A Review of Gamosepaly in the Brassicaceae and a Revision of Desideria, with a Critical Evaluation of Related Genera

15 Pages·2000·13.7 MB·

by  Ihsan A Al-Shehbaz

REVIEW GAMOSEPALY
A OF
Ihsan AUShehbaz^
A.
THE AND
BRASSICACEAE
IN
A REVISION OF
DESIDERIA,
WITH
A
CRITICAL
EVALUATION OF RELATED
GENERA^
Abstract
Ganiosepaly is reported in 12 genera of the Brassiiaeeae and is considered to have evolved indepcndenlly as many
conckided gamnsepaly
times. It is thai is not a useful character for the circumscription of genera in the family. The
boundaries of Desideria and several other genera are critically evaluated, and a taxonomic revision of Desideria
is
A
new name
presented. new
{D. haranerisis), eight condjinations {D. baiogionemis, D. flabellata, D. himalayensis, D.
incana, D. linearis, D. prolifera, D. pumila, and D. stewariii), and eight new synonyms (Christolea karakorumensis,
C.
Ermania
pinnatifida, C. scaposa, Desideria pamirica, hi/aria, E. kachoori, E. kashmiriana, and E. pa rkeri) are proposed.
As
herein delimited, Christolea consists of oidy two species; Ermania is reduced to synonymy of Melanidion, and
Ermaniopsis and Oreohlastus are reduced to synonymy of Desideria. The relationship and distinguishing characters of
Ermania,
Desideria, Christolea, Eurycarpus, hnospora, Melanidion. and Solmslaubachia are discussed.
Ke) words: Brassicaceae, Christolea, Desideria, Emiania, Eurycarpus, gamosepaly, Leiospora, MelanUlion, SolmsUiubachia.
During work on the Brassicaceae (Cruciferae) for America. was reported by Oliver (1893) in
It first
&
the Flora of China, Flora of Nepal, and Flora of Ka- Braya uniflora Hook. Thomson. Hooker and
f.
became Thomson
zahstan, evident that the hmits of several (1861) and Hooker and Anderson (1872)
it
Himalayan and Central Asian genera needed critical did not report gamosepaly in the species even
many
evaluation, and the nomenclature of species and though the type collection has flowers and
all fruits
needed The
infraspecific taxa adjustments. genera ad- with persistent, united sepals. Schulz (1924) trans-
dressed in the present paper are Christolea Cambess., ferred the species the monotypic Pycnoplinthus
to
Desideria Pamp., Emuinia Cham, ex Botsch., Emuin- 0. E. Schulz, a genus restricted China and Kash-
to
iopsis H. Hara, Eurycarpus Botsch., Leiospora (C. A. mir (Jafri, 1973; Kuan, 1987; Hajra et al., 1993).
Mey.) Dvorak, Melanidton Greene, Oreoblastus Sus- Desideria mirnhilis Pamp. (China, Kashmir, Ta-
lova, and Solmslaubarhia Muschl. They exhibit over- jikistan) the second species reported have a
is to
lapping similarities in several characters, and their gamosepalous calyx, and too was placed in a
it
have been
limits often confused. monotypic genus (Pampanini, 1926, 1930). Hedge
Because Desideria was based on a species with (1968b) described Sisymbrium gamosepalum
a gamosepalous calyx, a review of gamosepaly in Hedge and Arabidopsis gamosepala Hedge, both
of
the Brassicaceae presented to determine whether which endemic
is are to Afghanistan, but the latter
or not this character alone is sufficient to establish species was transferred by Al-Shehbaz and O'Kane
The &
genera. study led to the revision of Desideria Hedge
(1997) Neotorularia Leonard.
to Sis-
J.
and
also critically evaluated the limits of several ynibrium L. 50 species; Al-Shehbaz, unpub-
(ca.
presumably
related genera.
lished) represented by indigenous species on
is all
and
continents except Australia Antarctica
(Al-
Gamosepaly thk Bhassicaceae
in
Shehbaz, 1988), whereas Neotorularia includes
Gamosepaly has been reported in at least 12 about 15 species distributed primarily in Central
genera of the Brassicaceae from Asia and South Asia and the Middle East (Al-Shehbaz, unpub-
am
most grateful to Zhu Guanghua and Song Hong for their help in the translation of Chinese text and herbarium
I
Hank
labels, to Tatyana Shulkina for help with the Russian literature, and to van der Werff, Gerril Davidse, Nicholas
Turland, and Michael Gilbert for their advice on some nomenclalural problems. am profoundly thankful A. R.
to
I
Naqshi for sending duplicates of type material. Su/anne Warwick, Peter Heenan, and an anonymous reviewer are
I.
thanked for their critical review of the manuscript. I also thank the curators and directors of the herbaria cited in this
am my
paper. grateful to Oliver Appel and Juan Martfnez-Laborde for bringing to attention gamosepaly in Catadysia
I
Eudema
rosulans and respectively.
friesii,
2 Missouri Botanical Garden, P.O. Box 299, Louis, Missouri 63166-0299, U.S.A.
St.
Ann. Missouri Bot. Card, 549-563. 2000.
87:
550 Annals
the
of
Garden
Missouri Botanical
lished). Sisymbrium and Neotorularia each includes pals are free, but they appear connate because of
only a single species with a ganiosepalous calyx. interlocking stellate trichomes (Dudley, 1964).
Two
additional species of Desideria, pamirica In t'onclusion, two generalizations be
Z). 1
from Tajikistan (Suslova, 1973) and nepalensis regarding gamosepaly. First, the union of sepals
Z).
from Nepal (Hara, 1975), were described with a evolved independently several times in the Bras-
gamosepalous calyx. The reports of gamosepaly in sicaceae. not known whether evolvt-d one or
It is it
C
Christolea scaposa by Jafri (1973), karakorumen- more times within Desideria, but a phylogenetic
Wu
An
sis by and (1994), and D. pamirica are study based on molecular data should reveal that.
shown in the present study to be erroneously based With the critical examination of more genera of
on plants of D. mirahilis. Brussicaceae, likely that more species with
is
it
Gamosepaly was reported from South Amer- gamosepalous calyces be found. Second, ga-
will
first
&
lea by Al-Shehbaz (1990b) in Brayopsis Gilg mosepaly alone cannot be used to define the bound-
Muschl,, a genus of six species of which only B. aries of genera because occurs in several genera
it
gamosepala Al-Shehbaz (Bolivia) has united sepals. in which the majority of species have free sepals.
