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
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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. Fieldwork and herbarium studies in central
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013
KEW BULLETIN VOL. 68(3)
Asia and western China were generously supported by
the National Geographic Society grants 7121–01,
6286–98 to Al-Shehbaz and 8773 – 10 to German; by
the German Research Foundation (project BrassiBase
within priority program 1529 ‘Adaptomics’), and
Russian Foundation for Basic Research (grant N 1104-90702) to German.
References
Adams, M. F. (1817). Descriptiones plantarum minus
cognitarum Sibiriae praesertim orientalis, quas in
itinere ann. 1805 et 1806 observavit. Mém. Soc. Nat.
Moscou 5: 89 – 116.
Al-Shehbaz, I. A. (2010). Parrya. In: Flora of North
America Editorial Committee (eds), Flora of North
America 7: 511 − 514. Oxford, New York.
____ (2012). A generic and tribal synopsis of the
Brassicaceae (Cruciferae). Taxon 61: 931 – 954.
____ & Yang, G. (2000). A reconsideration of the genus
Eurycarpus (Brassicaceae). Novon 10: 346 – 348.
____, Beilstein, M. A. & Kellogg, E. A. (2006).
Systematics and phylogeny of the Brassicaceae: an
overview. Pl. Syst. Evol. 259: 89 − 120.
____, Grant, J. R., Lipkin, R., Murray, D. F. & Parker,
C. L. (2007). Parrya nauruaq (Brassicaceae), a new
species from Alaska. Novon 17: 275 – 278.
Appel, O. & Al-Shehbaz, I. A. (2003). Cruciferae. In:
K. Kubitzki (ed.), Families and Genera of Vascular
Plants 5: 75 − 174. Springer-Verlag, Berlin,
Heidelberg.
Bagdasarova, T. V. & Gubanov, I. A. (1975).
Specimina authentica taxorum a cl. Nicolao
Pavlovio descriptorum in herbario Universitatis
Mosquensis conservata. Novosti Sist. Vyssh. Rast.
12: 335 – 355.
Baranov, P. A. (1925). Sketch of the vegetation of
Chimgan. J. Turkest. Branch Russ. Geogr. Soc. 17: 1 – 34.
Boissier, P. E. (1856). Diagnoses Plantarurm Orientalium
Novarum. Ser. 2, vol. 5. Herrmann, Leipzig.
____ (1867). Flora orientalis. Vol. 1. H. Georg, Basel,
Genève.
Boivin, B. (1966). Énumération des plantes du Canada. Naturaliste Canad. 93: 583 – 646.
Botschantzev, V. P. (1955). De Cruciferis notae
criticae. Bot. Mater. Gerb. Bot. Inst. Komarova Akad.
Nauk. S.S.S.R. 17: 160 − 178.
____ (1965a). De cruciferis notae criticae, 4. Novosti
Sist. Vyssh. Rast. 1965 [2]: 123 – 125.
____ (1965b). Species novae ex Asia Media. Novosti
Sist. Vyssh. Rast. 1965 [2]: 268 – 281.
____ (1972). On Parrya R. Br., Neuroloma Andrz. and
some other genera (Cruciferae). Bot. Zhurn. (Moscow & Leningrad) 57: 664 − 673.
____ & Vvedensky, A. I. (1941). Species plantarum
novae. Bot. Mater. Gerb. Bot. Inst. Uzbekistansk. Fil.
Akad. Nauk S.S.S.R. 3: 3 – 20.
A SYNOPSIS OF THE GENUS PARRYA (BRASSICACEAE)
____ & ____ (1948). Cruciferae novae ex Asia Media.
Bot. Mater. Gerb. Inst. Bot. Zool. Acad. Nauk
Uzbekistansk. S.S.R. 12: 3 – 12.
Brown, R. (1823). Chloris melvilliana. W. Clowes,
London.
