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Notes on Melanoseris lessertiana (Lactucinae, Asteraceae) and Morphologically Allied Species in the PanHimalayan Region

菊科莴苣亚族黑苞毛鳞菊及其泛喜马拉雅地区相近种的分类杂记



全 文 :菊科莴苣亚族黑苞毛鳞菊及其泛喜马拉雅地区相近种的分类杂记
王泽欢1ꎬ Kilian Norbert2ꎬ 彭  华1∗
(1 中国科学院昆明植物研究所东亚植物多样性与生物地理学重点实验室ꎬ 昆明  650201ꎻ
2 Botanic Garden and Botanical Museum Berlin ̄Dahlemꎬ Freie Universität Berlinꎬ Berlinꎬ Germany)
摘要: 基于已有标本的详细研究和文献的追踪ꎬ 证实黑苞毛鳞菊 (或黑苞乳苣) 在中国没有分布ꎬ 国内相
关文献上的该物种实际应为后期发表的青海乳苣ꎬ 现已处理为青海毛鳞菊ꎬ 并首次报道不丹有分布ꎮ 青海
毛鳞菊的瘦果形态和花药管长度明显异于黑苞毛鳞菊ꎮ 作为标本稀有的云南特有种ꎬ 单头乳苣近期被暂时
处理为黑苞毛鳞菊的异名ꎬ 但本研究支持其作为一个独立种且与大花毛鳞菊复合群关系较近ꎮ 对于巴基斯
坦北部和克什米尔地区近期发表的与黑苞毛鳞菊形态相近的两个新种ꎬ Cicerbita alii 被证实为独立种且新
组合到毛鳞菊属ꎬ 而 C􀆰 astorensis被处理为黑苞毛鳞菊的异名ꎮ
关键词: 乳苣属ꎻ 毛鳞菊属ꎻ 岩参属ꎻ 莴苣属ꎻ 分类ꎻ 莴苣亚族ꎻ 菊科
中图分类号: Q 949            文献标志码: A              文章编号: 2095-0845(2015)04-401-06
Notes on Melanoseris lessertiana (Lactucinaeꎬ Asteraceae) and
Morphologically Allied Species in the Pan ̄Himalayan Region
WANG Ze ̄huan1ꎬ KILIAN Norbert2ꎬ PENG Hua1∗
(1 Key Laboratory for Plant Diversity and Biogeography of East Asiaꎬ Kunming Institute of Botanyꎬ Chinese Academy
of Sciencesꎬ Kunming 650201ꎬ Chinaꎻ 2 Botanic Garden and Botanical Museum Berlin ̄Dahlemꎬ
Freie Universität Berlinꎬ Berlinꎬ Germany)
Abstract: Based on detailed study of available specimens and perusal of literatureꎬ it is shown that Melanoseris les ̄
sertiana or Mulgedium lessertianum is not present in China and corresponding reports in the floristic literature of Chi ̄
na actually refer to a species originally described as Mulgedium qinghaicumꎬ and correctly placed as Melanoseris
qinghaicaꎬ a species here reported for the first time also for Bhutan. M􀆰 qinghaica can be clearly distinguished from
M􀆰 lessertiana by achene shape and anther tube length. The rare and little known Yunnan endemicꎬ originally de ̄
scribed as Lactuca monocephala and tentatively considered as conspecific with M􀆰 lessertiana recentlyꎬ is confirmed
as an independent species related to the M􀆰 atropurpurea complex. Of two recently described species from N Pakistan
and Kashmir morphologically allied to M􀆰 lessertianaꎬ Cicerbita alii is confirmed as a separate species and transferred
to Melanoserisꎬ while C􀆰 astorensis is considered as conspecifc with M􀆰 lessertiana.
