obovate to oblanceolate, 1 − 1.7 cm × 1.5 − 3 (− 4) mm; fruits not or slightly torulose, 1.5 − 2 mm wide; sepals glabrous; plants 10 – 35 cm tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14. Petals 1 − 1.5 cm × 1.5 − 3 mm, entire; plants usually glandular; N Tian Shan (Chu-Ili mts). . . . . 15. P. longicarpa 14. Petals 1.3 − 1.7 cm × 3 – 3.5 (− 4) mm, emarginate; plants eglandular or rarely sparsely glandular; W Tian Shan (Karatau and W part of Kirghis Alatau) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. P. australis 15. Leaves ﬂeshy; caudex surculose; ﬂowering stems not exceeding leaves; fruiting pedicels horizontal to descending, densely pubescent; fruits often curved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16. P. maidantalica 15. Leaves not or slightly ﬂeshy; caudex not surculose; ﬂowering stems exceeding leaves; fruiting pedicels erect to ascending or divaricate, glabrous to sparsely pubescent; fruits usually straight . . . . . . . . . . . . . . . . . . . . . . . . . .16 16. Subshrubs or if herbs not or loosely cespitose; caudices usually loose with elongated, often woody branches, with or without petiolar remains; petiole base long ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 16. Herbs, usually densely cespitose, sometimes pulvinate; caudices compact, often with petiolar remains; petiole base glabrous or rarely sparsely ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 © The Board of Trustees of the Royal Botanic Gardens, Kew, 2013 460 KEW BULLETIN VOL. 68(3) 17. Subshrubs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 17. Perennial herbs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22 18. Leaf blade sharply differentiated from petiole, obovate, coarsely dentate to sublyrate-pinnatiﬁd, apex obtuse or subacute, long ciliate with coarse straight trichomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6. P. darvazica 18. Leaf blade undifferentiated from petiole, lanceolate, linear or almost subulate, entire or runcinate-incised, apex acute, glabrous or pubescent with thin crisped trichomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 19. Fruits 2 − 3 mm wide; seeds 4 – 5.5 × 1.5 – 2.5 mm; leaves rigid, midvein prominent abaxially; sepals 3 − 5 (− 6) mm, crisped pubescent; anthers oblong, 1 − 1.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. P. albida 19. Fruits 2.5 − 4 mm wide; seeds 5 – 7 × 2 – 3.5 mm; leaves soft, midvein not prominent abaxially; sepals 6 − 11 mm, glabrous, papillose, or with straight trichomes; anthers linear, 2.5 − 4 mm . . . . . . . . . . . . . . . . . . 20 20. Plants densely to moderately glandular; leaves involute; fruiting pedicels 1.8 − 3.5 cm; S Kazakhstan . . . . . 25. P. pazijae 20. Plants eglandular or rarely sparsely glandular; leaves ﬂat; fruiting pedicels 1 − 2.5 cm; SW Kyrgyzstan, S Uzbekistan, Tajikistan, N Afghanistan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21 21. Petals purple to lavender, broadly obovate, 18 − 23 × (4 −) 5 − 7 mm, apex emarginate; fruit valves with a prominent midvein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. P. fruticulosa 21. Petals white, oblanceolate, 9 − 13 × 2 − 3 mm, apex rounded; fruit valves with obscure midvein . . . . 21. P. nuratensis 22. Fruits densely glandular, 2.5 – 5.5 cm, abruptly attenuate into a style 5 – 7 (− 10) mm . . . . . 4. P. asperrima 22. Fruits eglandular or sparsely glandular when young, (3 –) 5 – 13 cm, gradually attenuate into style, if abruptly so then style to 3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 23. Fruits (3.5 –) 4 – 5.5 (− 6) mm wide, abruptly attenuate to a style (0.5 –) 1.5 – 3 mm; petals (8 –) 11 – 15 mm wide; seeds orbicular to broadly ovate, wing equally developed all around . . . . . . . . . . . . . . . . . . . . . . 24 23. Fruits (2 –) 2.5 – 3.5 (− 4) mm wide, gradually attenuate to style (3 –) 4 – 8 (− 10) mm; petals 2 – 8 (− 10) mm wide; seeds oblong, wing predominantly distal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 24. Leaves entire or rarely few with 1 (or 2) proximal pairs of remote teeth or lobules . . . . . . . . . . . . 13. P. lancifolia 24. All or most leaves pinnately lobed to pinnatisect with 2 – 4 pairs of conﬂuent lobes . . . . . . . . 34. P. saurica 25. Plants eglandular; roots and caudex branches brownish or greyish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26 25. Plants often glandular; roots and usually caudex branches black . . . . . . . . . . . . . . . . . . . .17. P. minjanensis 26. Leaves dentate, runcinate, subentire, or rarely subpinnatiﬁd with 2 − 4 pairs of lateral lobes; Kyrgyzstan, Tajikistan, and Uzbekistan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31. P. runcinata 27. Stems usually glabrous; petals white to pale lavender; styles (3 –) 4 – 8 (− 10) mm; at least some leaves runcinate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31. P. runcinata 27. Stems usually with crisped trichomes; petals lavender to violet; styles 3 – 4.5 mm; leaves entire, rarely some pinnatiﬁd or dentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39. P. tianschanica 28. Petals white to lavender, (10 –) 12 – 16 (− 18) × 2 – 6 mm, obovate, slightly emarginate . . . . . . . . . . . . . 1. P. alba 28. Petals violet, (16 –) 18 – 22 × (5 –) 6 – 10 mm, obcordate . . . . . . . . . . . . . . . . . . . . . . . . . 37. P. stenocarpa 29. Sepals 2.5 − 5 mm; petals 6 − 13 mm; fruits obovate to narrowly oblong, 0.8 − 2.5 (− 3.5) cm . . . . . . . . . . . . . . . 30 29. Sepals 5 − 10 mm; petals 13 − 24 mm; fruits linear or rarely narrowly oblong, (2 –) 3 − 6 (− 8) cm . . . . . 32 30. Leaves rigid, coriaceous, prominently veined, apiculate; style c. 3.5 mm; W Tian Shan (Uzbekistan) . . . .35. P. saxifraga 30. Leaves soft, herbaceous, obscurely veined, not apiculate; style 0.2 – 1 mm; North America (Alaska, N Canada) . . . . 31 31. Petals 6 − 7 mm; fruits obovate to oblong, 0.8 − 1.4 (− 1.7) cm × 5 − 7 mm; ovules/seeds 6 − 8 per ovary/fruit; leaves obovate to broadly spatulate; plant densely glandular; W Alaska . . . . . . . . . . . . . . . . . . . . . . . . . . 19. P. nauruaq 31. Petals (8 −) 10 − 13 mm; fruits narrowly oblong, 1 − 2.5 (− 3.5) cm × (3 −) 3.5 − 5 mm; ovules/seeds 14 − 20 per ovary/fruit; leaves linear-oblanceolate; plant eglandular; N Canada . . . . . . . . . . . . . . . . . . . . . 3. P. arctica 32. Petioles undifferentiated from blade; leaves entire, linear to narrowly so, 1 – 3 (− 4.5) mm wide; plants eglandular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 32. Petioles usually differentiated from blade; leaves entire, dentate, serrate, sinuate, or incised, linear-oblanceolate to narrowly obovate, 4 – 35 mm wide; plants glandular or rarely eglandular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 33. Plants pulvinate; fruiting pedicels 0.6 − 1.3 cm; fruits torulose, 2 − 5.5 cm × 2 − 3 mm; stems 3 − 11 cm; leaf blades ﬂat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24. P. pavlovii 33. Plants not pulvinate; fruiting pedicels 2 − 5 cm; fruits not torulose, 5 − 7.8 cm × 3 − 4 mm; stems 9 − 30 cm; leaf blades involute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .29. P. popovii 34. Fruits 2 − 3 (− 3.5) mm wide; seeds up to 3 mm wide; C Asia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .35 34. Fruits 4 − 7 mm wide; seeds 3 – 6 mm wide; N Eurasia, C Asia, North America . . . . . . . . . . . . . . . . . . . . .37 35. Leaves entire to remotely denticulate; plants eglandular or very sparsely glandular with minute glands on petioles and sometimes leaf blades; fruits eglandular . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30. P. pulvinata 35. Leaves serrate to pinnatiﬁd or pinnatisect, rarely sinuate; plants moderately to densely glandular throughout with longstalked large glands (rarely eglandular but then pubescent); fruits usually glandular at least when young . . . . . . 