Bureau, E. & Franchet, A. (1891). Plantes nouvelles du
Thibet et de la Chine occidentale. J. Bot. (Morot) 5:
17 – 25.
Cafferty, S. & Jarvis, C. E. (2002). Typification of
Linnaean plant names in Brassicaceae (Cruciferae).
Taxon 51: 529 – 537.
de Candolle, A. P. (1821). Regni vegetabilis systema
naturale. Vol. 2. Treuttel & Würtz, Paris.
____ (1824). Prodromus systematis naturalis regni
vegetabilis. Vol. 1. Treuttel & Würtz, Paris.
Cardot, J. (1911). Deux genres noveaux de la région
magellanique. Rev. Bryol. 38: 49 – 52.
Czerepanov, S. K. (1973). Svod Dopolneni i Izmeneni k
‘Flore SSSR’ (tt. I – XXX). Science Press, Leningrad.
____ (1995). Vascular Plants of Russia & Adjacent States
(The Former USSR). University Press, Cambridge.
Danguy, P. (1908). Note sur une collection botanique
rapportée du Pamir par le commandant de
Lacoste. J. Bot. (Morot) 21: 49 – 53.
Don, G. (1831). A General History of the Dichlamydeous
Plants. Vol. 1. J. G. & F. Rivington, London.
Dunn, S. T. (1927). Parrya finchiana. Bull. Misc. Inform.,
Kew 1927: 247.
Ebel, A. L. (1999). Type specimens of the Brassicaceae taxa
kept in Krylov Herbarium of the Tomsk State University
(TK). Bot. Zhurn. (St. Petersburg) 84(6): 71 − 77.
Fedtschenko, B. A. (1904). Flora of West Tian Shan.
Botanical results of expeditions of 1897 and 1902
and an overview of previous studies. Trudy Imp. S.Petersburgsk. Bot. Sada 23: 249 – 532.
Fischer, F. E. L., Meyer, C. A. & Avé-Lallemant, J. L. E.
(1842). Index octavus seminum, quae Hortus Botanicus
Imperialis Petropolitanus pro mutua commutatione offert,
accedunt animadversione botanicae nonnulae.
Petropoli.
Franchet, A. (1896). Note sur une collection de
plantes rapportées du Pamir en 1894 par M. E. de
Poncins. Bull. Mus. Hist. Nat. (Paris) 2: 342 – 346.
German, D. A. (2005). Type specimens of Altai
Cruciferae taxa described in 1829 – 1841 by C. A.
Meyer, A. A. Bunge and C. F. Ledebour deposited
in the Herbarium of the Komarov Botanical
Institute (LE). Novosti Sist. Vyssh. Rast. 37: 233 – 267.
____ (2010). A check-list and the system of the
Cruciferae of Altai. Komarovia 6(2): 80 – 88.
____ (2012a). Typification of names of Cruciferae taxa
described by E. R. Trautvetter from Middle Asia
and Kazakhstan. Novosti Sist. Vyssh. Rast. 43: 111 –
131.
____ (2012b). Taxonomical confusions in the Cruciferae
of North and Central Asia. II. Sisymbrium lineare and
Arabis crassifolia. Turczaninowia 15(2): 37 – 43.
473
____ & Cherneva, O. V. (2008). Typification of
Cruciferae taxa described by A. G. Schrenk. Novosti
Sist. Vyssh. Rast. 40: 285 – 312.
____ & Veselova, P. V. (2011). Type specimens of
Brassicaceae (Cruciferae) taxa deposited in the
herbarium of Institute of Botany and
Phytointroduction of Kazakhstan (AA). Bot. Zhurn.
(St. Petersburg) 96: 1378 – 1388.
____, Grant, J. R., Lysak, M. A. & Al-Shehbaz, I. A.
(2011). Molecular phylogeny and systematics of the
tribe Chorisporeae (Brassicaceae). Pl. Syst. Evol.
294: 65 − 86.