Key words: Mulgediumꎻ Melanoserisꎻ Cicerbitaꎻ Lactucaꎻ taxonomyꎻ Lactucinaeꎻ Asteraceae
  Mulgedium lessertianum was first described by de
Candolle (1838)ꎬ and was subsequently transferred
by Decaisne (Jacquemontꎬ 1843) to the new genus
Melanoseris Decne.ꎬ but later to Lactuca L. (Clarkeꎬ
1876) and Cicerbita Wallr. ( Mamgain and Raoꎬ
1989). Its distribution area has been considered to
range from N Pakistan and Kashmir ( Bano and
Qaiserꎬ 2010) across the Himalayas (Mamgain and
Raoꎬ 1989ꎬ 1995ꎻ Hara et al.ꎬ 1982ꎻ Grierson and
Longꎬ 2001) to China ( Tangꎬ 1983ꎻ Shihꎬ 1988ꎬ
植 物 分 类 与 资 源 学 报  2015ꎬ 37 (4): 401~406
Plant Diversity and Resources                                    DOI: 10.7677 / ynzwyj201514136
∗ Author for correspondenceꎻ E ̄mail: hpeng@mail􀆰 kib􀆰 ac􀆰 cn
Received date: 2014-10-20ꎬ Accepted date: 2015-01-16
作者简介: 王泽欢 (1986-) 女ꎬ 博士ꎬ 主要从事植物分类与系统发育研究ꎮ E ̄mail: wangzehuan@mail􀆰 kib􀆰 ac􀆰 cn
1997). In the Flora of China (Shih et al.ꎬ 2011)ꎬ
the genus Melanoseris was revivedꎬ and the species a ̄
gain treated as M􀆰 lessertiana. A molecular phyloge ̄
netic analyses by Wang et al. (2013) confirmed that
Melanoseris represents a separate lineage within the
subtribe Lactucinae and revealed its sister group re ̄
lationship to Lactuca. From other species in the ge ̄
nus with capitula of similar flower number ( in the
range of 15 - 25)ꎬ M􀆰 lessertiana in the wide sense
previously applied differs by its low stature (usually
up to 30 cm)ꎬ the usual absence ( or else already
basal branching) of a dominant main stem and pres ̄
ence of several weak secondary flowering stemsꎬ and
a pappus without an outer row of minute hairs.
Since the incision of the leaves of M􀆰 lessertiana
shows strong variabilityꎬ the species has been subdi ̄
vided into three subspecies ( Stewartꎬ 1972ꎻ Dhar
and Kachrooꎬ 1983ꎻ Mamgain and Raoꎬ 1989ꎬ 1995)
or varieties (Bano and Qaiserꎬ 2010). Besidesꎬ sev ̄
eral species morphologically similar to M􀆰 lessertiana
have been described. Mulgedium qinghaicum S􀆰 W.
Liu & T􀆰 N. Ho (Liu and Hoꎬ 2001) from Qinghaiꎬ
Chinaꎬ of which the holotype and only specimen
(S􀆰 W. Liu 3536bꎬ HNWP) is lostꎬ and Lactuca
monocephala C􀆰 C. Chang (Changꎬ 1934) [ =Mulg ̄
edium monocephalum (C􀆰 C. Chang) C. Shih (Shihꎬ
1988)]ꎬ then only known from the type collections
from Yunnan in 1913ꎬ were tentatively treated as
conspecific with Melanoseris lessertiana by Shih and
Kilian (Shih et al.ꎬ 2011). In additionꎬ recently three
new species morphologically allied to M􀆰 lessertiana
were described from N Pakistan and Kashmir (Bano
and Qaiserꎬ 2010)ꎬ i􀆰 e. Cicerbita aliiꎬ C􀆰 astorensis
and C􀆰 gilgitensis.
1  Material and methods
The investigation is based on herbarium materi ̄
al from the herbaria of Bꎬ KUNꎬ MB and PEꎬ and
high resolution digital specimen images of further
type material available through JSTOR Global Plants
(2014)ꎬ or such provided by herbaria. We morpho ̄
logically investigated the three species reported from
Chinaꎬ Mulgedium lessertianumꎬ Lactuca monoceph ̄
ala and M􀆰 qinghaicum as well as Cicerbita alii and
C􀆰 astorensis from Pakistan. Micromorphological fea ̄
tures of herbarium specimens were examined under a
WILD M5 optical reflected ̄light microscope. Fea ̄
tures were documented with an Olympus DP72 digit ̄
al camera mounted on an Olympus SZX16 stereo
zoom optical reflected ̄light microscope equipped with
the Olympus analySIS docu software.