An
examination of other South American species Therefore, in the present delimitation of Desideria
revealed gamosepaly in Catadysia rosularis 0. E. gamosepaly ignored as a generic character, and
is
Eudema
Schulz (Appel, pers. comm.) and friesii 0. the overall similarities and relationships of species
E. Schulz (Martinez-Laborde, pers. comm.). Eude- are emphasized.
ma &
Humb. known
Bonpl. includes six species distrib- Nothing about the inheritance of ga-
is
uted from Ecuador into Argentina and Chile (Al- mosepaly in the family, but the occurrence of plants
Shehbaz, 1990a), of which only £. friesii has a with free and united sepals in the same population
gamosepalous calyx, whereas Catadysia 0. E. oi Phaeonychium jafrii is a good lead for conduct-
Schulz is a monotypic genus endemic to Peru ing a simple experiment to test the genetic basis of
(Schulz, 1929, 1936). this character.
Gamosepaly has recently been discovered in one
of six species of the Himalayan Pegaeophyton Hay- GKNKliic Rei.A'IIONSIUPS AND CiMCl MSCMlin lONS
&
ek Hand.-Mazz., Al-Shehbaz
tcatsonii of Sik-
P.
kim (Al-Shehbaz, 2000a), and in one of six species
of the Himalayan and Central Asian Phaeonychium Pampanini (1926) established the monotypic De-
0. E. Schulz, P. jafrii Al-Shehbaz (Al-Shehbaz, sideria solely on the basis of having a gamosepalous
2000b), although the type collection of the latter calyx. Although he indicated that mirahilLs re-
Z).
has plants with free and united sepals. Solmslau- sembles what was then known as Cheiranthus him-
bachia xerophyta (W. W. Sm.) Comber (China) also alayerisis Cambess., Schulz (1927a, 1936), Bot-
has calyces with either free or completely united schantsev (1955, 1956), and Jafri (1955) regarded
&
sepals, whereas gamosepala Al-Shehbaz G. gamosepaly as an anomaly and reduced mira-
5.
Z).
C
Yang (China), which is known (mly from the type bilis to synonymy of himalayensis, a species that
&
collection, has united sepals (Al-Shehbaz Yang, Schulz and Botschantsev assigned Ermania and
to
2000b). However,
Jafri to Christolea. these authors over-
At least one of the approximately 150 species of looked the significant features (see below) that dis-
Erysimum DC,
L., E. siliculosum (M. Bieb.) has a tinguish these two species. With the description of
gamosepalous The was
calyx. species previously two additional species in Desideria (Suslova, 1973;
recognized in Syrenia Andrz., a genus that I place Hara, 1975), the genus was recognized as distinct
synonymy Erysimum. some
in the of likely that in subsequent works Czerepanov,
It is floristic (e.g.,
of the species related to E. siliculosum also have 1995; Hara, 1979; Pachomova, 1974; Yunussov,
gamosepalous calyces, but have not examined ad- 1978), and remained be delimited primarily
I to
it
equate material of those. on the basis of having a gamosepalous calyx.
A
Pugionium
In all four species of Gaertn. (north- critical evaluation of all genera related to De-
em
and
China, Mongolia, adjacent Siberian Russia) sideria in this paper leads to the conclusion that
As
the sepals are connate. the fruit develops, the the genus should include 6 of the 10 species treat-
calyx ruptures basally along the lines of sepal con- ed in Ermania by Schulz (1936), 8 of the 10 spe-
Ermania
nation. cies recognized by Botschantsev
in
Finally, the genus Gamosepalum Hausskn. was (1955), and 5 of the 13 species assigned to Chris-
initially thought to have a gamosepalous calyx tolea by Jafri (1955). The species recognized by
(Schulz, 1927b, 1936). However, careful examina- these authors in Christolea or Ermania and exclud-
component
tion of its species revealed that the se- ed from Desideria in the present account are: Par-
Volume Number
4
87, Al-Shehbaz
551
2000
Gamosepaly
Brassicaceae
in
Ihiflorus regardless of the interpretation of effective date
its
Anderson, which now belong Phaeonychium
to (Al- of valid publication, does not affect the nomen-
it
Shehbaz, 2000b); Draba parryoides Cham, and Me-
clature of the unrelated taxa herein placed De-
in
lanidion boreale E. L. Greene, which are assigned
sideria.
Melanidion
to (s Although superficially resembling some species
if^
Cambess., which is retained in Christolea; and Par- of Desideria and Christolea, Ermania parryoides
is
&
rya lanuginosa Hook. Thomson, which
most closely related Melanidion
f. is to boreale E. L.
&
placed in Eurycarpus Botsch. (Al-Shehbaz Yang, Greene. Both species have Arctic and subarctic
2000a).
distribution (the Russian Far East for the former
As
herein delimited, Desideria consists of 11 Hi- and Alaska, Yukon, and Northwest
Territories for
malayan, Chinese, and Central Asian species char- the latter) and are similar in habit, foliage, pubes-
acterized by having well-defined basal rosettes, cence, flowers, and morphology. Huhen
fruit (1945)
slender and rhizome-like caudices, orbicular or fla- was the first to point out this close relationship, and
bellate to broadly ovate or obovate, often dentate he transferred M, boreale to Ermania, but his trans-
and
palmately veined basal leaves, simple and/or was because Ermania was
fer illegitimate invalidly
forked trichomes, linear to linear-lanceolate lati- published. The principal difference between these
septate fruits rectangular in cross section, nonto- species that E, parryoides has
is latiseptate fruits
and
rulose strongly veined valves with distinct mar- (flattened parallel to the septum) and M. boreale has
ginal veins, valve apices united with the replum, angustiseptate fruits (flattened at a right angle to
often obsolete styles, 2-lobed stigmas, and accum- the septum), but this difference not as significant
is
A
bent cotyledons. combination of fruits rectangu- as once thought because there are many genera
of
lar in cross section, valves with prominent marginal the Brassicaceae with both Drury and
fruit types.
veins, valve apices united with the replum, often Rollins (1952) and Rollins (1993) reduced Melan-
and
obsolete styles, dentate leaves often palmately idion synonymy Smelowshia
to of C. A. Mey., but
veined readily distinguish Desu their circumscription of the North American Sme~
genera discussed in this paper. lowskia was so broad that some of the species rec-
ognized are doubtfully congeneric. M.
boreale
If
and
CHRISTOLEA, ERMANIA, AND MELANIDION E, parryoides were kept in a genus distinct
from
Smelowskia, as presently support, then Er-
I
in his description oi Draba parryoides, Chamisso mania would have be abandoned and
to the earlier
"DRABA?