Goloskokov, V. P. (1963). Herbarium typorum
plantarum kazachstanicae. Bot. Mater. Gerb. Inst.
Bot. Acad. Nauk Kazakhst. S.S.R. 1: 4 – 67.
Greene, E. L. (1891). Flora Franciscana. Cubery & Co.,
San Francisco.
Grubov, V. I. (ed). (2000). Catalogue of the type specimens
of Central Asian vascular plants in the Herbarium of the
V. L. Komarov Botanical Institute. St. Petersburg
University Press, St. Petersburg.
Gubanov, I. A., Bagdasarova, T. V. & Balandina, T. P.
(1998). Scientific Heritage of Outstanding Russian
Florists G. S. Karelin and I. P. Kirilov. Moscow.
Gureyeva, I. I., Balashova, V. F., German, D. A. & Ebel,
A. L. (2012). Type specimens of Brassicaceae
Burnett in the Krylov Herbarium (TK). Sist. Zametki
Mater. Gerb. Krylova Tomsk. Gosud. Univ. 106: 3 – 23.
Holmgren, N. H. (2004). Lectotypification and new
combinations in North American Brassicaceae.
Brittonia 56: 245 – 248.
Hooker, J. D. (1861). Outlines of the distribution of
Arctic plants. Trans. Linn. Soc. London 23 (2): 251 –
348.
____ & Thomson, T. (1861). Praecurores ad Floram
Indicam. J. Proc. Linn. Soc., Bot. 5: 128 − 181.
Hooker, W. J. (1829). Flora Boreali-Americana. Vol. 1.
Treuttel & Würtz, London.
Hultén, E. (1928). Flora of Kamtchatka and the
adjacent islands. Kungl. Svenska Vetensk.-Akad.
Handl. Ser. III, 5(2): 1 – 1134.
____ (1945). Flora of Alaska and Yukon. Acta Univ.
Lund., N. F., Avd. 2, 40(1): 799 – 978.
____ (1968). Comments on the flora of Alaska and
Yukon. Ark. Bot. 7, ser. 2: 1 – 147.
____ (1971). The Circumpolar plants. II. Dicotyledons.
Kungl. Svenska Vetensk.-Akad. Handl. 13(1): 1 – 463.
Jafri, S. M. H. (1956). Some Cruciferae of W. Pakistan,
Afghanistan and N.W. Himalaya. Notes Roy. Bot.
Gard. Edinburgh 22: 95 − 119.
____ (1973). Brassicaceae. In: E. Nasir & S. I. Ali (eds),
Flora of West Pakistan. Vol. 5. Ferozson, Karachi.
Jepson, W. L. (1925). A Manual of the Flowering
Plants of California. Associated Students Store,
Berkeley.
____ (1936). A Flora of California. Vol. 2. The University
Press, Berkeley.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013
474
Kamelin, R. V. (1998). Materials on the history of the
flora of Asia (Altai Mountain Country). Altai State
University Press, Barnaul.
Karelin, G. & Kirilow, J. (1842). Enumeratio
plantarum in desertis Songoriae orientalis et in
jugo summarum alpium Alatau anno 1841
collectarum. Bull. Soc. Imp. Naturalistes Moscou 15:
129 – 180; 321 – 453; 503 – 542.
Kitamura, S. (1964). Plants of West Pakistan and
Afghanistan. Kyoto University, Kyoto.
Komarov, V. L. (1896). Materials to the flora of
Turkestan upland. Basin of Zeravshan. Trav. Soc.
Nat. Pétersb. Bot. 26: 31 – 162.
____ (ed.) (1939). Flora URSS. Vol. 8. Publishers of
Academy of Sciences of USSR, Moscow & Leningrad.
Korshinsky S. (1898). Fragmenta florae Turkestanicae.
Plantae novae vel minus cognitae Turkestaniae
describuntur. Bull. Acad. Imp. Sci. Saint-Pétersbourg
ser. 5, 9: 399 – 424.