36 © The Board of Trustees of the Royal Botanic Gardens, Kew, 2013 A SYNOPSIS OF THE GENUS PARRYA (BRASSICACEAE) 461 36. Leaves serrate to pinnatiﬁd, with up to 5 pairs of teeth or lobes; petals 4 − 5 mm wide; plants glabrous; Afghanistan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28. P. podlechii 36. Leaves usually regularly runcinate-pinnatiﬁd to pinnatisect, with (3 –) 4 – 7 (− 9) pairs of lobes, rarely shallowly sinuate-incised; petals 4 − 8 mm wide; plants often sparsely pubescent; elsewhere . . . . . . . . . . 26. P. pinnatiﬁda 37. Fruits narrowly oblong, linear-lanceolate, or linear, 5 − 8 mm wide, sometimes constricted between seeds; ovules/seeds 10 − 16 per ovary/fruit; Eurasia and North America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 37. Fruits linear, 4 − 5 mm wide, not constricted between seeds; ovules/seeds 20 − 40 per ovary/fruit; C Asia and Himalayas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 38. Plants pubescent; at least some leaves pinnatiﬁd, pinnatisect or deeply toothed, very rarely all leaves entire; C Asia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41. P. turkestanica 38. Plants glabrous; leaves entire, dentate, serrate, or incised; elsewhere . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 39. Plants often not cespitose; fruiting pedicels (1 −) 1.5 − 4 (− 6) cm; ﬁlaments 6 − 10 mm; Alaska, N Canada, Arctic Europe, Siberia, Russian Far East . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20. P. nudicaulis 39. Plants densely cespitose; fruiting pedicels 0.4 − 1.5 (− 2) cm; ﬁlaments 4 − 6 mm; Utah, Wyoming . . . . .32. P. rydbergii 40. Plants eglandular or distally glandular; fruits not torulose; fruiting pedicels (1.5 −) 2 − 7 (− 10) cm; Afghanistan, Bhutan, China, India, Pakistan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. P. glabra 40. Plants densely glandular throughout or only fruits eglandular; fruits torulose; fruiting pedicels 1 − 3.5 cm; Tajikistan, Uzbekistan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 41. Petals narrowly obovate, 4 − 5 mm wide, apex obtuse; fruits eglandular, seeds oblong, 5 − 7 mm . . . .12. P. kuramensis 41. Petals broadly obovate, 7 − 9 mm wide, apex emarginate; fruits densely glandular; seeds broadly ovate to suborbicular, 3.5 − 5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36. P. schugnana 1. Parrya alba Nikitina (1967: 113); Achoriphragma album (Nikitina) Czerep. (Czerepanov 1995: 126); Neuroloma album (Nikitina) Pachom. in Vved. (Vvedensky 1974: 118). Type: Kyrgyzstan, Central Tian Shan, jugum Ak-Shijrak, jugum Ketmen-Tjube, 11 July 1957, Ajdarova & Ubukeyeva s.n. (holotype FRU). Parrya simulatrix Nikitina (1967: 113), synon. nov. Achoriphragma simulatrix (Nikitina) Soják (1982: 106); Neuroloma simulatrix (Nikitina) Botsch. (Botschantzev 1972: 672), as ‘simulator’. Type: Kyrgyzstan, [Central Tian Shan], N slope of range Moldo-Too, upper reaches of R. Karakeche, left tributary, SE slope with wormwoods, stony soil, 17 May 1959, Z. Arbayeva [I. Sudnitsyna?] s.n. (holotype? FRU!; isotype? LE!). DISTRIBUTION. Kyrgyzstan. NOTES. Botschantzev (1972) treated the simultaneously published Parrya alba and P. simulatrix as questionably conspeciﬁc and placed the former in synonymy with the latter but with a question mark. We support uniting the two into one species, but we give priority to P. alba for two reasons. First, the questioning by Botschantzev of the synonymy with P. alba under P. simulatrix does not give the latter priority as determined in Art. 11.5 (McNeill et al. 2012). Although nothing is said in Art. 11.5 regarding such cases, it is clear that only direct synonymy has taxonomical power which is in agreement with the logic of the Code (conf. Art. 52.2, Note 1). Second, there is indirect evidence that P. simulatrix might not be validly published because more than one specimen may have been considered as a type, as Nikitina (1967: 76, 113) cited in the protologue a type (Arbayeva s.n.) that has not yet been found and may well represent two or more specimens collected on different dates (within 17 – 19 May 1959) and later annotated by her. By contrast, Botschantzev (1972) cited another (Sudnitsyna s.n.) as the type collection, which we examined and veriﬁed. If so, the name P. simulatrix was not validated by Nikitina under Art. 40.1, but a combination with relevant epithet was validly published by Botschantzev in 1972 (as Neuroloma simulatrix Botsch.) ﬁve years after the publication of P. alba. Based on these considerations, we accept the name P. alba for the combined species to avoid any future confusion. 2. Parrya albida Popov in Baranov (1925: 26); Achoriphragma albidum (Popov) Soják (1982: 105); Neuroloma albidum (Popov) Botsch. (Botschantzev 1972: 669). Type: [Uzbekistan], Syr-Darya Prov., Tashkent distr., Mt Tschimgan Major, rocky and stony slopes, 2,700 m, 22 Aug. 1924, [Pavel A.] Baranov 97 (lectotype TASH 9774!; isolectotypes, B × 2 sheets!, BP!, BRNU!, CAS!, K!, LE × 3 sheets!, MHA!, MO!, MW!, P!, TK!, W!), selected by Botschantzev (1972: 669) and Ebel (1999: 75). While designating a lectotype for P. albida, Botschantzev (1972) mentioned two herbaria, TAK (later incorporated in TASH) and LE. A second-step lectotypiﬁcation was done by Ebel who cited the specimen in TASH as the holotype. Parrya angrenica Botsch. & Vved. (Botschantzev & Vvedensky 1941: 14), synon. nov.; Achoriphragma angrenicum (Botsch. & Vved.) Soják (1982: 105); Neuroloma angrenicum (Botsch. & Vved.) Botsch. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2013 462 (Botschantzev 1972: 669). Type: [Uzbekistan], Angren, gorge Itelge, mountain top near village Tumen, not common, [2,900 m], 5 June 1931, Matskevych 59 (holotype TASH 172355!). Parrya villosula Botsch. & Vved. (Botschantzev & Vvedensky 1941: 16); Achoriphragma villosulum (Botsch. & Vved.) Czerep. (Czerepanov 1995: 126); Neuroloma villosulum (Botsch. & Vved.) Botsch. (Botschantzev 1972: 67). Type: [Kyrgyzstan], West Tian Shan, basin of river Chatkal, upper reaches of R. Sandalash, Achik-tash basin, NE slope of Ming-Bulak pass, 2,900 m, 17 Aug. 1938, Pjatayeva & Momotov 1062 (holotype TASH 183948!). KEW BULLETIN VOL. 68(3) DISTRIBUTION. W Tian Shan (Kazakhstan, Kyrgyzstan, Uzbekistan). NOTES. Of the six specimens of the type collection of Parrya asperrima at LE, the one carrying Fedtschenko’s handwriting as Parrya nudicaulis var. asper is taken here as the second-step lectotype. 5. Parrya australis Pavlov (1949: 28); Achoriphragma australe (Pavlov) Czerep. (Czerepanov 1995: 127); Neuroloma australe (Pavlov) Botsch. (Botschantzev 1972: 670). Type: Kazakhstan, Karatau Mts, cliffs along gorge Berkara, close to lake Bijlyu-Kul, 3 May 1939, Pavlov 129 (holotype AA!; isotypes LE!, MW × 2 sheets!). DISTRIBUTION. W Tian Shan (Kazakhstan, Kyrgyzstan, Tajikistan, Uzbekistan). NOTES. Pachomova (in Vvedensky 1974) reduced Parrya villosula to synonymy with P. albida, although she maintained P. angrenica as a distinct species. However, a critical study of the type collections and additional material of all three taxa reveals that they represent marginal forms of a species variable in leaf morphology (entire to serrulate or incised), indumentum (subglabrous to densely puberulent), development of glands (presence on all parts of plant to complete absence), and petal size (6 – 15 × 1.5 − 5 mm) and colour (whitish to violet). The presence of reticulate variation in all characters is the reason we recognise only one polymorphic species easily distinguished from the other subshrubby members of the genus by having leaves with a prominent midvein and crisped pilose sepals. DISTRIBUTION. W Tian Shan (Kazakhstan, Kyrgyzstan). 