(1831: 533) stated, parryoides n. sp. vel published Melanidion recognized. These two spe-
novum
potius genus e solo fructu, deficiente flore, cies will be dealt with in a subsequent publication.
baud
rite definiendum. Drabis dolichocarpis sub- In his original description of Cheiranthus hima-
jungimus pro tempore plantam aliquando
fors jure layensis Cambessedes
ifi
ERMANIAM
meritoque nomine
inventoris />arryoi- (1844) did not indicate anything about
their rela-
dem
slutandam." Several workers
(e.g., Schulz, tionship or similarities to each other. However, Jafri
1936; Botschantsev, 1955; Hedge, 1968a; Suslova, placed them and
(1955) several other species
in
&
1972; Ovczinnikov Yunussov, 1978; Greuter et Christolea and adopted a broad generic concept
al., 1993) considered the above statement as a valid that included species presently assigned the gen-
to
publication of the genus Ermania, while others era Christolea, Desideria, Eurycarpus, Melanidion,
(e.g., Jafri, 1955, 1973; Jurtsev, 1975; Berkutenko, Parrya R. and Phaeonychium. With such
Br., a
1988; Czerepanov, 1995) did not. According Ar- broad
to delimitation, several additional
es-
g
tide 34 of the Code (Greuter et al., 2000), Cham- pecially Pegaeophyton and Pycnoplinthus, could
isso s statement does not constitute valid publica- have been included
easily in Christolea without ex-
tion of the genus. Despite Schulz's (1936) detailed panding the generic limits any further. Unfortu-
description of Ermania, was in German, and the was
it nately, Jafri's delimitation of Christolea closely
genus remained invalidly published until Botschan- followed in some of the more recent An,
floras
(e.g..
tsev (1956) provided the Latin diagnosis. Therefore, 1987, 1995; Hajra 1993; Huang, 1997b;
et
al.,
Ermania
all transfers to proposed by Schulz Kuan,
1985).
(1927a, 1933a, b, c) and Botschantsev (1955) re- Ovczinnikov and Yunussov (1978) also adopted
mained As
invalid. it is presently delimited, Er- a rather broad concept of Ermania by including
mania
includes only E. parryoides (Cham.) Botsch., Christolea and Oreoblastus These
as sections. au-
and
the generic type, other species assigned from
all to thors differed Jafri (1955) primarily in their
it belong to other genera. Ermania does not occur decision about the effective date of valid publica-
in the Himalayas and Central Asia and, therefore, tion of Ermania. In my opinion, their vastly het-
Annals
the
of
Garden
Missouri Botanical
erogeneous generic circumscriptions of Christolea without marginal veins, broadly lanceolate ob-
to
Ermania
or are unacceptable. Christolea consists of long fruits narrowly elliptic in cross section, well-
two species, the Himalayan C. crassifolia and the defined subconical styles, and minute, entire stig-
Chinese endemic C. niyaensis Z. X. An, and dif- mas much narrower than the style. By contrast,
it
fers from Melanidion (including Ermania) by hav- Desideria almost always has dentate, palmalely
ing many-leaved stems, nonrosulate lower leaves, ined leaves, prominently veined valves with well-
exclusively simple trichomes, incumbent cotyle- developed marginal veins, linear linear-lanceo-
to
dons, apiculate anthers, and transversely oriented late fruits rectangular in cross section, obscurely
seeds. By contrast, Melanidion has leafless stems, differentiated or cylindric styles, and distinct, often
well-developed basal rosettes, dendritic trichomes 2-lobed stigmas as broad as the style.
mixed with simple ones, accumbenl or obliquely
accumbent cotyledons, obtuse anthers, and longi- OREOBLiSTVS
tudinally oriented
ids.
Although admitted
(1973) the of
Jafri artificiality
Desideria from both Melanidion and
differs
he
his delimitation oi Christolea, 155)
c 'lly
(p.
by having
Christolea fruits rectangular in cross sec-
"Even Camb. and
stated that, Christolea
if, (s. str.)
tion, valves with prominent marginal veins, and
Ermania Cham,
ex [Botschantsev] Schulz
(s. str.)
From
valve apices united with the replum. Chris-
are considered as separate genera, there can be no
Desideria by having well-developed
tolea^ differs a
doubt that Oreoblastus Suslova congeneric with
is
basal rosette, usually leafless stems, slender and
Desideria Pamp., where most of our species would
rhizome-like caudices, often palmately veined
7 7
nontorulose longitudinally oriented
leavt\s, fruits,
Suslova (1972) separated Oreoblastus from De-
and accumbent By
biseriate seeds, cotyledons. con-
sideria by having free instead of united sepals, a
Christolea has nonrosulate lower leaves, leafy
trast,
deciduous and
instead of persistent calyx, septate
stems, compact and woody caudices, pinnately
instead eseptate However, she must have
ol fruits.
veined
leaves, strongly torulose transversely
fruits,
overlooked
the persistent calyx in several speci-
and
oriented uniseriate seeds, bent cotyle-
i
and
that she annotated as Oreoblastus^ the
dons. The Himalayan and Central Asian Desideria
holotype of her D. pamirica (Suslova, 1973) hae
also differs from the Arctic and subarctic Melani-
septate instead of eseptate though the septa
fruits,
dion by lacking the dendritic trichomes and having
perforated but never lacking. Except for having
2-lobed stigmas, smooth biseriate
ile fruits,
free instead of united sepals, Oreoblastus indis-
is
and median
seeds, toothed nectaries lacking the
As
tinguishable from Desideria. indicated above,
glands. Melanidion has dendritic trichomes, entire
sepal connation alone insufficient for the estab-
is
and
capitate stigmas on distinct styles, torulose
lishment of genera and, therefore, Oreoblastus
is
and
uniseriate or rarely subbiseriale seeds,
fruits,
reduced herein to synonymy of the earlier pub-
median
annular nectaries with well-developed
lished Desideria.
A
glands. comparison oi Desideria with the presum-
ably related genera is summarized in Table 1.
EHMANIOPSIS
The presence absence on me-
EURYCARPUS vs. of a tooth the
dian stamens was considered by some Schulz,
(e.g.,
In establishing the genus Eurycarpus, Botschan- 1936; Kara, 1974; Golubkova, 1976) as an impor-
Ermania my
tsev (1955) separated from by having tant generic character. In opinion, feature
it this
biseriate instead of uniseriate seeds, broadly Ian- alone does not justify the segregation of genera.
ceolate instead of linear fruits, entire instead of Toothed and toothless filaments are found in Don-
&
and
dentate leaves, leafless instead of leafy scapes, tostemon Andrz. ex A. Mey. (Al-Shehbaz
C.
However, he probably compared only the type spe- Ohba, 2000), whereas winged or wingless, toothed
cies of both genera because most of the differences and appendaged unappendaged
or toothless, or
fil-
above do not hold if one compares Eurycarpus with aments are found mAlyssum L. (Al-Shchbaz, 1987;
Erman-
the ten species Botschantsev recognized in Dudley, 1964).