2  Results and discussion
Morphological analyses of the material revealed
that in addition to achene shapeꎬ anther tube length
is a valuable character to distinguish macromorpho ̄
logically else similar taxa.
Melanoseris lessertiana: The type collection of
M􀆰 lessertianum DC. from Kumaon [ a region of E
Uttarakhand]ꎬ India ( Wallich Cat no. 3254 / 364
with ripe achenes e􀆰 g. in the holotype at G ̄DC (G
00469963) and the isotype at K ( K 000250933)
clearly shows that the achenes of this species have a
stout beak of the kind as illustrated for M􀆰 lyrata
Decne. ( Jacquemontꎬ 1843)ꎬ an early synonym of
M􀆰 lessertiana. Our analysis of specimens from the
Himalayas revealed that this shape is characteristic
for M􀆰 lessertiana in the sense of the type ( see also
Fig􀆰 1A1-2) and is correlated with an anther tube of
3􀆰 7-4􀆰 7 mm length (Table 1). The involucre has
some pubescence (Fig􀆰 1A3) of varying degree. We
cannot confirm the statement by Bano and Qaiser
(2010) of the presence of an outer row of minute
pappus hairs in this species.
Mulgedium qinghaicum: Although the type ma ̄
terial is lostꎬ the authors provided an illustrationꎬ
which is quite instructiveꎬ characterizing the species
by a long ̄beaked acheneꎬ a pappus without an outer
row of minute hairs and flowers with an anther tube
(excluding appendages) of approximately only 2 mm
length. Detailed study on the specimens and perusal
of literature ( Tangꎬ 1983ꎻ Shihꎬ 1997ꎻ Liu and
Hoꎬ 2001) revealed that all material known from
China and referred to Melanoseris lessertiana ( or
204                                  植 物 分 类 与 资 源 学 报                            第 37卷
Mulgedium lessertianum) well matches Mulgedium
qinghaicum in the combination of features characteri ̄
zing this species. M􀆰 qinghaicum differs from M􀆰 less ̄
ertiana mainly in achene shape and anther tube
lengthꎬ while it agrees in capitulum size and pubes ̄
cence of the involucre ( Fig􀆰 1B3). The achenes of
M􀆰 qinghaicum have a thin beak which is distinctly
longer than the body (Fig􀆰 1B1-2)ꎬ while M􀆰 lesser ̄
tiana has a much shorter and stouter beak (Fig􀆰 1A1
-2). The anther tube of M􀆰 qinghaicum is only 2􀆰 1
-2􀆰 4 mm longꎬ compared with 3􀆰 7 - 4􀆰 7 mm for
M􀆰 lessertiana (Table 1). Molecular phylogenetic an ̄
alyses (Wang et al.ꎬ 2013) confirmed the inclusion
of the species in the Melanoseris lineage but are not
conclusive with respect to its relationshipꎬ because
nuclear and chloroplast phylogenies show incongru ̄
ent placements within Melanoseris. The species is
here reported for the first time also for Bhutanꎬ based
Fig􀆰 1  Achenesꎬ pappus and capitula of Melanoseris lessertiana and three morphologically allied species
A: Melanoseris lessertiana ( =Mulgedium lessertianum)ꎻ B: Mulgedium qinghaicumꎻ C: Cicerbita aliiꎻ D: Cicerbita astorensis. ∗1: achene
shape (bars at 2 mm)ꎻ 2: details of pappus base (bars at 500 μm)ꎻ 3: capitula (bars at 5 mm). A from G. Watt 2172 (B)ꎻ
B from G. & S. Miehe 00 ̄346 ̄01 (MB)ꎻ C from O. Polunin 56 / 567 (B)ꎻ D from O. Polunin 56 / 717 (B)
3044期      WANG Ze ̄huan et al.: Notes on Melanoseris lessertiana (Lactucinaeꎬ Asteraceae) and Morphologically 􀆺     
on the collection by G. & S. Miehe 00 ̄346 ̄01 at
Marburg (MB): “Bhutanꎬ Floristic Province N 19
( Upper Pho Chu)ꎬ District Gasaꎬ Gecha La W
sideꎬ above Wocheꎬ 28°1′Nꎬ 90°1′Eꎬ 4 200-4 300 mꎬ
Rhododendron anthropogon dwarf ̄scrub pastures on
SW ̄facing slopeꎬ 1 Sep 2000” ).