6. Parrya darvazica Botsch. & Vved. in Botsch. (Botschantzev 1965b: 277); Achoriphragma darvazicum (Botsch. & Vved.) Soják (1982: 105); Neuroloma darvazicum (Botsch. & Vved.) Botsch. (Botschantzev 1972: 670). Type: [Tajikistan], Turkestania, Darvaz: Sary-Dasch [Sagyr-Dascht], rocky cliffs, 2,000 – 2,100 m [6,500 – 7,000 ft], 1 June 1897, Korshinsky s.n. (holotype LE!; isotypes LE × 5 sheets!). DISTRIBUTION. Pamir (Tajikistan). 3. Parrya arctica R. Br. (Brown 1823: 110); Neuroloma arcticum (R. Br.) Spreng. (Sprengel 1825: 888). Type: Canada, Mellville Island, Captain Parry’s voyage, 1820, Parry s.n. (lectotype BM!; isolectotype GH!), selected by Botschantzev (1972: 667). 7. Parrya fruticulosa Regel & Schmalh. in Regel (1877: 237); Achoriphragma fruticulosum (Regel & Schmalh.) Soják (1982: 105); Neuroloma fruticulosum (Regel & Schmalh.) Botsch. (Botschantzev 1972: 670). Type: [Uzbekistan], Zaravschanskaya valley, Mt Aksai, 800 – 2,150 m [2,732 − 6,986 ft], 15 May 1869, Fedtschenko s.n. (holotype LE!; isotype LE!). Of the two sheets at LE, the one with a label is taken here as the type. Parrya fruticulosa var. subintegra Regel & Schmalh. in Regel (1877: 237). Type: same as for the species. DISTRIBUTION. Canada (Northwest Territories, Nuna- DISTRIBUTION. Afghanistan, Kyrgyzstan, Tajikistan, Uz- vut, Yukon). bekistan. NOTE. The ﬁrst record for Afghanistan is discussed under Parrya minjanensis. 4. Parrya asperrima (B. Fedtsch.) Popov (1924: 143); Parrya nudicaulis var. asperrima B. Fedtsch. (Fedtschenko 1904: 384); Achoriphragma asperrimum (B. Fedtsch.) Soják (1982: 105); Neuroloma asperrimum (B. Fedtsch.) Botsch. (Botschantzev 1972: 669). Type: [Kyrgyzstan], W Tian Shan, [cliffs in valley of river] Ashutur, 7 Aug. 1897, Fedtchenko s.n. (lectotype LE!; isolectotypes, LE × 5 sheets!), selected by Botschantzev (1972: 669) and here. Parrya golenkinii Lipsch. & Pavlov in Lipschitz (1936: 319). Type: [Kazakhstan], Kara-tau, NW part, Kurdjuk (Ak-sumbe Mts), NE slope, calcareous rocks, 750 m, 21 May 1934, Tekutiew 103 (holotype MW!; isotype LE!). © The Board of Trustees of the Royal Botanic Gardens, Kew, 2013 8. Parrya glabra (Royle) D. A. German & Al-Shehbaz, comb. nov. Type: Northwest India, ‘Nako in Kunawar’, Royle s.n. (holotype K (n.v.)). Perhaps it is the same collection on which the synonym Hesperis kunawarensis is based. http://www.ipni.org/urn:lsid:ipni.org:names:77128837-1 Hesperis glabra Royle, Ill. Bot. Himal. Mts: 72 (1834), non Schur (1853: 65), nec Boiss. & Noë in Boiss. (Boissier 1856: 22), nec Kuntze (1891: 936). A SYNOPSIS OF THE GENUS PARRYA (BRASSICACEAE) Hesperis kunawarensis Royle ex Regel (1870: 272), synon. nov. Neuroloma kunawarense (Royle ex Regel) Botsch. (Botschantzev 1972: 670). Type: Northwest India, no locality data, Royle s.n. (holotype LE (n.v.); isotype K!). Parrya nudicaulis var. dasycarpa Regel (1870: 260), synon. nov. Type: Tibet Occ[identalis]. Regio alp[ina], 4,250 – 5,500 m [14,000 – 18,000 ft], Thomson s.n. (lectotype LE!; isolectotype LE!), selected here. Neuroloma grifﬁthii Botsch. (Botschantzev 1972: 670), synon. nov. Type: Afghanistan, without locality, Grifﬁth 1351 (holotype LE (n.v.); isotypes K!, P!, W!). Two sheets collected by Grifﬁth from Afghanistan are at K. The unnumbered one from the Lemann Herbarium is a possible isotype. The other carries two numbers, ‘1523 Affghanistan Grifﬁth’ [sic] and ‘1351 Cat. Griff.’. The latter sheet is the isotype. Unfortunately the holotypes of Hesperis kunawariensis and Neuroloma grifﬁthii, both at LE (see Botschantzev 1972), were lost or misplaced, and despite several attempts, neither could be found (pers. com., Rudolf Kamelin, Olga Cherneva, and Vladimir Dorofeyev). DISTRIBUTION. Afghanistan, Bhutan, China (Sichuan, Tibet), India, Pakistan. 9. Parrya gracillima (Popov ex Botsch. & Vved.) D. A. German & Al-Shehbaz, comb. nov. Type: [Uzbekistan], along river Ugam, near Khumsan, 16 − 17 May 1922, Popova s.n. (holotype TASH 75032!). 463 Clausia hispida var. leiocarpa Lipsky (1904: 43). Type: [Tajikistan], Chodschent, 1880, Regel s.n. (lectotype LE!; isolectotype LE!), selected here. DISTRIBUTION. W Tian Shan, NW Pamir-Alai (Kazakh- stan, Tajikistan, Uzbekistan). 11. Parrya junussovii D. A. German & Al-Shehbaz, nom. nov. Type: N Tajikistan, ‘ad declive australe jugi Kuraminici, 4 km supra pagum Pangaz, in vicinis foinae Czorby, ad saxa’, 800 m, 24 June 1970, Astanova, Czukavina & Kinzikayeva 3360 (holotype TAD!). http://www.ipni.org/urn:lsid:ipni.org:names:77128840-1 Pseudoclausia kuramensis Ovcz. & Junussov in Ovczinnikov, Fl. Tadzhikskoi SSR. 5: 627 (1978), non Parrya kuramensis Botsch. (Botschantzev 1965b: 278). DISTRIBUTION. W Tian Shan (Tajikistan). 12. Parrya kuramensis Botsch. (Botschantzev 1965b: 278); Achoriphragma kuramense (Botsch.) Soják (1982: 106); Neuroloma kuramense (Botsch.) Botsch. (Botschantzev 1972: 670). Type: Tajikistan, South slope of Kuraminsky range, upper reaches of R. Pangaz, N slope, between boulders, 8 Aug. 1947, Sidorenko 167 (holotype LE!; isotype LE!). DISTRIBUTION. W Tian Shan (Tajikistan, Uzbekistan). http://www.ipni.org/urn:lsid:ipni.org:names:77128838-1 Clausia gracillima Popov ex Botsch. & Vved., Bot. Mater. Gerb. Bot. Inst. Uzbekistansk. Fil. Akad. Nauk S.S.S.R. 3: 13 (Botschantzev & Vvedensky 1941); Pseudoclausia gracillima (Popov ex Botsch. & Vved.) A. N. Vassiljeva (1961: 244). DISTRIBUTION. W Tian Shan (Kazakhstan, Kyrgyzstan, 13. Parrya lancifolia Popov (1938: 86); Achoriphragma lancifolium (Popov) Soják (1982: 106); Neuroloma lancifolium (Popov) Botsch. (Botschantzev 1972: 670). Type: [Kazakhstan], Kungey Alatau, gorge of R. ChetMerke, alpine belt, N gravelly slopes with Kobresia, 1 July 1937, Goloskokov s.n. (lectotype AA!; isolectotypes AA!, LE × 2 sheets!), selected by Goloskokov (1963: 22, as type). Tajikistan, Uzbekistan). DISTRIBUTION. Saur; N, C and E Tian Shan (China (Xinjiang), Kazakhstan, Kyrgyzstan). 10. Parrya hispida (Regel) D. A. German & Al-Shehbaz, comb. nov. Type: [Kazakhstan], Turkestan, prope Tschemken, Sewerzow s.n. (lectotype LE!), selected here. 14. Parrya lipskyi D. A. German & Al-Shehbaz, nom. nov. http://www.ipni.org/urn:lsid:ipni.org:names:77128839-1 http://www.ipni.org/urn:lsid:ipni.org:names:77128841-1 Chorispora hispida Regel, Bull. Soc. Imp. Naturalistes Moscou 43: 266 (1870); Diptychocarpus hispidus (Regel) Regel & Schmalh. in Regel (1877: 230); Clausia hispida (Regel) Lipsky (1904: 43); Pseudoclausia hispida (Regel) Popov (1955: 18). Clausia hispida var. lasiocarpa Lipsky (1904: 43). Type: same as for the species (holotype, LE!). Clausia turkestanica Lipsky, Trudy Imp. S.-Peterburgsk. Bot. Sada 23: 41 (1904), non Parrya turkestanica (Korsh.) N. Busch in Kom. (Komarov 1939: 644); Pseudoclausia turkestanica (Lipsky) A. N. Vassiljeva (1961: 244). Type: [Uzbekistan, Gissar range], Bukhara, Shakhrisyabs, Shut, 2,050 m [6,800 ft], 6 © The Board of Trustees of the Royal Botanic Gardens, Kew, 2013 464 June 1896, Lipsky 233 (lectotype LE!; isolectotype LE!), selected by Botschantzev (in sched.) and here. Clausia turkestanica var. glandulosissima Lipsky (1904: 43). Type: [Uzbekistan, Gissar range], Bukhara. Shakhrisyabs. Gorge of river Gul’bas, 2,450 – 2,750 m [8,000 – 9,000 ft], 10 June 1896, Lipsky 235 (holotype LE!). Lipsky mentioned two plants from one locality both of which are mounted on one sheet and thus represent a single specimen. Clausia turkestanica var. subintegrifolia Lipsky (1904: 42). Type: [Turkmenistan], Gaudan, stony tops and slopes, 13 April 1895, Korshinsky s.n. (lectotype LE!; isolectotypes LE × 2 sheets!), selected here. DISTRIBUTION. Afghanistan, China (Xinjiang), Iran, Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, Uzbekistan. 