As
ia. indicated above, eight of Botschantsev's ten Although Kara (1974) provided a detailed dis-
Ermania
species of are presently assigned to De- cussion to distinguish Ermaniopsis from Ermania
A
sideria. comparison of Desideria with Eurycarpus and related genera, the single character
that sets
(two species) sensu Al-Shehbaz and Yang (2000a) Ermaniopsis apart the presence of a lateral tooth
is
shows that the latter differs by having entire and on the filaments of median stamens. On the basis
pinnately veined leaves, obscurely veined valves of all other characters, Ermaniopsis pumila H. Hara
I
Volume Number
4 Al-Shehbaz 553
87,
2000 Gamosepaly
Brassicaceae
in
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554
Annals
the
of
Garden
Missouri Botanical
home
is perfectly at in Desideria, In fact, Hara in- Desideria Pamp., Bull. Soc. Bol. Ilal. 1926: 111.
TYPE;
dicated that E, pumila resembles Desideria (as Par- 1926. Desideria mirahilis Pamp.
A
pumila
rya) in vegetative characters. close ex-
Ermani}mis
amination of flower and fruit characters clearly H, Hara, J. Jap. Bot. 49: 198. 1974. TYI'E:
Ermcniopsis pumila H. Hara.
shows that the two species are congeneric, and Er-
&
(Moscow
Oreobhistus Suslova, Bol. Zlmrn. Leningrad)
maniopsis reduced herein to synonymy of Desi-
is
TYPE:
57: 648. 1972. Oreoblastusflabcllatus (Regel)
deria. Unfortunately, both species have the same
Suslova.
named
and pumila
epithet, E, hereafter as D.
is
Herbs many-
The median pumila perennial, with a slender, often
haranetLsis. filaments of both D.
and D. haranensis are dilated, and only the latter branched, rhizome-like caudex often covered with
species shows a minute to prominent tooth on the remains of basal rosettes. Trichomes simple and/or
mixed
median with short-stalked forked ones. Stems sim-
staminal filaments.
ple, leafy or leafless, sometimes absent. Basal
AND
SOLMSIAVBACHIA LEIOSPORA leaves petiolate, rosulate, simple, 3- to 9(to 11)-
^^*^" palmately vemed, persisting whole or
^^^*^^^'
On
the basis of morphology, Desideria
fruit
is
persistent. Cauline leaves similar
to
^"^^
8 P^^^^^^'
most Solmslaubachia
closely related to (9 spp.:
^^^^^ ^"^^^^ ^^^^^^^^' sul)sessile or petio-
""'
endemic China and extending Bhutan
to also into
1
''''^f^
Racemes
late, or absent. 3- to 30-flowered, dense
and Sikkim) and the Central Asian and Himalayan
or lax, bracteate throughout or ebracteate, corym-
Leiospora (6 spp.). All three g h fruit
^«^^' elongated or not elongated in someti
fruit,
detached from and
readily the pedicel, their valves
^^^^'^ '^^^*^^ ^" originating from basal
Upon P^^^^^^'
are adnate apically to the replum. maturity,
'"^^^**^- ^^pals ovate to oblong, free or united, de-
the fruit falls off the plant and apex remains
its
^^^^"^^^ persistent, erect, equal, base of inner
The ""'
tardily dehiscent. three genera also have ob-
margins membranous.
"^^ saccate, Petals pur-
solete or no styles, and their fruit valves are strong- P^^""
purple-green, or rarely white, sometimes yel-
angled the margins and completely conceal P^^'
ly at
^^^^^^ ^^^^ ^^^^^^ ^^^^'^^^ spatulate,
the replum. These combinations of characters are ^* ^^ ^^^^^^^ ^^
^P^^ ""^^^^^ *« subemarginate; claw strongly differ-
not found in any Himalayan or Central Asian gen-
entiated from blade, subequaling or longer than se-
era of the Brassicaceae.
Stamens erect, tetradynamous; filaments
6,
Desideria easily separated from Solmslauha- P^^^'
is
wingless or rarely winged, toothless or rarely
chia by having palmately veined leaves apically 3-
toothed, free, dilated at base: anthers ovale to ob-
to 9(to ll)-toothed, ovate to oblong anthers often
mm
apiculate apex. Nectar glands 2 and
less than long, and forked trichomes some- ^^^g' "*^* at
1
and
or confluent outside bases of
mixed By lateral, 1 all sta-
times with simple ones. contrast, Solms-
median
nectaries present or absent. Ovules
lauhachia has entire, pinnately veined leaves, lin- s;
ear-oblong anthers more than mm long, and 10 to 70 per ovary. Fruit dehiscent siliques, line*
1
to lanceolate, latiseptate, rectangular in cross
exclusively simple trichomes. <
inflated, sessile; valves papery, with a
Desideria readily distinguished from Leiospora ^i^'^' '^«t
is
prominent midvein and marginal
by having wingless seeds, equal sepals with the distinct veins, gla-
lat-
b^ous or pilose, smooth, adnate with replum
at fruit
eral pair nonsaccate, palmately veined leaves api-
apex; replum rounded, often concealed by valve
3-
cally to 9(to ll)-loothed, oblong-linear anthers
mm
0.4-l(-L6) long, and capitate, slightly 2-lobed margin; septum complete, perforated, or reduced to
membranous,
a rim, translucent, veinless, rarely
stigmas with neither decurrent nor connivent lobes.
Leiospora often has winged or margined seeds, un- absent; style obsolete; stigma capitate, slightly 2-
equal sepals with the lateral pair strongly saccate, lobed. Seeds uniseriate or biseriate, wingless, ob-
entire or marginally dentate leaves, linear anth long to ovate, often flattened; seed coal obscurely
mm
when
2.5-3 long, and conical, prominently 2-lobed reticulate, not mucilaginous wetted; cotyle-
don
;cumbent.
stigmas with connivent, decurrent
lobes.
Eleven species: Himalayas, western China, and
Taxonomic Tkkatmfni
adjacent central Asia.
Kky to Spkciks ok Desideria
tiik
la. Sepals united, persistent till or after frnil deliiscence; septum absent or reduced to a rim.
11-13 X 5-6 mm; 5-6 nun 2—
2a. Petals calyx lori^; (lowers appearing solitary; Ne[>al 11. D. nvpaleiisis
1, ....
2b. Petals and ealyx smaller; flowers more than 4, in distinct racemes; China, Kashmir, Tajikistan
10. mirdbilis
I),
Volume Number 4 Al-Shehbaz 555
87,
2000 Gamosepaly
Brassicaceae
in
lb. Sepals free, caducous or rarely persisting about fruit maturity; septum complete or rarely perforated
til]
apically.
3a. Flowers solitary from a basal rosette.
mm
6—9
4a. Fruit ovate to broadly lanceolate, wide, prominently reticulate veined 9. D. baiogionensis
.._
mm
2—5
4b. Fruit linear linear-lanceolate, wide, obscurely veined.
to
3^ mm
5a. Leaf trichomes forked and simple; replum retrorsely pilose; valves glabrous; sepals
mm
6—8
long; petals long 8. D. piimila
mm
6—7
5b. Leaf trichomes exclusively simple; replum and valves pilose to villous; sepals long;
mm
11—14
petals long 7. D. prolifera
3b. Flowers (3 to)6 to 30 in a raceme.
