全 文 ：书中国唇形科草糙苏属分类学研究杂记
*
向春雷1，董洪进1，胡国雄1，2，周 静3，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，云南 昆明 650201;2 中国科学院大学，
北京 100049;3 昆明医科大学药学院暨云南省天然药物药理重点实验室，云南 昆明 650500)
摘要:最近的研究结果表明，国产糙苏属 (Phlomis)的所有种类 (除了栽培种橙花糙苏 P. fruticosa以外)
以及沙穗属 (Eremostachys)、独一味属 (Lamiphlomis)、钩萼草属 (Notochaete)均应并入草糙苏属 (Phlo-
moides)。新界定的草糙苏属因其草本习性等特征而与糙苏属易于区别。尽管已有不同作者对国产草糙苏
属大部分物种名称进行了新组合，但仍遗漏了 11个物种名称。为方便使用，本文对这 11个名称进行了新
组合，同时提供了新界定的草糙苏属国产物种名录及检索表，共 50种 9变种。
关键词:糙苏属;糙苏族;草糙苏属;唇形科;新组合
中图分类号:Q 949 文献标识码:A 文章编号:2095－0845(2014)05－551－10
Taxonomic Notes on the Genus Phlomoides
(Lamiaceae:Lamioideae)from China
XIANG Chun-Lei1，DONG Hong-Jin1，HU Guo-Xiong1，2，ZHOU Jing3，PENG Hua1＊＊
(1 Key Laboratory for Plant Diversity and Biogeography of East Asia，Kunming Institute of Botany，Chinese Academy
of Sciences，Kunming 650201，China;2 University of Chinese Academy of Sciences，Beijing 100049，China;
3 School of Pharmaceutical Science ＆ Yunnan Key Laboratory of Pharmacology for Nature Products，
Kunming Medical University，Kunming 650500，China)
Abstract:Ｒecent phylogenetic studies have investigated the tribe Phlomideae and，in particular，the degree of“in-
clusiveness”of the genus Phlomoides． As a result，Eremostachys，Lamiophlomis，Notochaete and all Chinese Phlo-
mis，excepted for cultivated P. fruticosa，were transferred to the latest redefined Phlomoides． The newly circum-
scribed Phlomoides is easily distinguished from Phlomis by its herbaceous habit and several other traits． Although the
necessary new combinations for the Chinese species have been made elsewhere，unfortunately，eleven names were o-
mitted by previous researchers． Therefore，this paper proposes 11 new nomenclatural combinations in Phlomoides，
and provides a checklist and a key to all Chinese Phlomoides．
Key words:Phlomis;Tribe Phlomideae;Phlomoides;Lamiaceae;New combinations
The genus Phlomoides Moench (Phlomideae，
Lamioideae，Lamiaceae) is the species-rich genus
with about 150－170 species，extending from central
Europe to the Ｒussian Far East (Mathiesen et al．，
2011;Salmaki et al．，2012a，b)． The genus was
separated from Phlomis L． by Moench (1794)who
placed Phlomis tuberosa L． in his monotypic genus
Phlomoides． However，in the past 200 years，most
taxonomists (Bentham，1832－1836;Boissier，1879;
Briquet，1895－1897;Wu and Li，1977)had treated
植 物 分 类 与 资 源 学 报 2014，36 (5):551～560
Plant Diversity and Ｒesources DOI:10．7677 /ynzwyj201413228
＊
＊＊
Funding:The National Natural Science Foundation of China(31110103911，31100164) ;Knowledge Innovation Project of the Chinese Acade-
my of Sciences (KSCX2-EW-Z-1) ;the Ministry of Science and Technology China，Basic Ｒesearch Project (2013FY112100)
Author for correspondence;E-mail:hpeng@mail. kib. ac. cn
Ｒeceived date:2013－11－25，Accepted date:2014－07－14
作者简介:向春雷 (1984－)男，博士，副研究员，主要从事植物分类与系统发育研究。E-mail:xiangchunlei@mail. kib. ac. cn
Phlomoides as one of two sections within Phlomis un-
til it was reinstated as a genus (Adylov et al．，
1986;Adylov and Machmedov，1987;Kamelin and
Machmedov，1990a，b)． The reinstated Phlomoides
includes some species with the upper corolla lip
not laterally compressed and with tuberous lateral
roots (Salmaki，2012a)． Meanwhile，some other
genera such as Eremostachys Bunge， Notochaete
Benth．，Paraeremostachys Adylov，Kamelin ＆ Ma-
khm．，Lamiophlomis Kud were established or sepa-
rated from Phlomis (Bentham，1829;Bunge，1830;
Kud，1929;Adylov et al．，1986) ，and all of them
were considered closely related to Phlomis． Based
on pericarp structure，Ｒyding (2008)agreed that
Phlomoides should be treated as a separate genus，
including the species from Lamiophlomis and No-
tochaete，but he didn’t make normal taxonomical
treatments．
Pan et al． (2009)supported the inclusion of
Lamiophlomis in Phlomis based on their molecular
phylogenetic study，however only fourteen species of
Phlomis were included in their study． Ｒecently，u-
sing DNA sequences of the plastid trnL intron，trnL-
trnF spacer and rps16 intron，Phlomis s. l． were split
into two lineages， i. e． “Phlomoides group” and
“Phlomis group”，and therefore Mathiesen et al．
(2011)recognized two genera，i. e．，Phlomis s. str．
and Phlomoides s. l．，and the latter included Pseud-
eremostachys，Lamiophlomis as well as Notochaete
hamosa Benth． In the following molecular phyloge-