15. Parrya longicarpa Krasn. (Krasnov 1887: 18); Achoriphragma longicarpum (Krasn.) Soják (1982: 106); Neuroloma longicarpum (Krasn.) Botsch. (Botschantzev 1972: 671). Type: [Kazakhstan], [Near Mt] Kuldja Bassy, [14 April] 1886, Krasnov (as Krassnow) s.n. (lectotype LE!; isolectotypes, K, LE × 2 sheets!, P!), selected by Botschantzev (1972: 671, as type) and here. Botschantzev indicated the collection date as ‘10 – 25 IV 1886’, although only one of the three Krasnov sheets at LE bears the date ‘14 IV’, and the others were not dated. All plants were collected from the same locality and apparently represent one gathering. The sheet with a specimen bearing fruits of the previous year is taken here as the second-step lectotype. Parrya siliquosa Krasn. (Krassnoff 1888: 12). Type: same as that of P. longicarpa (lectotype LE!), selected here. DISTRIBUTION. Tian Shan (Kazakhstan). 16. Parrya maidantalica Popov & P. A. Baranov (1923: 175); Achoriphragma maidantalicum (Popov & P. A. Baranov) Soják (1982: 106); Christolea maidantalica (Popov & P. A. Baranov) N. Busch in Kom. (Komarov 1939: 330); Neuroloma maidantalicum (Popov & P. A. Baranov) Botsch. (Botschantzev 1972: 671); Leiospora maidantalica (Popov & P. A. Baranov) Kamelin (1998: 22). Type: [Kazakhstan], Syr-Daryinsakaya province, Talasskiy Alatau, Maidantal Pass, alpine zone, [2,750 − 3,050 m], 5 Aug. 1921, Abolin & Popov 8848 (lectotype TASH 74978!; isolectotypes LE!, TASH 74979!, TASH 74977!), selected by Botschantzev (1972: 671) and here. KEW BULLETIN VOL. 68(3) the most complete one is designated here as a secondstep lectotype. 17. Parrya minjanensis Rech. f. (Rechinger 1951: 62); Neuroloma minjanensis (Rech. f.) Botsch. (Botschantzev 1972: 671); Parrya stenocarpa var. minjanensis (Rech. f.) Kitam. (Kitamura 1964: 76). Type: Afghanistan, Minjan Pass, 3,650 m [12,000 ft], 26 July 1937, Koelz 12679 (holotype W!; isotype US!). Parrya stenocarpa var. pinnatisecta O. E. Schulz (1927: 1092). Type: Pakistan, Chitral, 3,300 m, 3 June 1895, Harriss 15910 (lectotype K!), selected here. Parrya chitralensis Jafri (1956: 115). Type: Pakistan, Chitral, 3,300 m, 3 June 1895, Harriss 15910 (holotype K!). Parrya stenocarpa subsp. gilgitica Jafri (1956: 115). Type: Kashmir, Gilgit, Tin pass, 3,600 – 4,200 m, 12 July 1886, Giles 476 (holotype K!). Parrya chitralensis Rech. f. (Rechinger 1959: 41). Type: Pakistan. Chitral. Barum Göl, Marmano Shal, 3,700 m, 19 June 1950, Wendelbo s.n. (holotype W!; isotype O). DISTRIBUTION. Afghanistan, Pakistan. NOTES. In its cespitose habit and foliage, Parrya minjanensis resembles P. stenocarpa, especially in having pinnately divided older leaves and entire younger ones. Indeed, Jafri (1973) was correct in considering the two species as closely related based solely on morphology. However, the two species differ substantially in ﬂower size, and P. minjanensis has smaller sepals 5 − 7.5 (vs 8 − 10) mm, petals 15 − 18 × 3 − 5 (vs 20 − 25 × (6 −) 7 − 11) mm, and anthers 2.8 − 3.2 (vs 3.5 − 4) mm, always eglandular (vs sparsely to densely glandular), and appears to be restricted to Afghanistan and Pakistan (vs China, Kazakhstan and Kyrgyzstan). However, the limits of P. minjanensis were confused by Rechinger (1968) who cited from Afghanistan at least three collections of the shrubby P. fruticulosa. These include Podlech 11188 (MSB, W), Podlech 11715 (M, W), and Hedge & Wendelbo 5218 (E, M, O). The last species occurs in the neighbouring Tajikistan and Uzbekistan, and the collections represent its ﬁrst recording from Afghanistan although it is somewhat disjunct. 18. Parrya mollissima (Lipsky) D. A. German & AlShehbaz, comb. nov. Type: [Uzbekistan], Saylyk in Tashkent Alatau, 1,500 m [5,000 ft], 1 – 15 April 1882, Regel s.n. (lectotype LE!; isolectotypes LE × 3 sheets!), selected here. DISTRIBUTION. W Tian Shan (Kazakhstan, Kyrgyzstan, http://www.ipni.org/urn:lsid:ipni.org:names:77128842-1 Uzbekistan). NOTE. Botschantzev (loc. cit.) mentioned ‘type’ (i.e., lectotype) in TAK (= TASH) and LE. Of three specimens with identical label information in TASH, Clausia mollissima Lipsky, Trudy Imp. S.-Peterburgsk. Bot. Sada 23: 44 (1904); Pseudoclausia mollissima (Lipsky) A. N. Vassiljeva (1961: 243). © The Board of Trustees of the Royal Botanic Gardens, Kew, 2013 A SYNOPSIS OF THE GENUS PARRYA (BRASSICACEAE) DISTRIBUTION. W Tian Shan (Kazakhstan, Uzbekistan). NOTE. The sheet with two complete plants and ﬁve tips of racemes is taken here as the lectotype. 19. Parrya nauruaq Al-Shehbaz, J. R. Grant, R. Lipkin, D. F. Murray & C. L. Parker (2007: 277). Type: USA, Alaska. Nome Quadrangle, Seward Peninsula, 12 km NNE of Cape Rodney, Moon Mts, 64º44'N, 166º20'W, 0 − 100 m, 8 June 2005, Parker, Lipkin, & Meyers 15987 (holotype ALA!; isotype MO!). DISTRIBUTION. USA (Alaska). 20. Parrya nudicaulis (L.) Regel (1862: 176); Cardamine nudicaulis L. (Linnaeus 1753: 654); Achoriphragma nudicaule (L.) Soják (1982: 106); Arabis nudicaulis (L.) DC. (de Candolle 1821: 240); Matthiola nudicaulis (L.) Trautv. (Trautvetter 1871: 51); Neuroloma nudicaule (L.) DC. (de Candolle 1824: 156); Parrya nudicaulis (L.) Boiss. (Boissier 1867: 159), comb. superﬂ. Type: ‘Habitat in Sibiria, D. Gmelin’ (lectotype Herb Linn. No. 835.4 (LINN)), selected by Botschantzev (1972: 671). Arabis grandiﬂora L. (Linnaeus 1753: 665). Type: ‘Habitat in Sibiria’ (lectotype Herb Linn. No. 842.3 (LINN)), selected by Marhold in Cafferty & Jarvis (2002: 531). Hesperis arabidiﬂora DC. (de Candolle 1821: 454), nom. illegit.; Neuroloma arabidiﬂorum (DC.) DC. (de Candolle 1824: 156), nom. illegit.; Parrya arabidiﬂora (DC.) Sweet (1830: 24), nom. illegit. Type: same as that of Arabis grandiﬂora L. Hesperis arabidiﬂora var. aspera DC. (de Candolle 1821: 454); Neuroloma arabidiﬂorum var. asperum (DC.) DC. (de Candolle 1824: 156); Parrya macrocarpa var. aspera (DC.) Hook. (Hooker 1829: 47). Type: n.v. Hesperis arabidiﬂora var. glabra DC. (de Candolle 1821: 454); Neuroloma arabidiﬂorum var. glabrum (DC.) DC. (de Candolle 1824: 156); Parrya macrocarpa var. glabra (DC.) Hook. (Hooker 1829: 47). Type: same as that of Arabis grandiﬂora L. Cardamine articulata Pursh (1814: 439). Type: n.v. Parrya macrocarpa R. Br. (Brown 1823: 12), nom. illegit. Type: same as that of Cardamine nudicaulis. Cheiranthus scapiger Adams (1817: 112); Hesperis scapigera (Adams) DC. (de Candolle 1821: 454); Neuroloma scapigerum (Adams) DC. (de Candolle 1824: 156); Parrya scapigera (Adams) G. Don (1831: 173). Type: [Russia, Yakutia, mouth of Lena, Bykovsky peninsula], ad Lenam [Adams s.n.] (lectotype LE!), selected here. Parrya nudicaulis subsp. interior Hultén (1945: 890); P. nudicaulis var. interior (Hultén) B. Boivin (1966: 644). Type: USA, Alaska, near Coldfoot, on the high divide of the northeast between the Yukon River and the Arctic Sea, July − Aug. 1904, Swift 1 (holotype GH!). 465 Parrya nudicaulis subsp. septentrionalis Hultén (1968: 67). Type: [USA, Alaska], Cape Beaufort. 5 – 8 Aug. 1961, Hultén s.n. (holotype n.v.). Parrya nudicaulis var. grandiﬂora Hultén (1945: 892). Type: Alaska, McKinley, Park Scamman 631 (holotype n.v.). Parrya linnaeana Ledeb. (Ledebour 1841: 131), nom. illegit. Type: same as that of Arabis grandiﬂora L. Parrya ajanensis N. Busch in Kom. (Komarov 1939: 646); Achoriphragma ajanensis (N. Busch) Soják (1982: 105); Neuroloma ajanense (N. Busch) Botsch. (Botschantzev 1972: 669); Type: [Russia, Far East, Khabarovsky krai], Ayansky district, vicinities of Ayan, valley of R. Kilkinoy, 20 Aug. 1935, Vasilyev 334 (lectotype LE!; isolectotype LE!), selected here. Busch (loc. cit.) cited the type as: ‘Ajan, marshy meadow at river Sivakcza, V. Vassiljev; in herb. Ac. Sc. URSS conservatur’ and gave no date or collection number. Three Vasilyev collections from Ayan are at LE, only the above ﬁt the diagnosis of P. ajanensis, and the sheet annotated by Busch in February 1937 is designated herein as lectotype. DISTRIBUTION. Canada, Russia (eastern part of Arctic Europe, Siberia, Far East), USA (Alaska). 