Lactuca monocephala: While the protologue and
the holotype in Kew give no collecting data other
than just “Yunnan”ꎬ the label of the isotypes at Edin ̄
burgh (E) and Beijing ( PE) are much more in ̄
formativeꎬ reading “ Plant of 6 - 9 inchesꎬ flowers
pale blueꎬ on cliffs and humus ̄covered boulders on
the mountains in the N. E. of the Yangtze bendꎬ Lat.
27°50′ Nꎬ Alt. 10. ̄11. 000 ft.ꎬ Sept. 1913”. The
type collection bears no ripe achenes and the anther
tube has a similar length as M􀆰 lessertiana (see Table
1). The species was rediscovered for the first time
by Niu Yang in NW Yunnan in 2010 and the molec ̄
ular data ( generated by Jiangwen Zhangꎬ Kunming
Institute of Botany) confirm its position in Melano ̄
seris indicating a relationship with the M􀆰 atropurpurea
complex (unpublished data). The specimenꎬ collected
in flowerꎬ subsequently got lost. Photographs of the
plant are shown in the Cichorieae Portal (ICN Portal
2014 under M􀆰 monocephala).
Cicerbita alii: The isotype at Berlin reveals that
the unusually thickꎬ short stem and its achenes with
the distinctively shouldered body below the beak
(Fig􀆰 1C1) and a pappusꎬ in contrast to the state ̄
ment by Bano and Qaiser (2010)ꎬ without an outer
row of minute hairs ( Fig􀆰 1C2) are correlated with
the possession of the longest anther tube (of c. 6 mm)
so far known in Melanoseris (Table 1). This feature
combination leaves no doubt at its independent spe ̄
cies status. The involucreꎬ at least of the type collec ̄
tionꎬ shows a particularly dense and long indumen ̄
tum (Fig􀆰 1C3). Morphologicallyꎬ the species well
matches the Melanoseris clade and should be treated
as a member of that genus. The species seems to have
a restricted distribution in N Pakistan and Kashmir.
Cicerbita astorensis: The paratype material at
Berlin cannot reliably be distinguished from M􀆰 lesser ̄
tianaꎬ neither with respect to achene shape (Fig􀆰 1
D1-2) nor to anther tube length (Table 1)ꎬ nor to
any other feature investigatedꎬ including the habitual
difference given by its authors. We cannot confirm
the statement by Bano and Qaiser ( 2010) of the
presence of an outer row of minute pappus hairs.
Therefore this species should be treated as conspecif ̄
ic with M􀆰 lessertiana.
As no specimen of Cicerbita gilgitensis was avai ̄
lableꎬ this species is not considered here.