Racemes
6a. bracteate throughout.
7a. Stem and pedicel trichomes forked 2. D. stewartii
7b. Stem and pedicel trichomes exclusively simple or absent.
mm 3^
X mm;
8a. Fruit lanceolate to linear-lanceolate, (3—)4—6 wide; petals (6—)6.5—8
mm
X
seeds biseriate, (1.5-)1.8-2(-2.3) 1-1.4 himalayeriMs
1. /).
mm
4— X mm;
8b. Fruit linear, (0.8-)l-1.7(-2) wide; petals 5(-5.5) 1.5-2.5 seeds uni-
mm
X
seriale, 0.8-1.1 0.5-0.8 3, D. linearis
Racemes
6b. ebracteate.
mm
9a, Filaments flattened, subapically toothed; petals 6.5—8 long; leaf trichomes minutely
forked, mixed with short simple ones 6. D. haraneiisis
mm
11-18
9b. Filaments terete, toothless; petals long; leaf trichomes either exclusively simple
or distinctly forked.
10a. Plants canescent; leaf trichomes almost exclusively branched; leaves 3(to 5)-toothed
D, incaria
5.
mm
10b. Plants greenish; leaf trichomes exclusively simple, to 1.5 long; leaves (3 to)5- to
9(to ll)-toothed 4. D. Jlabellala
2^ mm mm
1. Desideria himalayensis (Cambess.) Al-Sheh- pair long; anthers ovate, ca. 0.6 long,
baz, comb. nov. Basionym: Cheiranthus him- Ovules 7 to 12 per locule. Fruit lanceolate to lin-
cm X mm,
alayensis Cambess., in Jacquemont, Voy. Inde ear-lanceolate, (1.7-)2-3.5(-4) (3-)4-6
Ermania
14. 1844. himalayensis (Cambess.) strongly flattened; valves pilose or glabrous, dis-
4:
septum membranous;
0. E. Schulz, Notizbl. Bot. Gart. Berlin-Dah- tinctly veined; complete,
lem 1080. 1927. Oreoblastus himalayensis style obsolete; stigma 2-lobed. Seeds brown, ovate,
9:
& X mm,
(Cambess.) Suslova, Bot. Zhurn. (Moscow (1.5-)1.8-2(-2.3) 1-1.4 biseriate, minutely
TYPE: [W
Leningrad) 57: 652. 1972. Tibet.] reticulate.
Kioubrung-
"In declivitate orientali jugi vulgo
Phenology. Flowering June through August.
Jacquemont
ghauti in Tartaria sinensi," Victor
mid
Fruiting July October.
to
1782
(holotype, P!; isotypes, K!, P!).
Habitat and distribution, Alpine tundra, open
sandstone 4300-5300 m. China (Qing-
4-20 cm scree;
Plants densely pilose throughout ^^i^l^^
tall,
mm
^ai, Xizang), India, Kashmir, Nepal.
to subglabrous. Trichomes simple, to 1.5 long.
Stems simple, pilose or glabrous. Basal leaves not ^.
,- r> , ? speci- mens examin• ed.i .L-HtliPitsvltA*. I^inghi ai.: tLei•-
n c>elected
. A
^ 0.4-
fleshy, pilose or glabrous, persistent; petiole &
Huang (HNWP).
Yang K~890 Xizang:
^ie.wu-den, Wu^
cm &
1.6(—3) long, not ciliate; leaf blade broadly ob- Baingoin Xian. Whale Lake, Wu, Ohba, Wu Fei 4075
&
ovate spatulate, 4-14 X 3-9 mm, base cuneate (KUN, MO, TI); E of Moincer, 3ri4'N, 80°56'E, G, S,
to
NW
Miehe 9643/17 (GOET, MO); 34°55'N,
Tibet,
margins apex
to attenuate, (3 to)5-toothed, acute.
Ho
82*^24^, Pike 842 Aksu, Deasy 92 (BM); Shuang
(K);
Stem
leaves similar to basal or linear to lanceolate,
Xian, Qinghai-Xizang Team 12009 (PE); Ritu Xian, Qing-
X mm,
5-17 1-4 &
often entire, short petiolate to hai-Xi=ang Team 76-9061 (KUN, PE), Li Bosheng
subsessile. Racemes 6- to 25-flowered, bracteate Zhang Du 10980 (PE). INDIA. Punjab: Lahul, Kangra,
KASHMIR.
Bara Laeh La. Koelz 6738 (GH). Hamag, Up-
throughout; bracts similar stem leaves but small-
to
per Lidder Slewart 9349 G, K, MO). NEPAL.
valley, (B,
sometimes adnate to pedicel. Fruiting pedicels
er,
Dhampus
Dhaulagiri Himal, hidden valley, between pass
mm
3-10
ascending, straight or curved, long, pilose
^nd French pass, Wald 65 (BM); Thorong La, Marsyandi
3^
X
or glabrous. Sepals free, oblong, 1.2—1.5 Valley, A/cBeaf/i i4S6 (E); Naurgaon, Marsyandi, Mcfiea//i
^^^^ Annapuma Himal. between N Annapurna Gla-
mm, (E);
caducous, pilose or with a terminal tuft of
camp N Annapuma
cier base and top of Glacier icefall,
base not saccate, margins membranous. Pet-
hairs,
km N
170-175 Pokhara, Komarkova 18 (GH).
of
als purple or lilac with yellowish center, broadly
3^
X mm,
spatulate, (6-)6.5-8 apex subemargin- Desideria himalayensis most frequently con-
is
mm
3-4 and some
ate; claw long. Filaments white, slightly fused with D, linearis, authors (e.g., Jafri,
mm
median 3-4 1955) considered them be conspecific. Three
dilated base, pairs long, lateral to
at
556
Annals
the
of
Garden
Missouri Botanical
mem-
collections of Desideria linearis {Lyon 44, Stainton brous, distinctly veined; septum complete,
3055, Stainton 3241) were cited by Jafri (1973) as branous; style obsolete; stigma 2-lobed. Seeds
X
Christolea himalayensis, and the first two were list- brown, ovale, 1.4—2.2 0.8-1.1 mm, biseriale, mi-
ed by Hedge (1968a) as Ermania himalayensis. The nutely
reticulate.
two species can be readily separated by petal
size,
Phenology, Flowering unknown.
Fruiting
in
width, and seed arrangement and
fruit (see
si
August.
Mixed
key). collections of the two species
(e.g..
and 4100—
Habitat Scree
distribution. slopes;
Koelz 6738) are not but no intemiedi-
, 5000 m. China (Xizang), India, Kashmir.
ates have been found. Both species can be
distin-
CHINA.
guished from the related D, stewartii by having leaf Selected specimens examined. Xizang: Ali,
Qinghaijihet Tea^^^
and stem trichomes exclusively simple instead of ^^y^
Bara Lacha Pass, Lanul, Cooper 5490
(E). IIiiiia<*lial
forked.