21. Parrya nuratensis Botsch. & Vved. (Botschantzev & Vvedensky 1941: 15); Achoriphragma nuratense (Botsch. & Vved.) Soják (1982: 106); Neuroloma nuratense (Botsch. & Vved.) Botsch. (Botschantzev 1972: 671). Type: [Uzbekistan], Nuratinskiye Mts, Takhku-Tau, gravelly slopes, 7 May 1913, Korovin s.n. (holotype TASH 74970!; isotypes LE!, TASH 74968!, TASH 74969!). DISTRIBUTION. NW Pamir-Alai (Uzbekistan). NOTE. More collections are desirable to conﬁrm distinctness of Parrya nuratensis from P. fruticulosa. 22. Parrya olgae (Regel & Schmalh.) D. A. German & AlShehbaz, comb. nov. Type [Uzbekistan], Zaravschanskaya valley, Mt Aksai, 850 – 2,150 m [2,732 − 6,986 ft], 15 May 1869, Fedtschenko s.n. (holotype LE!). http://www.ipni.org/urn:lsid:ipni.org:names:77128843-1 Diptychocarpus olgae Regel & Schmalh., Trudy Imp. S.Peterburgsk. Bot. Sada 5: 230 (1877); Clausia olgae (Regel & Schmalh.) Lipsky (1904: 44); Pseudoclausia olgae (Regel & Schmalh.) Botsch. (Botschantzev 1965a: 125). DISTRIBUTION. NW Pamir-Alai (Tajikistan, Uzbekistan). 23. Parrya papillosa (Vassilcz.) D. A. German & Al-Shehbaz, comb. nov. Type: [Kazakhstan]. Turkestansky district, Mts Karatau, stony plateau Ak-kuz, 25 May 1930, Ju. Lipschitz 330 (holotype LE!; isotypes MW × 2 sheets!, TASH!). © The Board of Trustees of the Royal Botanic Gardens, Kew, 2013 466 http://www.ipni.org/urn:lsid:ipni.org:names:77128844-1 Clausia papillosa Vassilcz. in Komarov, Fl. U.R.S.S. 8: 641 (1939); Pseudoclausia papillosa (Vassilcz.) A. N. Vassiljeva (1961: 246). DISTRIBUTION. W Tian Shan (Kazakhstan). 24. Parrya pavlovii A. N. Vassiljeva (1969: 31); Parrya linearifolia Pavlov (1949: 29), non P. linearifolia W. W. Sm. (Smith 1920: 219); Achoriphragma pavlovii (A. N. Vassiljeva) Soják (1982: 106); Neuroloma pavlovii (A. N. Vassiljeva) Botsch. (Botschantzev 1972: 671). Type: Kazakhstan, Dzam[bulskaya] prov., Karatau Mt, in crevices of cliffs above river Taldy-Bulyk, close to Aktogai, 1 June 1948, Pavlov 113 (holotype AA!; isotype MW!). DISTRIBUTION. W Tian Shan (Kazakhstan). 25. Parrya pazijae (Pachom.) D. A. German & AlShehbaz, comb. nov. Type: [Kazakhstan], W Tian Shan, Karatau Mts, rocky shallow soil SW slope, Kara-Sai, 14 May 1934, Pjatayeva 39 (holotype TASH 125437!). http://www.ipni.org/urn:lsid:ipni.org:names:77128845-1 Neuroloma pazijae Pachom., Bot. Mater. Gerb. Inst. Bot. Akad. Nauk Uzbeksk. S.S.R. 19: 44 (Pachomova 1974); Achoriphragma pazijae (Pachom.) Soják (1982: 106). DISTRIBUTION. W Tian Shan (Kazakhstan). 26. Parrya pinnatiﬁda Kar. & Kir. (Karelin & Kirilow 1842: 147); Achoriphragma pinnatiﬁdum (Kar. & Kir.) Soják (1982: 106); Neuroloma pinnatiﬁdum (Kar. & Kir.) Botsch. (Botschantzev 1972: 672). Type: ‘In ﬁssuris rupium summarum alpinum Alatau ad ﬂ. Lepsa et Sarchan’, [June] 1841, Karelin & Kirilov 1198 (lectotype LE!; isolectotypes B!, K!, KFTA!, KW!, LE × 5 sheets!, M!, MW!, P × 2 sheets!, PRC!), selected by Botschantzev (1972: 672) and Gubanov et al. (1998: 29). Parrya pinnatiﬁda var. kizyl-arti Korsh. (Korshinsky 1898: 409). Type: [Kyrgyzstan], Turkestania. On cliffs along R. Kizyl-Art, 3,500 – 3,650 m [11,500 − 12,000 ft], 12 July 1895, Korshinsky 267 (lectotype LE!; isolectotypes LE × 2 sheets!), selected here. Parrya pinnatiﬁda var. oligadenia Trautv. (Trautvetter 1860: 97). Type: [Kazakhstan Dzhungaria], Dshabyk, 20 July 1841, Schrenk s.n. (Herb. Trautvetter 8913) (lectotype LE!; isolectotypes LE × 2 sheets!, P!), selected by German (2012a: 128). Neuroloma botschantzevii Pachom. (Pachomova 1974: 43), synon. nov. Achoriphragma botschantzevii (Pachom.) Soják (1982: 105). Type: Kyrgyzstan, Osh province, © The Board of Trustees of the Royal Botanic Gardens, Kew, 2013 KEW BULLETIN VOL. 68(3) Alay range, Kurbel, 7 km S Khaidarkan, Juniperus forest on cliffs, 28 [23] June 1968, Arbayeva, Ubukeyeva, Mursaliev & Sultanova s.n. (holotype LE!; isotype FRU!). The collection date on the printed holotype label is 28 June, but that of the hand-written label of the isotype is 23 June. DISTRIBUTION. China (Xinjiang, Tibet), Kazakhstan, Kyrgyzstan, Tajikistan, Uzbekistan. NOTES. Parrya pinnatiﬁda is easily recognised by its regularly runcinate-pinnatiﬁd to pinnatisect leaves with (3 or) 4 – 7 (− 9) pairs of somewhat conﬂuent, oblong-triangular lobes. This morphology is rather stable throughout the species range except for in the Alai and Trans-Alai ranges, where the variation encompasses forms with deeply pinnatisect leaves and oblong to linear lobes, as well as forms with shallowly incised, toothed or even subentire leaves. The latter form was described as Neuroloma botschantzevii, but the presence of intermediates and the molecular data (German unpubl.) justiﬁes the merge with P. pinnatiﬁda. 27. Parrya pjataevae (Pachom.) D. A. German & AlShehbaz, comb. nov. Type: [Uzbekistan], Gissarsky range, basin of Tupolang River, right bank of its tributary Tamshush, NNE and NNW rocky slopes, 10 July 1948, Pjatayeva 881 (holotype TASH!). http://www.ipni.org/urn:lsid:ipni.org:names:77128846-1 Neuroloma pjataevae Pachom., Bot. Mater. Gerb. Inst. Bot. Akad. Nauk Uzbeksk. S.S.R. 19: 47 (Pachomova 1974); Achoriphragma pjataevae (Pachom.) Soják (1982: 106). DISTRIBUTION. NW Pamir-Alai (Uzbekistan). 28. Parrya podlechii (Dvořák) D. A. German & Al-Shehbaz, comb. nov. Type: Afghanistan, Kakhar, Khost-o-Fereng, NE Kotali-i-Yawnu, Kalatal, 3,200 m, 10 July 1965, Dieter Podlech 11718 (holotype M!; isotypes MSB!, W!). http://www.ipni.org/urn:lsid:ipni.org:names:77128847-1 Clausia podlechii Dvořák in Rech. f., Fl. Iran. 57: 275 (Rechinger 1968). DISTRIBUTION. Afghanistan. NOTES. Rechinger (1968) cited one collection, Hedge & Wendelbo 5208 (E, M, O), under Parrya nudicaulis that was placed in the tribe Matthioleae, but this collection clearly belongs to P. podlechii, a species placed in that account in Clausia of the tribe Hesperideae. 29. Parrya popovii Botsch. (Botschantzev 1965b: 280); Achoriphragma popovii (Botsch.) Soják (1982: 106); A SYNOPSIS OF THE GENUS PARRYA (BRASSICACEAE) Neuroloma popovii (Botsch.) Botsch. (Botschantzev 1972: 672). Type: [Kazakhstan], Zailiysky Alatau, western part of Mt Turaigyr, E of Mt Zhanalan, N slope, 2,400 m, on rocks, 31 May 1953, Goloskokov s.n. (holotype LE!; isotypes AA!, LE!). Parrya nudicaulis var. linearifolia Regel (1870: 257). Type: [Kazakhstan, Kungei Alatau, in the valley of river Merke, Semenov s.n.] (holotype LE!). DISTRIBUTION. N & W Tian Shan (Kazakhstan, Kyrgyzstan). 30. Parrya pulvinata Popov in Kom. (Komarov 1939: 646); Achoriphragma pulvinatum (Popov) Soják (1982: 106); Neuroloma pulvinatum (Popov) Botsch. (Botschantzev 1972: 672). Type: Kazakhstan, Talasski Alatau, valley of river Kish-Aksu, near mouth of river Bugulen-sai, 2,800 m, 11 July 1933, Linczevski s.n. (holotype LE!). Parrya korovinii A. N. Vassiljeva (1969: 23), synon. nov. Achoriphragma korovinii (A. N. Vassiljeva) Soják (1982: 105); Neuroloma korovinii (A. N. Vassiljeva) Botsch. (Botschantzev 1972: 670). Type: Kyrgyzstan, [West Tian Shan], alpine belt of Talassky Alatau, pass from Bish-Tash to Karagoin, on marble, 30 July 1922, Korovin 1961a (holotype TASH 75025!). Parrya minuta Botsch. & Vved. (Botschantzev & Vvedensky 1948: 9); Achoriphragma minutum (Botsch. & Vved.) Czerep. (Czerepanov 1995: 126); Neuroloma minutum (Botsch. & Vved.) Botsch. (Botschantzev 1972: 671). Type: [Kazakhstan], Syr-Daryinsakaya province, Tashkentsky district, Talassky Alatau, mountain close to source of river Maindantal, 17 Aug. 1922, Baranov 1775 (holotype TASH!). DISTRIBUTION. N, C & W Tian Shan (Kazakhstan, Kyrgyzstan, Uzbekistan). NOTES. In her original description of Parrya korovinii, Vassiljeva (1969) distinguished the species from P. pulvinata on minor differences in ‘habit [robust plant 22 cm alt.], form and dimentions of leaf blades [oblong, 4 – 7 × 1 – 1.5 cm], and ecology [on marble]’ but most essentially by the presence of one or two cauline leaves vs their absence in the latter species. Pachomova (in Vvedensky 1974: 113 – 114) did not take into account any of these characters but separated the two species based on the intensity and localisation