Table 1  Comparison of the anther tube length of Melanoseris lessertiana and morphologically allied species
Species name Voucher (herbarium) Collection location
Anther tube length
(excluding appendages)
/ mm
Mulgedium lessertiana G. Watt. 2172 (B) Chumba Stateꎬ India 3􀆰 7-4􀆰 0
Mulgedium lessertiana M. Nüsser 157 (B) N Pakistan 4􀆰 0-4􀆰 3
Mulgedium lessertiana J. H. Lace 1657 (B) Chambaꎬ India 4􀆰 5-4􀆰 7
Mulgedium lessertiana var. lyrata O. Polunin 56 / 192 (B) Khelanmargꎬ Kashmir 4􀆰 2-4􀆰 4
Cicerbita astorensis O. Polunin 56 / 717 (B) Mengandobꎬ Kashmir 3􀆰 9-4􀆰 1
Mulgedium qinghaicum G. & S. Miehe 00 ̄346 ̄01 (MB) Upper Pho Chuꎬ Bhutan 2􀆰 1-2􀆰 2
Mulgedium qinghaicum G. & S. Miehe 05 ̄047 ̄06 (MB) Xizangꎬ China 2􀆰 3-2􀆰 4
Mulgedium qinghaicum G. & S. Miehe 05 ̄086 ̄06 (MB) Xizangꎬ China 2􀆰 1   
Lactuca monocephala G. Forrest 11231 (E) Yunnanꎬ China 4􀆰 1-4􀆰 4
Cicerbita alii O. Polunin 56 / 567 (B) Kolahoiꎬ Kashimir 5􀆰 9-6􀆰 1
404                                  植 物 分 类 与 资 源 学 报                            第 37卷
Taxonomic treatment
Key to the four morphologically allied species of Melanoseris
1a. Perennial rosette herbs with subscapiform only basally leafyꎬ unbranched flowering stemsꎬ involucre glabrousꎬ anther tube ex ̄
cluding appendages 4􀆰 1-4􀆰 4 mm long Melanoseris monocephala􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺
1b. Perennial herbs with branched and ± leafy flowering stemsꎬ synflorescence racemiformꎬ involucre pubescent to varying
degrees
2a. Achenes with a distinctively shouldered body below the black beakꎬ anther tube excluding appendages c. 6 mm long
Melanoseris alii􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺
2b. Achene body attenuate to weakly contracted into a pale beakꎬ anther tube excluding appendages less than 5 mm long
3a. Achenes with a stout beakꎬ anther tube excluding appendages 3􀆰 7-4􀆰 7 mm long Melnoseris lessertiana􀆺􀆺􀆺􀆺
3b. Achenes with a thin beak distinctively longer than the bodyꎬ anther tube excluding appendages 2􀆰 1-2􀆰 4 mm long
Melanoseris qinghaica􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺􀆺
    Melanoseris lessertiana (DC.) Decne. in Jac ̄
quemontꎬ Voy. Inde 4: 102. 1843ꎻ Mulgedium les ̄
sertianum DC.ꎬ Prodr. 7: 251. 1838ꎻ Lactuca lesser ̄
tiana (DC.) C􀆰 B. Clarkeꎬ Compos. Ind.: 270. 1876ꎻ
Cicerbita lessertiana ( DC.) Mamgain & Rao in J.
Bombay Nat. Hist. Soc. 86: 273. 1989. -Holotype:
[India] “montanis ad Kamaon et Emodum”ꎬ Walli ̄
ch s􀆰 n.ꎬ cat. comp. [3254 / ]364 (G ̄DC [mounted
on two sheetsꎬ G 00469962ꎬ G 00469963ꎻ isotypes:
K 000250933ꎬ NY 00230698).
=Cicerbita astorensis Bano & Qaiser in Pakistan J.
Bot. 42: 39. 2010. -Holotype: Pakistanꎬ Astor Dist.ꎬ
Shaban top above Dombabhoꎬ 11􀆰 9􀆰 2006ꎬ Ali Noor
& al. 584 (KUH).
Notes: (1) We consider the two sheets preserved
in the Candolle herbarium at Genève (G ̄DC) as a
single specimen according to Art 8􀆰 3 of the Mel ̄
bourne Code (McNeill et al.ꎬ 2012): they are num ̄
bered (by the curators?) “1” ( =G 00469962) and
“2” ( =G 00469963)ꎻ “1” carries the original label
while “2” only has the pencil number “364”ꎻ only
“2” has the label with the revised determinationꎬ a
paper capsule with loose fruiting heads and achenesꎬ
and carries the statement by Auguste Monnier men ̄
tioned in the protologue “Certé non Hieracium / Genus
novum?”ꎬ all being indications that already Candolle
considered them as a single entity. (2) For further syn ̄
onyms see ICN Portal (ICN (Hand et al.)ꎬ 2009+).