Krassnoff, A. [Krasnov, A. N.] (1887). Enumeratio
Plantarum Anno 1886 in Tian Schan Orientali
Lectarum. Petropoli.
____ (1888). Descriptiones plantarum novarum vel
minus cognitarum anno 1886 a A. Krassnow in
regionibus Thian Schanicis lectarum. Script. Bot.
Hort. Univ. Imp. Petrop. 2, 1: 9 – 22.
Krylov, P. N. (1931). Flora Sibiriae Occidentalis. Vol. 6.
Tomsk.
Kuntze, O. (1891). Revisio generum plantarum
vascularium omnium atque cellularum multarum
secundum leges nomenclaturae internationales cum
enumeratione plantarum exoticarum in itinere mundi
collectarum. Part 2. Leipzig.
Kurz, S. (1872). Drei neu Tibetische Pflanzen. Flora 55:
284 – 286.
Ledebour, C. F. (1829). Icones plantarum novarum vel
imperfecte cognitarum floram Rossicam, imprimis
Altaicam, illustrantes. Cent. 1. Pl. 1 – 100. Rigae
apud I. Deubner, Londoni Parisiis et Argentorati
apud Treuttel et Würtz, Bruxellae in Libraria
Parisiensi.
____ (1841). Flora rossica. Vol. 1, part 1. Sumtibus
librariae E. Schweizerbart, Stuttgartiae.
Linnaeus, C. (1753). Species Plantarum. Vol. 2. Impensis
Laurentii Salvii, Stockhom.
Lipschitz, S. J. (1935). Parrya schugnana Lipsch., sp.
nova. Repert. Sp. Nov. Regni Veg. 38: 31.
____ (1936). Generis Parryae species novae e
Turkestania rossica. Repert. Spec. Nov. Regni Veg. 39:
318 – 319.
Lipsky, V. I. (1904). Contributio ad floram Asiae
Mediae. II. Trudy Imp. S.-Peterburgsk. Bot. Sada 23: 1
– 247.
Maximowizc, C. J. (1889). Flora tangutica, sive
enumeratio plantarum regionis Tangut (Amdo)
provinciae Kansu, nec non Tibetiae praesertim
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013
KEW BULLETIN VOL. 68(3)
orientaliborealis adque Tsaidon: ex collectionibus N.M.
Przewalski atque G.N. Potanin. (Fasc. I. Thalamiflorae
et Disciflorae, 1889). Typis Academiae Imperialis
Scientarum Petropolitanae, Petropoli.
McNeill, J., Barrie, F. R., Buck, W. R., Demoulin, V.,
Greuter, W., Hawksworth, D. L., Herendeen, P. S.,
Knapp, S., Marhold, K., Prado, J., Prud’homme van
Reine, W. F., Smith, G. F., Wiersema, J. H. &
Turland, N. J. (2012). International Code of
Nomenclature for Alge, Fungi, and Plants (Melbourne Code). Regnum Veg. 154.
Nabiev, M. M. (1972). Botschantzevia Nabiev genus
novum Cruciferarum. Novosti Sist. Vyssh. Rast. 9: 186
– 187.
Nelson, A. (1912). Contributions from the Rocky
Mountain herbarium. XI. New plants from Idaho.
Bot. Gaz. 54: 136 – 151.
Nikitina, E. V. (1967). Cruciferae. In: E. V. Nikitina, V.
M. Dzhanayeva, R. A. Aidarova, A. U. Ubukeyeva, Z.
S. Arbayeva & V. I. Tkachenko. Flora Kyrgyzsko SSR,
Suppl. 1: 72 − 78, 112 − 113. Ilim, Frunze.
Ovczinnikov, P. N. (ed) (1978). Flora Tadzhiksko SSR.
Vol. 5. Academy of Sciences USSR, Leningrad.
Pachomova, M. G. (1974). Species novae familiae
Cruciferae ex Asia Media. Bot. Mater. Gerb. Inst.