of glandular cover on leaves — sparse throughout in P. korovinii vs very sparse and usually restricted to petioles or sometimes absent in P. pulvinata. Indeed, the specimens she annotated as P. korovinii (e.g. Botbayeva s.n. and Sultanova s.n. — FRU!), both of which are from its type locality, are plants with predominantly (except for one plant) leaﬂess stems 5 – 10 cm tall with linear-oblanceolate to narrowly obovate leaves to 4 × 0.7 cm. These are typical forms of P. pulvinata but with leaves sparsely glandular throughout. The presence of eglandular and densely to sparsely glandular forms is known within many species of Parrya and, therefore, this feature is not taxonomically 467 useful in this genus. A careful examination of ample material of this complex reveals that a single species is involved and that there are no sound morphological differences between the two. It is important to emphasise that the stem leaves mentioned by Vassiljeva (1969) are highly reduced and borne on the pedicels of some scapes of the holotype of P. korovinii and on one of several plants of Sultanova s.n. Therefore, P. korovinii apparently represents a local, slightly more glandular plant of P. pulvinata. Although the holotype of Parrya korovinii represents a luxurious form of P. pulvinata, that of P. minuta is a diminutive fruiting form of the same species. It is interesting to note that Vassiljeva (1974) placed (as Neuroloma) P. korovinii and P. pulvinata in different sections separated mainly by the presence of white vs lavender to purple ﬂowers, respectively. However, ﬂower colour in P. korovinii was not known to her, and such sectional assignments clearly demonstrate the artiﬁciality of Vassiljeva’s (1974) infrageneric classiﬁcation of the genus. 31. Parrya runcinata (Regel & Schmalh.) N. Busch in Kom. (Komarov 1939: 645); Parrya fruticulosa var. runcinata Regel & Schmalh. in Regel (1877: 237); Achoriphragma runcinatum (Regel & Schmalh.) Soják (1982: 106); Neuroloma runcinatum (Regel & Schmalh.) Botsch. (Botschantzev 1972: 672). Type: [Tajikistan], Seravshan basin, Pass Kadzhara, 3,300 m [10,900 ft], 20 June 1870, Fedtschenko s.n. (holotype LE!). Parrya stenocarpa var. major Kom. (Komarov 1896: 87). Type: [Tajikistan], Seravshan, Varsoüt, talus slopes of alpine belt, 2,750 m [9,000 ft], 20 July 1892, Komarov s.n. (lectotype LE!), selected here. DISTRIBUTION. Pamir-Alai (Kyrgyzstan, Tajikistan). 32. Parrya rydbergii Botsch. (Botschantzev 1955: 178); Parrya platycarpa Rydb. (Rydberg 1912: 326), non Hook. f. & Thomson (Hooker & Thomson 1861: 136); Neuroloma rydbergii (Botsch.) Botsch. (Botschantzev 1972: 672); P. nudicaulis subsp. rydbergii (Botsch.) Hultén (1971: 318); P. nudicaulis var. rydbergii (Botsch.) N. H. Holmgren (2004: 247). Type: USA, Utah, ‘near the summit of one of the highest peaks of the Uintas, above Bear River Cañon’, Uinta Mts, 3,650 m [12,000 ft], Aug. 1869, Watson 54 (holotype NY!; isotypes GH!, US!). DISTRIBUTION. USA (Utah, Wyoming). 33. Parrya sarawschanica (Regel & Schmalh.) D. A. German & Al-Shehbaz, comb. nov. Type: [Uzbekistan], Zaravschanskaya valley, Dzhizmanskoye gorge, 600 – 900 m [2,040 − 2,874 ft], 2 May 1869, Fedtschenko s.n. (holotype LE!). http://www.ipni.org/urn:lsid:ipni.org:names:77128849-1 © The Board of Trustees of the Royal Botanic Gardens, Kew, 2013 468 Diptychocarpus sarawschanicus Regel & Schmalh. in Regel, Trudy Imp. S.-Peterburgsk. Bot. Sada 5: 231 (1877), non Parrya seravschanica A. N. Vassiljeva (1969: 21), nom. invalid; Pseudoclausia sarawschanica (Regel & Schmalh.) Botsch. (Botschantzev 1965a: 125); Clausia sarawschanica (Regel & Schmalh.) Lipsky (1904: 45). DISTRIBUTION. Pamir-Alai (Uzbekistan). 34. Parrya saurica (Pachom.) D. A. German & Al-Shehbaz in German (2010: 85); Neuroloma sauricum Pachom. (Pachomova 1974: 41); Achoriphragma sauricum (Pachom.) Soják (1982: 106). Type: [Kazakhstan], Semipalatinsk province, Zaissan district, Saur, pass Taz, rocky alpine slope, 21 July 1914, Shishkin & Genina s.n. (holotype LE!; isotypes AA!, LE × 4 sheets!, TK!). DISTRIBUTION. Saur (Kazakhstan). 35. Parrya saxifraga Botsch. & Vved. (Botschantzev & Vvedensky 1941: 15); Achoriphragma saxifraga (Botsch. & Vved.) Soják (1982: 106); Neuroloma saxifraga (Botsch. & Vved.) Botsch. (Botschantzev 1972: 672). Type: [Uzbekistan], W Tian Shan, basin of river Angren, upper reaches, along road from Orta-Ailyksai towards Pass Kem-Saz, on gravelly soil, 30 July 1938, Pjatayeva & Momotov 401 (holotype TASH!). KEW BULLETIN VOL. 68(3) selected by German in German & Cherneva (2008: 305). The fruiting specimen marked ‘Herb Fischer’ is the lectotype. Parrya nudicaulis var. ciliata Regel (1870: 259). Type: [Kazakhstan], Dzungaria, Schrenk s.n. (holotype LE!). Parrya stenophylla Popov in Kom. (Komarov 1939: 646); Achoriphragma stenophyllum (Popov) Czerep. (Czerepanov 1995: 126); Neuroloma stenophyllum (Popov) Botsch. (Botschantzev 1972: 673). Type: [Kazakhstan], Zailiysky Alatau, Small Almatinka River, glacier near gates of Tyuyuksu, moraine gravel, 7 July 1936, Popov s.n. (holotype LE!; isotypes AA × 2 sheets!). Parrya stenocarpa var. glabra Krylov (1931: 1394). Type: [Kazakhstan, S Altai, Sarym-Sakty range], vicinity of Katon-Karagai, N slope of Altaisky range, alpine meadows, 10 June 1923, Sumnevich s.n. (lectotype TK!), selected by Gureyeva et al. (2012: 19). DISTRIBUTION. S Altai, Saur, Tarbagatai, Tian Shan [China (Xinjiang), Kazakhstan, Kyrgyzstan]. NOTES. Except for being eglandular (vs often glandular) and having narrower leaves (1 − 2.5 vs 3 − 5 (− 9) mm wide), Parrya stenophylla is basically indistinguishable from P. stenocarpa and therefore the two species are merged herein following Pachomova (in Vvedensky 1974). However, glandular and eglandular forms often occur within populations of many species of the genus, and leaf width is not sufﬁciently different between them. DISTRIBUTION. W Tian Shan (Uzbekistan). 36. Parrya schugnana Lipsch. (Lipschitz 1935: 31); Achoriphragma schugnanum (Lipsch.) Soják (1982: 106); Neuroloma schugnanum (Lipsch.) Botsch. (Botschantzev 1972: 672). Type: [Tajikistan], Shugnan, cliff above river Gunt, vicinity of village Demiona, 22 July 1931, Yu. Lipschitz 611 (holotype MW!; isotypes LE!, MW × 3 sheets!). DISTRIBUTION. Pamir-Alai (Tajikistan). 37. Parrya stenocarpa Kar. & Kir. (Karelin & Kirilow 1842: 147); Achoriphragma stenocarpum (Kar. & Kir.) Soják (1982: 106); Neuroloma stenocarpum (Kar. & Kir.) Botsch. (Botschantzev 1972: 673). Type: [Kazakhstan], Mts Alatau, between river Baskan and Sarchan, [July] 1841, Karelin & Kirilov 1197 (lectotype LE!; isolectotypes B!, K!, KW!, LE × 3 sheet!, M!, P × 2 sheets!, PRC!, US, W!), selected by Botschantzev (1972: 673). Parrya stenoloma Schrenk ex Fisch., C. A. Mey. & AvéLall. (Fischer et al. 1842: 69). Type: [Kazakhstan], Alps of Tarbagati, [June 1841, Schrenk s.n.] (lectotype LE!; isolectotypes AA!, LE × 7 sheets!, WU!), © The Board of Trustees of the Royal Botanic Gardens, Kew, 2013 38. Parrya subsiliquosa Popov in Kom. (Komarov 1939: 645); Achoriphragma subsiliquosum (Popov) Soják, (1982: 106); Neuroloma subsiliquosum (Popov) Botsch. (Botschantzev 1972: 673). Type: Kazakhstan, Zailiysky Alatau, river Talgar, rocky cliffs close to meteorological station, 25 May 1936, Popov s.n. (lectotype LE!; isolectotype AA!), selected by Botschantzev (1972: 673, as type) and German & Veselova (2011: 1385). Hesperis kunawarensis var. hirtula Regel (1870: 273). Type: [Kazakhstan, Zailiysky Alatau], Mts near Wernoje, 29 April [1867], [?] Sewerzow s.n. (lectotype LE!; isolectotype LE!), selected here. DISTRIBUTION. N Tian Shan (Kazakhstan, Kyrgysztan). 