Distribution: Melanoseris lessertiana in the sense
of its type is distributed in the Himalayas from N Paki ̄
stan (Baltistanꎬ Gilgit) and Kashmir (Bano and Qais ̄
er 2010) across NE India (Jammu ̄Kashmirꎬ Himachal
Pradeshꎬ Uttarakhandꎬ Sikkim) (Hajra et al.ꎬ 1995)
and Nepal (Hara et al.ꎬ 1982)ꎬ but apparently does
not enter China. Its presence in Bhutan (Grierson and
Springateꎬ 2001) needs confirmation for possible
confusion with M􀆰 qinhaica.
Melanoseris qinghaica (S􀆰 W. Liu & T􀆰 N. Ho)
N. Kilian & Ze H. Wang in PLoS One 8 ( 12)ꎬ
e82692: 18. 2013ꎻ Mulgedium qinghaicum S􀆰 W. Liu
& T􀆰 N. Ho in Acta Phytotax. Sin. 39: 556. 2001. -
Holotype: “China. Qinghai: Hualongꎬ in poplar for ̄
estsꎬ alt. 2 600 m”ꎬ 18 Sep 1988ꎬ S. W. Liu 3536b
(HNWPꎻ lost)ꎻ lectotype (designated here): [i ̄
con] Acta Phytotax. Sin. 39: 557ꎬ fig. 2. 2001.
Distribution: To our current knowledgeꎬ M􀆰 qing ̄
haica is restricted in its distribution to China (Xi ̄
zangꎬ Qinghai and Yunnan) and Bhutan.
Melanoseris monocephala (C􀆰 C. Chang) Ze
H. Wang in PLoS One 8 (12)ꎬ e82692: 18. 2013ꎻ
Lactuca monocephala C􀆰 C. Chang in Contr. Biol.
Lab. Sci. Soc. Chinaꎬ Bot. Ser. 9: 132. 1934ꎻ Mulged ̄
ium monocephalum (C􀆰 C. Chang) C. Shih in Acta
Phytotax. Sin. 26: 391. 1988. -Holotype: Chinaꎬ
Yunnanꎬ [mountains in the N. E. of the Yangtse
bendꎬ 27° 50′ Nꎬ 10. ̄11. 000 ft.ꎬ Sep 1913]ꎬ G.
Forrest 11231 (K 000808102ꎻ isotypes: E 00394949ꎬ
PE 00552631).
5044期      WANG Ze ̄huan et al.: Notes on Melanoseris lessertiana (Lactucinaeꎬ Asteraceae) and Morphologically 􀆺     
Note: Locality data from isotypes at E & PE.
Distribution: This rare species is probably en ̄
demic to NE Yunnan.
Melanoseris alii (Bano & Qaiser) N. Kilian &
Ze H. Wangꎬ comb. nov.ꎻ Cicerbita alii Bano &
Qaiser in Pakistan J. Bot. 42: 39. 2010. -Holotype:
Pakistanꎬ Kashmirꎬ Kolohoi valleyꎬ 27􀆰 8􀆰 1956ꎬ O.
Polunin 56 / 567 (BMꎻ isotype B).
Distribution: The species is known only from
N Pakistan (Gilgit) and Kashmir.
Acknowledgements: The authors are grateful to Sabine &
Georg Miehe (Marburg) who provided material of their col ̄
lections for our studies and to the curators of the herbaria G
(Genèveꎬ Switzerland) and PE (Beijingꎬ China) for their
kind help. We also thank Julia Pfitzner (Berlinꎬ Germany)
for the photographs of Fig􀆰 1. The first author thanks the Ger ̄
man Academic Exchange Service ( Deutscher Akademischer
Austausch Dienstꎬ DAAD) for providing a grant for a four
month research stay in 2012 in Berlin.
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604                                  植 物 分 类 与 资 源 学 报                            第 37卷