Bot. Acad. Nauk Uzbeksk. SSR 19: 34 − 49.
Pavlov, N. V. (1949). Plantae novae florae
kazachstanicae. Vestn. Akad. Nauk. Kazakhsk. S.S.R.
1949(1): 28.
Peck, M. E. (1941). A Manual of the Higher Plants of
Oregon. Binfords & Mort, Portland.
Polatschek, A. (1994). Nomenklatorischer Beitrag zur
Gattung Erysimum (Brassicaceae). Phyton (Horn)
34(2): 189 – 202.
Popov, M. G. (1924). Cruciferae. In: M. G. Popov
(ed), Key for Plants of Vicinities of Tashkent 2: 137 –
160.
____ (1938). Species novae plantarum ex Asia Media.
Bull. Soc. Imp. Naturalistes Moscou. Ser. Biol. 47: 84 –
88.
____ (1955). Clausia aprica (Steph.) K.-Trotz. Spisok
Rast. Gerb. Fl. S.S.S.R. Bot. Inst. Vsesojuzn. Akad.
Nauk. 13: 18.
____ & Baranov, P. A. (1923). Descriptiones plantarum
novarum in regione alpina montium Talas Alatau
collectarum. Bot. Mater. Gerb. Glav. Bot. Sada
R.S.F.S.R. 4: 175 – 176.
Pursh, F. T. (1814). Flora Americae Septentrionalis. Vol. 2:
439. White, Cochrane & Co., London.
Rafinesque, C. S. (1819). Prodrome Des nouveaux
Genre de Plantes observés en 1817 et 1818 dans
l’intéreur des États-Unis d’Amérique. J. Phys. Chim.
Hist. Nat. Arts 89: 96 – 107.
Rechinger, K. H. (1951). Cruciferae iranicae novae vel
minus cognitae. Phyton (Horn) 3: 44 – 68.
____ (1959). Cruciferae. Dansk. Vid. Selsk., Biol. Skrift.
10, 3 (Symb. Afghan. IV): 13 – 52.
A SYNOPSIS OF THE GENUS PARRYA (BRASSICACEAE)
____ (1964). Notizen zur Orient-Flora, 63 – 69: Neue
Cruciferen aus Afghanistan. Anz. Oesterr. Akad.
Wiss., Mat.-Naturwiss. 1964: 422 – 427.
____ (1968). Parrya. In: K. H. Rechinger (ed), Flora
Iranica. Vol. 57. Akademicsche Druck-u.
Verlagsanstalt, Graz.
Regel, E. (1862). Aufzaehlungder von Radde in
Baikalien, Dahurien und am Amur sowie der vom
Herrn von Stubendorff auf seiner Reise durch
Sibirien nach Kamtschatka, und der von Rieder,
Kussmisscheff und anderen in Kamtschatka
gesammelten Pflanzen. I. Abteilung.
Dicotyledoneae. Polypetalae. Bull. Soc. Imp.
Naturalistes Moscou 34: 1 – 211.
____ (1870). Supplementum II ad enumerantionem
plantarum in regionibus cis- et trans-iliensibus a cl.
Semenovio anno 1857 collectarum. Fasc. 1.
Ranunculaceae, Berberideae, Nymphaeaceae,
Papaveraceae, Fumariaceae et Cruciferae. Bull.
Soc. Imp. Naturalistes Moscou 43: 237 – 283.
____ (1877). Descriptiones plantarum novarum et
minus cognitarum. Fasciculus V. Trudy Imp. S.Peterburgsk. Bot. Sada 5: 217 – 272.
Rollins, R. C. (1993). The Cruciferae of Continental North
America. Stanford University Press, Stanford.
Royle, J. F. (1834). Illustrations of the botany and other
branches of the natural history of the Himalayan
Mountains and of the flora of Cashmere. Wm. H. Allen
& Co., London.