39. Parrya tianschanica (Botsch.) D. A. German (2012b: 40); Neuroloma tianschanicum Botsch. (Botschantzev 1972: 673); Parrya tianschanica Nikitina (1967: 112), nom. inval. Achoriphragma tianschanicum (Nikitina) Soják, (1982: 106), comb. inval. Type: Kyrgyzstan, [Central] Tian Shan, Toguz-Torovsky region, Kyldoo, subalpine zone, NE slope, in rock crevices, 24 July 1957, Aidarova & Ubukeyeva s.n. (holotype FRU!; isotype FRU!). A SYNOPSIS OF THE GENUS PARRYA (BRASSICACEAE) DISTRIBUTION. C Tian Shan, NE Pamir-Alai (Kyrgyzstan). NOTES. By citing two collections as ‘type’, Nikitina (1967) invalidly published the name Parrya tianschanica under Art. 40.1 (McNeill et al. 2012). A combination with the same epithet was validated later by Botschantzev (1972: 673) by citing the single collection as ‘lectotype’ (i.e., holotype; Art. 40.3) in the genus Neuroloma, which was used as the basionym for validating the name P. tianschanica (German 2012b). 40. Parrya tschimganica (Popov ex Botsch. & Vved.) D. A. German & Al-Shehbaz, comb. nov. Type: [Uzbekistan], Tschimgan, by river from Tschimgan Mt, 6 July 1926, Popov 12 (holotype TASH 75072!). http://www.ipni.org/urn:lsid:ipni.org:names:77128850-1 Clausia tschimganica Popov ex Botsch. & Vved., Bot. Mater. Gerb. Bot. Inst. Uzbekistansk. Fil. Akad. Nauk S.S.S.R. 3: 13 (Botschantzev & Vvedensky 1941); Pseudoclausia tschimganica (Popov ex Botsch. & Vved.) A. N. Vassiljeva (1961: 246). DISTRIBUTION. W Tian Shan (Kazakhstan, Kyrgyzstan, Uzbekistan). 41. Parrya turkestanica (Korsh.) N. Busch in Kom. (Komarov 1939: 644); Parrya macrocarpa var. turkestanica Korsh. (Korshinsky 1898: 407); Achoriphragma turkestanicum (Korsh.) Soják (1982: 106); Neuroloma turkestanicum (Korsh.) Botsch. (Botschantzev 1972: 673); Parrya nudicaulis subsp. turkestanica (Korsh.) Hultén (1945: 892). Type: [Tajikistan], Alaisky Range, over Pass Tengiz-bai, slopes and cliffs, 3,050 – 3,350 m [10,000 − 11,000 ft], 19 July 1895, Korshinsky s.n. (lectotype LE!; isolectotype LE!), selected by Botschantzev (1972: 673). 469 Taxa excluded from Parrya Names in boldface are the currently accepted placements of excluded taxa, and their tribal assignments are given in square brackets [ ] at the end of each entry. Parrya arenicola (Richardson ex Hook.) Hook. f. (Hooker 1861: 285, 315). Type: deep sand upon shores of Arctic America between longitude 107° and 150°, Richardson s.n. (holotype (n.v.); isotype CAN!). = Arabidopsis arenicola (Richardson ex Hook.) Al-Shehbaz, R. Elven, D. F. Murray & Warwick [Camelineae]. Parrya beketovii Krasn., Enum. Pl. Tian-Schan Or. 18. (Krasnov 1887). Type: [SE Kazakhstan, Tian Shan], in valley of Tscharyn, [30 – 31 May] 1886, Krassnow s.n. (lectotype LE!; isolectotypes LE × 5 sheets!), selected by Botschantzev (1972: 670) and here = Leiospora beketovii (Krasn.) D. A. German & Al-Shehbaz, comb. nov. [Euclidieae]. http://www.ipni.org/urn:lsid:ipni.org:names:77128853-1 As shown by German et al. (2011), the species is nested within Leiosopora of the tribe Euclidieae and therefore is transferred herein. The sheet with three ﬂowering specimens carrying the handwriting of Krasnov is designated as the second-step lectotype. Parrya Danguy (1908: 51). Type: Bourr-Teppe, 3380 m, 4 July 1906, Lacoste 1 (holotype P) = Leiospora bellidifolia (Danguy) Botsch. & Pachom. [Euclidieae]. Parrya cheiranthoides (Nutt.) Jepson (1936: 75). Type: USA, Washington, Walla Walla, Nuttall s.n. (holotype BM!; isotypes K!, NY!, PH!) = Phoenicaulis cheiranthoides Nutt. [Boechereae]. DISTRIBUTION. Pamir-Alai (Afghanistan, Kyrgyzstan, Tajikistan, Uzbekistan). 42. Parrya vvedenskyi (Pachom.) D. A. German & AlShehbaz, comb. nov. Type: Kazakhstan, Syr Darya Province, Aulie-Atinsky distr., S Karatau, basin of river Asy, gravelly E slope of Berkary sai on the Karatau plateau. 20 June 1925, Sovietkina 495 (holotype TASH 75051!). http://www.ipni.org/urn:lsid:ipni.org:names:77128851-1 Pseudoclausia vvedenskyi Pachom., Bot. Mater. Gerb. Inst. Bot. Akad. Nauk Uzbeksk. S.S.R. 19: 39 (Pachomova 1974). DISTRIBUTION. W Tian Shan (Kazakhstan). Parrya cheiranthoides var. glabra (Jepson) Jepson (1936: 76). Type: USA, California, June 1892, Bruce 2250 (holotype UC!) = Phoenicaulis cheiranthoides Nutt. [Boechereae]. Parrya cheiranthoides var. lanuginosa (S. Watson ex B. L. Rob.) M. Peck (1941: 352). Type: Canada, N.W.T., bluffs E side of the Columbia below the Chelan, 14 Oct. 1880, Watson 28 (holotype GH!) = Phoenicaulis cheiranthoides Nutt. [Boechereae]. Parrya ciliaris Bureau & Franch. (Bureau & Franchet 1891: 20). Type: China, Tibet [Xizang]: route de Batang, 12 May 1890, Bonvalot & d'Orléans s.n. (holotype P!). = Solmslaubachia pulcherrima Muschl. [Euclidieae]. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2013 470 Parrya codringtonii (Rech. f.) Rech. f. (Rechinger 1964: 424). Type: Afghanistan, Baiman, Band-i-Amir, 3,650 m [12,000 ft], 4 April 1940, Codrington s.n. (holotype BM!). = Matthiola codringtonii Rech. f. [Anchonieae]. Parrya crassifolia Botsch. & Vved. (Botschantzev & Vvedensky 1948: 9). Type: Ferganisky Range, pass Kugart, W slope, 11 Aug. 1929, Titov & Ioffe 250 (holotype TASH-180230!; fragments LE!) = Leiospora crassifolia (Botsch. & Vved.) A. N. Vassiljeva. [Euclidieae]. Parrya eriocalyx Regel & Schmalh. in Regel (1877: 234). Type: [Kyrgyzstan, C Tian Shan], on the pass Terekty and in Aksai valley, July 1872, Kaulbars s.n. (holotype LE!) = Leiospora eriocalyx (Regel & Schmalh.) Dvořák. [Euclidieae]. Parrya ermanii Ledeb. (Ledebour 1841: 132), nom. illegit. Type: Same as that of Draba parryoides Cham. (see P. parryoides). [Smelowskieae]. Parrya eurycarpa (A. Gray) Jepson (1925: 434), non P. eurycarpa Maxim. (Maximowizc 1889: 56). Type: USA, California. Peak near Sonora Pass, 3,500 m [11,500 ft], 1863, Brewer 1909 (holotype GH!; isotypes B!, US!). = Anelsonia eurycarpa (A. Gray) J. F. Macbr. & Payson [Boechereae]. Parrya eurycarpa Maxim. (Maximowizc 1889: 56), non P. eurycarpa (A. Gray) Jepson (1925: 434). Type: [China], Tibet, near Jagem-Gol, 20 July 1884, Przewalski s.n. (lectotype LE!; isolectotype PE!), selected by Buzunova in Grubov (2000: 72, as holotype). = Solms-laubachia eurycarpa (Maxim.) Botsch. [Euclidieae]. Parrya exscapa C. A. Mey. in Ledeb. (Ledebour 1829: 21). Type: [Russia, Altai]. In lapidosis summarum alpinum Kuraicae et Tscheganense ad Tschujam legit Dr. Bunge. Floret Majo, Junio [1826] (lectotype LE!), selected by Botschantzev (1972: 669, as type). = Leiospora exscapa (C. A. Mey.) Dvořák [Euclidieae]. Parrya exscapa var. tianschanica Popov & P. A. Baranov (1923: 175). Type: [Uzbekistan], Syr-Daryinskaya province, Tashkent distr., Talassky Alatau, valley of Maidantal, right bank below the red rocks on the opposite of mouth of Korum-bel, 16 – 17 Aug. 1922, Baranov 1350 (lectotype TASH 9791!; same as lectotype of Parrya subscapigera), selected here. = Leiospora subscapigera (Botsch. & Vved.) Botsch. & Pachom. [Euclidieae]. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2013 KEW BULLETIN VOL. 68(3) Parrya ﬁnchiana Dunn (1927: 247). Type: [China, Xizang] Tibet, along Chongphu torrent, c. 20 miles NE of Mt Everest, 5,200 m [17,000 ft], 8 June 1922, Norton 41 (holotype K!). = Solms-laubachia platycarpa (Hook. f. & Thomson) Botsch. [Euclidieae]. Parrya ﬂabellata Regel (1870: 261). Type: [Kyrgyzstan], southern Tian Shan, Dschaman-Daban, 21 July [1867, Fr. Osten-Sacken s.n.], Sewerzow s.n. (holotype LE!; isotypes LE × 2 sheets!). = Solms-laubachia ﬂabellata (Regel) J. P. Yue, Al-Shehbaz & H. Sun [Euclidieae]. As revealed by Botschantzev (1955: 165), the holotype of Parrya ﬂabellata was collected by H. F. von der Oster-Sacken, not Sewerzow as mentioned in the original publication, and the two isotypes at LE have the correct collector name. Parrya forrestii W. W. Sm. (Smith 1914: 195). Type: China, Yunnan, western ﬂank of the Lichiang Range, 27°25'N, Aug. 1910, Forrest 6518 (lectotype E!; isolectotype K!), selected by Polatschek (1994: 200, as holotype). = Erysimum forrestii (W. W. Sm.) Polatschek [Erysimeae]. Parrya grandiﬂora (C. A. Mey.) Schischk. in Krylov (1931: 1394). Type: [Russia, Siberian