Pradesh: McBeath 2105
Ziiigzinfibar, (E).
Desideria himalayensis was reported Chris-
(as
known
An Desideria steivartii a very rare species
tolea) from Xinjiang by (1995), but have not is
I
One ^^^^ ^^^ f^^^" *he few collections cited above. Re-
seen any material from that part of China. col-
&
action {Polunin, Sykes Williams 37) was cited P^^^ ^^ ^^^ species from China (Kuan, 1985; An,
l^^"^) ^^^ ^^^^ ^^^^^ based on misidentified plants
by Hara
(1979) as this species, but this collection
of D. himalayensis,
clearly belongs Desideria
to linearis.
doubted
J^^^ (1973) the distinction of Desideria
Desideria himalayensis was erroneously
illustrat-
stewartii from D. himalayensis (both as Christolea),
ed
in Jafri (1973) with ebracteate inflorescences.
It
^"^ ^e confused the limits of the two species by
likely that the plant illustrated belongs D.fa-
is to
^^^ under
^^**"g collection, Stewart 9349, the for-
bellata, a species that occurs in bordering Afghan-
'"^^ instead of the latter species. have not seen
istan, China, Kyrgyzstan, and Tajikistan but not I
is
any flowering material of the species, and the de-
yet reported from Kashmir. Desideria himalayensis,
^^^ption of the flowers by which was
D, stewartii, and D, linearis are the only three spe- Jafri (1973),
An
fallowed by Hajra and was
cies of Desideria that consistently have racemes et al. (1993) (1987),
almost certainly based on a small flowering branch
bracteate throughout.
of D. linearis mounted on the holotype sheet of D,
4 4
stewartii.
Desideria
2. stewarlii Anderson) Al-Shehbaz,
(T.
In overall aspects of foliage and Desideria
fruit,
comb. nov. Basionym: Cheiranthus stewartii
T.
stewartii most closely resembles himalayensis.
Z).
Anderson, in D. Hooker, FL India
Brit.
J. 1:
However,
D. stewartii readily separated by the
is
Ermania
132. 1872. stewartii Anderson) 0.
(T.
presence of forked, stalked, intermingled trichomes
E. Schulz, Bot. Jahrb. Syst. 66: 98. 1933.
instead of exclusively simple straight ones.
Christolea stewartii Anderson) Notes
(T. Jafri,
Roy. Bot. Card. Edinburgh 22: 53. 1955. Or-
Desideria
3. linearis Busch) Al-Shehbaz,
(N.
eoblastus stewartii Anderson) Suslova, Bot.
(T.
comb. Basionym:
nov. Christolea linearis N.
&
Zhum. (Moscow
Leningrad) 57: 653. 1972.
Busch, in Komarov, Fl. URSS. 8: 636. 1939.
TYPE:
Kashmir. Ladak, 15,000-16,500 /
ft., Ermania
Busch) Ma-
linearis (N. Botsch., Bot,
L
Stewart s.n, (holotype, K!; isotype, E!).
Gerb. Bot. Komarova Akad. Nauk
ter. Inst.
S.S.S.R. 17: 166. 1955. Oreoblastus linearis
8-20 cm
Plants densely Trichomes
pilose.
tall,
&
(N. Busch) Suslova, Bot. Zhurn. (Moscow
some
stalked forked, rarely simple near the stem
TYPE:
Leningrad) 57: 652. 1972. Tajikistan.
Stems
base. simple, pilose or glabrous. Basal leaves
mm Pamir: Schugnan, At)chary, 2 Aug. 1904, B.
2-10
subfleshy, pilose, persistent; petiole long,
Fedtschenko
s.n. (holotype, LE!).
not ciliate; leaf blade broadly obovate spatulate.
to
X
2-15 2-10 mm,
base cuneate attenuate, mar- Ermania
to parkeri 0. E. Sclmlz. Repert. Sp. Nov. Regni
gins 3- to 5-toothed or subentire, apex acute. Stem Veg. 31: 333. 1933. Christolea parheri (0. E. Scluilz)
Jafri, Notes Roy. Bot. Card. Edinburgh 22: 1955.
7^2.
leaves similar to basal or linear to lanceolate, often
Oreoblastus parkeri (O. E. Schulz) Suslova,
liot.
Racemes
entire. 8- 15-flowered, bracteate
to
&
Zhurn. (Mosrow Leningrad) 57: 653. 1972. Syn.
throughout; bracts similar to stem leaves but small- TYPE:
nov. Kashmir. Sonamarg, Luderwas,
13,()00
U
er, often adnate to pedicel. Fruiting pedicels as- ft., Aug. 1928, R. R. Stewart 9874A (liulolype,
mm
4-12
cending, straight or slightly curved, l<mg, ^•)*
pilose. Flowers not seen. Ovules 12 per
7 to locule.
11. ^'"'"f '''''tZf'['k!^^'<^ ^''^^'•i-^l''t"\ ^"'S^l'
^** Tl
i, ^^^' ^^'*' 1990. byn. nov. PE: Kashmir. Shal-
, , o
. , , r, cm X
r-
Fruit lanceolate lanceolate-hnear, 1.7-3.5
to
Sonamarg 3900 20
j^^^,^ (Sind Valley), m, Aug.
3-5 mm,
strongly flattened; valves pilose or gla- 1983, Dar 7786 KASH).
G. (hoh)iype,
//.
Volume Number
4
87, Al-Shehbaz 557
2000
Gamosepaly
Brassicaceae
in
&
Ermania kachrooi Dar Bombay
Naqslii, Nat. Hist. ms
J.
Soc. 87: 277. 1990. Syn. riov. TYPE: Kashmir. Bal- KISTAN. Pamir: N slope, river Zor-Chechekly, 12 Aug,
&
Sonamarg 3200
tal, (Sind Valley), m, 2 Sep. 1982, \9^^. Stanjukovich Klshkovsky sm. (LE); Chechekty,
riv-
Dar 3934 KASH; KASH,
G. H. (holotype, isolypes, er Zor-Chechekty, Raikova 228 (LE).
MO!).
Although have not seen the holotype of Er-
I
cm
4-15
Plants densely pilose throughout
tall,
mania
kashmiriana, the original description and
mm il-
to subglabrous. Trichomas simple, to 1.5 long.
lustration, as well as the examination of a paratype
Stems
simple, pilose or glabrous. Basal leaves not
{Dar 8301), clearly support the placement of the
2—7
fleshy, pilose or glabrous, persistent; petiole
synonymy
species in of Desideria Dar and
mm linearis,
(-12) long, not ciliate; leaf blade broadly ob-
Naqshi compared
kashmiriana and
(1990) E, £".