Rydberg, P. A. (1912). Studies on the Rocky Mountain
flora — XXVII. Bull. Torrey Bot. Club 39: 301 – 328.
Schulz, O. E. (1927). Beiträge zur Kenntnis der
Cruciferen des nordwestlichen Himalaya-Gebirges
Notizbl. Bot. Gart. Berlin-Dahlem 9: 1057 – 1095.
____ (1936). Cruciferae. In: A. Engler & H. Harms
(eds), Die näturlichen Pflanzenfamilien. Ed. 2, 17B:
227 − 658. Verlang von Wilhelm Engelmann,
Leipzig.
Schur, F. (1853). Sertum florae Transsilvaniae. Verh.
Mitth. Siebenbürg Vereins Naturwiss. Hermannstadt. 4:
65 – 72.
Smith, W. W. (1914). Diagnoses specierum novarum in
herbario Horti Regii Botanici Edinburgensis
cognitarum. LI – CII. Notes Roy. Bot. Gard. Edinburgh
8: 173 – 212.
475
____ (1920). Diagnoses specierum novarum in
herbario Horti Regii Botanici Edinburgensis
cognitarum. CCCCI – CCCCL. Notes Roy. Bot. Gard.
Edinburgh 11: 191 – 232.
Soják, J. (1982). Einige Bemerkungen zur Flora der
UdSSR (1). Sborn. Nár. Mus. v Praze, Řada B, Příř.
Vědy. 38B (1, 2): 101 − 109.
Sprengel, K. (1825). Systema Vegetabilium. Vol. 2.
Librariae Dieterichianae, Göttingen.
Sweet, R. (1830). Hortus Britannicus. Ed. 2. James
Ridgway, London.
Tidestrom, I. (1925). Flora of Utah and Nevada. Contr.
U.S. Natl. Herb. 25: 1 – 665.
Trautvetter, E. R. (1860). Enumeratio plantarum
songoricarum a Dr. Alex. Schrenk annis 1840 –
1843 collectarum. Bull. Soc. Imp. Naturalistes Moscou
33: 55 – 162.
____ (1871). Conspectus florae insularum NowajaSemlja. Trudy Imp. S.-Petersburgsk. Bot. Sada 1(1): 43
– 88.
Vassiljeva, A. N. (1961). Cruciferae. In: N. V. Pavlov (ed),
Flora Kazakhstana 4: 171 – 339. Izd-vo-Akademii nauk
Kazakhstoi SSR, Alma-Ata.
____ (1969). Species novae generum Parrya R. Br.,
Ermania Cham., Rhammatophyllum O. E. Schulz ex
Asia Media. Bot. Mater. Gerb. Inst. Bot. Acad. Nauk
Kazakhst. S.S.R. 6: 17 − 26.
____ (1974). Commentationes de systematica generis
Neuroloma Andrz. Bot. Mater. Gerb. Inst. Bot. Acad.
Nauk Kazakhst. S.S.R. 8: 27 − 31.
Vvedensky, A. I. (ed) (1974). Conspectus Florae Asiae
Mediae. Vol. 4. FAN Press, Tashkent.
Watson, S. (1895). Parrya. In: A. Gray & S. Watson (eds),
Synoptical flora of North America. Vol. 1, pt. 1. American
Book Co., New York, Cincinnati & Chicago.
Yue, J. P., Sun, H., Li, J. H. & Al-Shehbaz, I. A. (2008).
A synopsis of an expanded Solms-laubachia
(Brassicaceae) and the description of four new
species from western China. Ann. Missouri Bot.
Gard. 95: 524 – 542.
Zhou, T. Y., Lu, L. L., Yang, G. & Al-Shehbaz, I. A.
(2001). Brassicaceae (Cruciferae). In: C. Y. Wu & P.
H. Raven (eds), Flora of China. Vol. 8. Science Press,
Beijing & Missouri Botanical Garden Press, St.
Louis.
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013