Altai], 990 [Prodr. Fl. Alt.], In summa alpe ad desertum editum amnis Tschuja legit Junio 1826 cel. Dr. Bunge [s.n.] (Hb. Meyer) (lectotype LE!; isolectotypes B!, LE × 5 sheets!, M!, P?, W!), selected by German (2005: 242). = Pachyneurum grandiﬂorum (C. A. Mey.) Bunge [Arabideae]. Previous designation of lectotype by Botschantzev (1972: 668, as type) cannot be followed because it is based on a later (1832) collection by A. A. Bunge. Parrya huddelliana A. Nelson (1912: 139). Type: USA, Idaho: Custer Co., Macay, Bear Canyon, 3,050 m [10,000 ft], 31 July 1911, Nelson & Macbride 1466 (holotype RM!; isotypes DS!, GH!, MO!, NY!, POM!, UC!, US!) = Anelsonia eurycarpa (A. Gray) J. F. Macbr. & Payson [Boechereae]. Parrya karatavica Lipsch. & Pavlov in Lipsch. (Lipschitz 1936: 318). Type: [Kazakhstan], Kara-Tau range, Boroldaiskiye Mts, Terseilau near Mikhailovka, stony slopes, 27 May 1934, Chilikina 327 (lectotype LE!), selected by Nabiev (1972: 187). = Botschantzevia karatavica (Lipsch. & Pavlov) Nabiev [Arabideae]. Although all syntypes were initially deposited at MW, the choice of lectotype by Bagdasarova & Gubanov (1975: 343, as holotype), cannot be followed because one syntype transferred to LE was designated by Nabiev (1972) as the lectotype. A SYNOPSIS OF THE GENUS PARRYA (BRASSICACEAE) Parrya lanuginosa Hook. f. & Thomson (Hooker & Thomson 1861: 136). Type: [India], Lanjar, 5,350 m [17,500 ft], Strachey & Winterbottom 7 (lectotype K!; isolectotypes BM, GH!, LE!), selected by Al-Shehbaz & Yang (2000: 347, as holotype). = Solms-laubachia lanuginosa (Hook. f. & Thomson) D. A. German & AlShehbaz [Euclidieae]. Parrya linearifolia W. W. Sm. (Smith 1920: 219), non Pavlov (1949: 29). Type: China. Yunnan: Beima Shan, MekongYangtze divide, 28°20'N, 4,400 m [14,500 ft], Aug. 1914, Forrest 13,235 (lectotype E!; isolectotype E!), selected by Yue et al. (2008: 535). = Solms-laubachia linearifolia (W. W. Sm.) O. E. Schulz [Euclidieae]. Parrya menziesii (Hook.) Greene (1891: 253). Type: USA, California, Menzies s.n. (holotype K!). = Erysimum menziesii (Hook.) Wettst. [Erysimeae]. Parrya menziesii var. glabra Jepson (1925: 434). Type: USA, California, June 1892, Bruce 2250 (holotype, UC!) = Phoenicaulis cheiranthoides Nutt. [Boechereae]. Parrya menziesii var. lanuginosa S. Watson (1895: 152). Type: Canada, N.W.T., bluffs E side of the Columbia below the Chelan, 14 Oct. 1880, Watson 28 (holotype GH!) = Phoenicaulis cheiranthoides Nutt. [Boechereae]. Parrya microcarpa Ledeb. (Ledebour 1841: 132), nom. illegit. Type: same as that of Parrya grandiﬂora = Pachyneurum grandiﬂorum (C. A. Mey.) Bunge [Arabideae]. Parrya pamirica Botsch. & Vved. in Botsch. (Botschantzev 1965b: 279). Type: [Tajikistan], Zaalaisky range, 250th km of Pamir highway, granite gravel, 29 June 1948, Lavrenko & Rodin 586 (holotype LE!; isotype LE!) = Leiospora pamirica (Botsch. & Vved.) Botsch. & Pachom. [Euclidieae]. 471 Parrya pedicellata (A. Nelson) Tidestr. (Tidestrom 1925: 247, 248). Type: USA Nevada, Hunter Creek Canyon, 5 miles W of Reno, 16 May 1903, Kennedy & True 705 (holotype RM!; isotypes DS!, UC!) = Phoenicaulis cheiranthoides Nutt. [Boechereae]. Parrya platycarpa Hook. f. & Thomson (Hooker & Thomson 1861: 136), non Rydb. (Rydberg 1912: 326). Type: [China, Xizang]: Tibet, along Chongphu torrent, c. 20 miles NE of Mt Everest, 5,200 m [17,000 ft], 8 June 1922, Norton 41 (holotype K!). = Solmslaubachia platycarpa (Hook. f. & Thomson) Botsch. [Euclidieae]. Parrya prolifera Maxim. (Maximowizc 1889: 56). Type: China, Tibet, Kon-chun-ua, 4,400 m [14,500 ft], 3 July 1884, Prezwalski s.n. (lectotype LE!; isolectotypes K!, P!, PE!), selected by Buzunova in Grubov (2000: 72, as holotype). = Solms-laubachia prolifera (Maxim.) J. P. Yue, Al-Shehbaz & H. Sun [Euclidieae]. Parrya pumila Kurz (1872: 285). Type: [Kashmir], Rupschu, 4,500 − 5,500 m [15,000 − 18,000 ft], not dated, Stoliczka s.n. (holotype CAL; isotype K!) = Solmslaubachia pumila (Kurz) Dvořák [Euclidieae]. Parrya ramosissima Franch. (Franchet 1896: 343). Type: [Tajikistan, Pamir], ‘Kara Djilga river, 4,300 m, 27 June 1893, M. E. de Poncins’ (holotype P) = Christolea crassifolia Cambess. [Euclidieae]. Parrya saposhnikovii A. N. Vassiljeva, Bot. Mater. Gerb. Inst. Bot. Akad. Nauk Kazakhst. S.S.R. 6: 19 (1969). Type: [Kyrgyzstan, Central Tian Shan], Semirechenskaya province, Przewalsky distr., river Kaindy on the opposite of mouth of river Karabel, rocky gorge, 5 Aug. 1912, Saposhnikov & Shishkin s.n. (holotype TK!; isotype TK!). = Leiospora saposhnikovii (A. N. Vassiljeva) D. A. German & Al-Shehbaz, comb. nov. [Euclidieae]. http://www.ipni.org/urn:lsid:ipni.org:names:77128855-1 Parrya parryoides (Cham.) Hultén (1928: 173). Type: Russia, Kamchatka, ‘in rupestribus montis ignivomi Schiwelutsch, 4,000 m’, Erman s.n. (lectotype LE; isolectotype B!), selected by Botschantzev (1972: 668). = Smelowskia parryoides (Cham.) Polunin [Smelowskieae]. Although Botschantzev (1955: 163; 1972: 668) mentioned the type seen by him at LE, thorough searches by both of us failed to ﬁnd it. Ledebour (1841: 132) reported Erman’s specimen at B, and we take that as the isolectotype. Molecular data (German et al. 2011) clearly demonstrated that this species is nested within Leiospora which is supported by morphology. Parrya subscapigera Botsch. & Vved. (Botschantzev & Vvedensky 1948: 9). Type: [Uzbekistan], SyrDaryinskaya province, Taskkent distr., Talassky Alatau, valley of Maidantal, right bank below the red rocks on the opposite of mouth of Korum-bel, 16 – 17 Aug. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2013 472 1922, Baranov 1350 (lectotype TASH 9791!), selected by Botschantzev (1972: 669). = Leiospora subscapigera (Botsch. & Vved.) Botsch. & Pachom. [Euclidieae]. Parrya surculosa N. Busch in Kom. (Komarov 1939: 647). Type: Tajikistan, Schugnan, Kaldok ad jugum Abkhary, 2 Aug. 1904, Fedtschenko s.n. (lectotype LE!; isolectotypes LE × 5 sheets!), selected here = Solms-laubachia surculosa (N. Busch) D. A. German & Al-Shehbaz. [Euclidieae]. Busch (loc. cit.) erroneously listed the year of collection as 1894. Of the six sheets of the type collection at LE, the single one annotated in Busch’s handwriting is taken here as the lectotype. Parrya villosa Maxim. (Maximowizc 1889: 55). Type: [China], Tibet borealis, ad. ﬂ. Kon-tschun-tschu, regio alpina frequens, 1 – 13 June 1884, Preszwalski s.n. (lectotype LE!), selected by Grubov (2000: 72) = Solms-laubachia villosa (Maxim.) D. A. German & AlShehbaz [Euclidieae]. Parrya xerophyta W. W. Sm. (Smith 1920: 217). Type: China, Yunnan, NE of Chungtien, 27°55'N, July 1918, Forrest 16444 (holotype E!; isotypes E!, K!, P!, W!) = Solmslaubachia xerophyta (W. W. Sm.) H. F. Comber [Euclidieae]. Both Parrya michaelis A. N. Vassiljeva and P. seravschanica A. N. Vassiljeva (1969: 17, 21, respectively) were invalidly published because under each, more than one specimen was listed as the type (McNeill et al. 2012: Art. 40.1). They were accepted by Czerepanov (1973) who cited their types as originally published and thus did not validate them, nor were they subsequently validated (German 2012b). We agree with Botschantzev (1972) and Pachomova (in Vvedensky 1974) in reducing them to synonymy with P. beketovii (= Leiospora beketovii) and L. subscapigera, respectively [Euclidieae]. Acknowledgements We are most grateful to the directors and curators of AA, ALA, B, BM, BP, BRNU, CAN, CAS, DS, E, FRU, GH, K, KFTA, KW, LE, M, MHA, MO, MSB, MW, NY, O, P, PE, PH, POM, PRC, RM, TAD, TASH, TK, UC, US, W, and WU for loans and access to their collections. We thank Olga V. Cherneva, Vladimir I. Dorofeyev and Dmitry V. Geltman (all at LE), Alexander L. Ebel (TK), Alexander P. Shalimov (ALTB) and Tatyana V. Shulkina (MO) for their help, as well as Alexander N. Sennikov (H) for taxonomic advice. 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