X
4—15 2-12 mm,
ovate to spatulate, base cuneale
kachrooi with D, stewartii and D. himalayensis
(all
to attenuate, margins 3- to 5-toothed or rarely sub-
as Ermania)^ but they
failed to relate their novelties
apex Stem
entire, acute. leaves similar to basal or
^^"7'"" ^^h a
5-10 X mm, f ^" "^^ ''P'"^""' ^- ^<''^'^<^^ i«
1-3
linear lanceolate,
to often entire, *^
D
form
S'^brous oi linear a species with.n a giv-
us,
short petiolate subsessile. Racemes 8- 20-
to to
en population which one can and
of find glabrous
flowered, bracteate throughout; bracts
il
P^^^^^ent plants. In general, plants of Desideria,
stem leaves but smaller, often adnate to pedicel.
'"^lu^^ng grow shaded
mm ^- linearis, that in partly
2^(-12)
Fruiting pedicels ascending,
straight,
under
especially large boulders, h ave
^^^^^' oft
long, pilose or glabrous. Sepals oblong
free, to
^^^^^^ elongated
2-3 X mm, *^^ ^P^^ ^^ *^^ so that the leaf
1-1.5
ovate, caducous, pilose or with
much
appears
^^^^"^ less congested.
a terminal tuft of hairs, base not saccate, margins
Several authors An, 1987, 1995; Kuan,
membranous. (e.g..
Petals purple or lavender with paler
4— X mm, 1985) followed Jafri (1955) in listing Desideria
base, narrowly spatulate, 5(-5.5) 1.5-2.5 li-
mm synonym
nearis ^^ oi Christolea himalayensis, hni
apex rounded; claw 2-2.5
Filaments a.
long.
white, slightly dilated base, median pairs 2.5- ^^^^^ authors erroneously recognized C. parkeri as
at
mm mm my
^ distinct species. In opinion, the last species
3.5 long, lateral pair 1.8-2.5 long; anthers
C
mm
ovate, 0.4-0.5 long. Ovules 8 to 13 per locule. is only a glabrescent form of Z). linearis. In fact,
Fruit linear, (1.5-)2-3,5(-4,2) cm X (0.8-)l-1.7 Parkeri is based on Ermania parkeri, an invalidly
mm, published species assigned Ermania
to the invalid
(-2) flattened; valves pilose or glabrous, dis-
Greuter under
tinctly veined; septum complete, membranous; (^^^ et al. (2000) Article 43.1).
Although maintained
style obsolete; stigma 2-lobed. Seeds brown, ovate, Jafri (1973) ChrLstolea par-
X
mm, he
0.8-1.1 0.5-0.8 minutely correctly indicated that not different
uniseriate, retic- f^^^h it is
from
the earlier-published Desideria
ulat (as Christolea)
e.
linearis, a species that he did not recognize for Pak-
Phenology. Flowering June through August.
j^^an and Kashmir. However, Suslova (1972) main-
Fruiting July through September.
tained both species Oreoblastus) and separated
(as
and
Habitat distribution. Gravelly or sandy
^j^^^ ^^j^j^ ^y the presence in the former of a
morame
slopes scree, gravelly below
glacier;
.^bapical tuft of hairs on the sepals instead of
its
3200-5200 Chma
m. (Xinjiang, Xizang), Kashmir,
^ ^
^^^^^^^ -^ ^.^^^^^ ^^j
Obvi-
himalayer^sLs.
Nepal,
Tajikistan.
ously, this distinction and taxa have
is artificial, all
pubescent more
sepals that often are densely hairy
CHINA.
Selected specimens examined. Xinjiang:
below The
Yecheng Xian, Li Bosheng et al. 1278 (PE); Tagdum- the apex. restriction of trichomes to the
1
basch-Pamir, Pistan near Saryokol, Alexeenko 2729 (LE). sepals and leaf apices quite frequent in glabres-
is
cent forms of D. linearis and D. himalayensis.
S^^r^:
mT^^'' i^'^'''^^'^''™
n-''^T' n^o ?m^m^Tk;T.^
cui
kh, Qinghai-Xizang Team 76-7948
(Wn'^'V).
?); /looiv -j j j
r.
• f-
^^^"'^ (^^^l) considered Desideria
linearis (as
Punjab: Lahul, Kangra, Bara Lath La, Koelz 6738 (GH).
Ermania
KASHMIR. be
Fshkuman Aghost, Schmid 2449 Chitral: parkeri) to closely related to E. nlbifio-
(G).
UsF)ur (Hachin). 36°2'N, 72°27'F. /,von 44 (A, E); Siro- ra (T. Anderson) 0. E. Schulz, but the nearest rel-
gol S of Shah Jinali Pass, Slainlon 3055 (E); Dorah Pass,
^fi^g ^f the D. himalayensis. As shown by
first is
Lulko 3241 8709
\siUt-y, Stainton (E); Aniiiindlh, StairUon aicl l n
/nn.f\nt
Al-Shehi baz \ r- ji t i i
(2000b), E. albiflora belongs to the ge-
Sonamarg, Luderwas, 9874 MO). Ka-
(E); Stewart (B, G,
"^s Phaeonychium
O.
rakonini: Gharesa Glacier base ramp, 13 mi. E of Nagar, E. Schulz.
Polunin 6133 (BM); Karakorum, Oct. 1877, Clarke s.n. Desideria linearis is extremely variable in the oc-
Ladak:
(K). above Stok, /Wa^u,e// 92 (E); Zanskar, glacier
currence and indumentum,
density of the
fruit
Sentik, 34°N, 76°E, Oe/owc/ic 27 Thui region: .,, \,
(P). ca.
Mw , i and, m• du, mentum, and, numb, er r
i^ ri . n^ . widtri, lengtri, ot ieai
zoiI\Kij\ km JNW olf (Tilgi•t., near wat.ersuhedI separating Gilgit ' &
'
'
from Broadhead 39 Taklung 6500 teeth. However, an examination of material from the
Chitral, (E); I^, Koelz
NEPAL.
(GH). Naur Pass, Lowndes 1159 (BM); 5 mi. S of various parts of the species range clearly negates
558
Annals
the
of
Garden
Missouri Botanical
the need to subdivide the species into infraspecific *^'«J« *''• (LE, MO); Aklu Xian, Qiaornong, 5. C. Wu, Y.
&
My K 4641 (HNWF, KUN). KYKCiYZSTAN.
Fei
//.
taxa.
Seinirechie: Piz<*walsk, river Kayche, 30 July 1913,
Shishkin (LE), 2 1913, Capoznnikov (MO).
s.n. Auf^. s.n.
4. Desideria flabellata (Kegel) Al-Shehb
Tianshan: Glacier Brocherel 338 TAJIKIS-
Kaliul'i", (G).
comb. nov. Basionym: Parrya flabellata Kegel, TAN. Shugnan: E Bukhara, near Pass Garin-Chashniy,
Kumshdy
Bull. Soc. Imp. Naturalistes Moscou 43: 261. Tuturin ISO (A, EE). Pamir: Glacier, Gorbanoi
^^^^•
^^^
1870. Christolea flabellata (Kegel) N. Busch,
URSS
Komarov, 330. 1939. Ermania
in Fl. 8:
In every aspect of trichome morphology, flower
flabellata (Kegel) 0. E. Schulz, Bot. Jahrb.
^nd and
^[^^ color, habit, Christolea pinnatlfida
is
Syst. 66: 98. 1933. Oreoblastus flabellatus (Re-
indistinguishable from plants of Desideria flabella-
&
gel) Suslova, Bot. Zhurn. (Moscow Lenin- The
fa. type of the former has no fruits and is rather
TYPE:
grad) 57: 651. 1972. Southern Tian
immature. only from
differs slightly typical plants
It
Dschaman-Daban,
Shan, Sewerzow
(holo-
s.n.
ot D, flabellata by having sHghtly elongated, spat-
type, LE!).
ulate leaves instead of typically flabellate ones.
Huang
(1997a) considered be
C. pinnatlfida to re-
Christolea phinalifida R. Huang, Acta Phytotax.
Sin.
E.
35: 556. 1997. Syn. nov. TYPE: China. Qinghai: Ma- lated to C. karakorumensis, but the latter is a syn-
qen, Anyema<ien Ml., IIJOO m, 25 June 1981, R. F. onym of mirabilis and has sepals typically united
Z).
Huang
CG-81-154 (holotypr, IINWP!).
-^^^^^^
^f f^^^ jje indicated that the ovaries are
Plants greenish, 4-15 cm tall Trichomes simple, glandular mamillate, but this observation was based
mm
on developing trichomes, and neither Df'-s/V/ma nor
Stems
straight, to 1.5 long. distinct, simple,
2-7 Christolea has any glandular trichomes or papillae,
densely Basal heaves
pilose. subfleshy; petiole
mm
long, pilose; leal blade flabellate to broadly
X
obovate, rarely spatulate, 0.6-2.5 0.3-2.5 cm, 5. Desideria incana Al-Shehbaz, comb.
(Ovcz.)
pilose, base cuneate to attenuate, margins (3 to)5- nov. Basionym: Christolea incana Ovcz., Sov-
&
to 9(to ll)-toothed, rarely lowermost entire, apex etsk. Bot. 1941(1 151. 1941. Ermania
2):
X
mm.
acute; teeth to 10 3 Stem leaves similar to incana (Ovcz.) Botsch., Bot. Mater. Gerb. Bot.
Kacemes
basal. 7- to 12-flowered, ebracteate. Inst. Komarova Akad. Nauk S.S.S.K. 17: 164.
Fruiting pedicels ascending, curved,
straight to 1955. Oreoblastus incanus (Ovcz.) Suslova,
cm &
(0.5-)0.7-1.5(—2.5) long, spreading pilose. Se- Bot. Zhurn. (Moscow Leningrad) 57: 652.
5-8 X mm,
pals free, narrowly oblong, 1.5-2.5 of- 1972. TYPE: Tajikistan. Darvaz: Mt. Masar,
ten persistent, pilose, base not saccate, margins glacier Abdul Gassan, 11,000-12,000 23
ft.,
membranous.
Petals purple, broadly spatulate, 1.1 July 1899, Lipsky 1936 (holotype, LE!).
V. I.
cm X mm,
1.5 3.5—6 apex subemarginate; claw 7
4—15 cm
Plants densely tomentose through-
tall,
9 nun long. Filaments white mauve,
to slightly di-
mm Trichomes short-stalked forked and simple,
^"*- to
median 4.5-6
lated at base, pairs long, lateral
™"
3^ mm mm tomentose. Basal leaves
^^"g- ^^^'''^
^
pair long; anthers oblong, 0.9-1.3 ^'"^P^^'
subfleshy, canescent, densely tomentose, persistent;
long. Ovules 7 to 12 per locule. Fruit lanceolate to
^'^-^ """ unexpanded and
cm X "^* ^''^''^^""^
2.5-5 P^^^^^^ ^^"S-
lanceolate-linear, (1.7-)2.5-3.5(-4.5)
mm, "^^ P^P^^ '^^"^' '^^f '^^^^^ ^^^^'"y ^^^^^^^'^ ^^
^*
strongly flattened; valves pilose, distinctly
4^13 X 2-8 mm,
^P^tulate, base cuneate to atten-
septum membranous;
veined; complete, style ob-
margins sometimes
3(to 5)-toothed, subentire
"^*^'
solete; stigma 2-lobed. Seeds brown, ovate, 1.3-2
^" ^^^"^^ branches, apex acute. Stem leaves similar
X mm,
0.9-1.2 uniseriate, minutely reticulate.
Racemes
basal. 6- 20-flowered, only basally
to to
Phenology Flowering early July and August.
bracteate. Fruiting pedicels ascending, 2
straight,
mm
Fruiting late July through early September.
7 long, tomentose. Sepals narrowly ob-
free,
and
Habitat Alpine
distribution. gravelly 5-7 X mm,
J^^g, 1.5-2 caducous, densely tomen-
slopes, moraine slopes; 3300-5100 m. Afghanistan,
tose, base not saccate, margins membranous. Petals
China
(Xinjiang), Kyrgyzstan, Tajikistan.
12-
purple with paler or yellowish base, spatulate,
X 4-6 mm, 7-10
18 apex rounded; claw nun
AFGHANISTAN. long.
Selected specimens examine
d.
Filaments median
Mountain above Salang tunnel, Gibbons 823 (MO); Hindu white, slightly dilated at base,
& mm mm
Kush, Gilbert H8 (E). Parvaii: Panjshir, Hedge Wen- pairs 5-6 long, lateral pair 3-4 long; an-
delbo 5451 (E). Kapisa: Mir Saniir area. Gibson 211 (E). mm
thers narrowly oblong, 1.2—1.5 long. Ovules 25
it^-i-
Takliar; Khost-o-Eereng, valley Echani-Tai, E of Clnju- „r o.ri- ^rkc-i^
^™^
i ^^'^
^^ ^ 2.5-3.0
^^^"^^- ^^^^^'
P^'
duk, Podlech 11832 CHINA. Xinjiang: Kashgaria, *^ ^'"^
((i).
mm,
354 strongly flattened; valves tomentose, distinctly
Tian Slmn, 1889, Koborouski sjh (LE), Merzbachcr
(LE): Kashgaria. Billuli Pass, 13 June 1909, Dirnogor- veined; septum complete, membranous; style ob-