netic study on the tribe Phlomideae，Salmaki et al．
(2012b)confirmed the inclusion of the members of
Eremostachys，and other mono- or oligotypic genera
(e. g． Lamiophlomis，Notochaete，Paraeremostachys)
in Phlomoides，and some necessary new combina-
tions are proposed therein．
Now，circumscription for Phlomoides has been
clear，and the relationships between Phlomoides and
Phlomis were resolved by molecular data and suppor-
ted by several morphological characters (Table 1)as
well as chromosome numbers (Azizian and Culter，
1982;Astanova，1984;Brullo et al．，1990;Ghaf-
fari，2006，Fang et al．，2007)． According to the
current circumscription of Phlomoides，there are for-
ty-nine species and ten varieties in China;of which
forty-two species and nine varieties were previously
in Phlomis，four species and one variety were previ-
ously in Eremostachys，two were previously in No-
tochaeta and one species previously in Lamiophlomis．
Although the necessary new combinations for most
Chinese species have been made elsewhere (e. g．
Adylov and Makhmedov，1987;Matheisen et al．，
2011;Salmaki et al．，2012b) ，eleven recognized
taxa are omitted by these authors． In order to aid fu-
ture systematic work on these taxa，here we propose
eleven new combinations，and for the first time，we
provide a checklist and a key to all Chinese Phlo-
moides species in its new definition．
Table 1 Characteristics of Phlomoides and Phlomis (see also Salmaki et al． 2012b)
Phlomoides Phlomis
Numbers of species in China 49 1
Growth form
Tall perennial herbs with woody rhizomes and /or
tubers at tip
Perennial herbs or small shrubs
Petioles 2－5 (－7)cm 1. 5－5 cm
Calyx lobe
equal or occasionally 3 /2 bilabiate，abruptly narrow
to acute apex
equal
Posterior lip of corolla long，shallowly hooded long，hooded，often deeply concave
Indumentum on upper corolla lip Bearded with simple hairs at margin Hardly bearded with simple hairs at margin
Nutlet apex Stellate hairy or sometimes glabrous Glabrous，papillose or stellate hairy
Chromosome number 2n= 22 2n= 12，14，20，22，40，42
255 植 物 分 类 与 资 源 学 报 第 36卷
1 New combinations
Phlomoides cuneata (C. Y． Wu)C. L． Xiang
＆ H． Peng，comb． nov．
Basionym:Phlomis cuneata C. Y． Wu，Fl． Ｒei-
publ． Popularis Sin． 65 (2) :434，594 (1977)．
TYPE:China，Xizang:Cuijila Shan，6 Sep． 1952，
P. C． Tsoong 5335 (PE)．
Distribution:Endemic to Xizang (Tibet) ，and
currently known only from the type locality (Cuijila
Shan)．
Note:The style of this species is equally 2-
cleft at apex，which is very rare in genus Phlomoides
from China．
Phlomoides dentosa (Franch．)Kamelin ＆ Ma-
khm． var． glabrscens (Danguy)C. L． Xiang ＆ H．
Peng，comb． nov．
Basionym:Phlomis dentosa Franch． var． glabr-
scens Danguy，Bull． Mus． Natl． Hist． Nat． 17 (5) :
345 (1911)． TYPE:China，Gansu:Si-Ning-Fou，
2 400 m，18th Jul． 1908，Anonymous s. n． (P)．
Distribution:Endemic to China，known from
Gansu，Hebei，Inner Mongolia，and Qinghai．
Note:Morphological characters distinguished
this variety from Phlomoides dentosa var． dentosa are
that stems sparsely minutely hispid to subglabrous，
leave subglabrous，bracts and calyx sparsely bristly
and dusty puberulent (vs． plants stellate hairy
throughout in variety dentosa)．
Phlomoides mongolica (Turcz．)Kamelin ＆ Ma-
khm． var． macrocephala (C. Y． Wu)C. L． Xiang ＆
H． Peng，comb． nov．
Basionym:Phlomis mongolica Turcz． var． ma-
crocephala C. Y． Wu，Fl． Ｒeipubl． Popularis Sin． 65
(2) :444，595 (1977)． TYPE:China，Inner Mon-
golia:precisely locality unknown，Ju Ud League
s. n． (PE)．
Distribution:Endemic to Inner Mongolia．
Note:It’s easy to distinguish this variety from
variety mongolica． Plants of this variety are ca． 15
cm tall，verticillasters solitary，terminal or axillary，
and sometimes inserted at base of stem． In contrast，
plants of Phlomoides mongolica var． mongolica are
40－ 70 cm tall，verticillasters numerous，axillary，
inserted on apical part of stems．
Phlomoides oreophila (Kar． ＆ Kir．)Adylov．
Kamelin ＆ Makhm． var． evillosa (C. Y． Wu)C. L．
Xiang ＆ H． Peng，comb． nov．
Basionym:Phlomis oreophila Kar． ＆ Kir． var．
evillosa C. Y． Wu，Fl． Ｒeipubl． Popularis Sin． 65
(2) :437，594 (1977)． TYPE:China，Xinjiang:
Fuhai，2 100 m，11 Jul． 1957，C. L． Chen 186 (PE)．
Distribution:Endemic to Xinjiang．
Note:This variety with stellate pannose bracts
and calyx is easily distinguished from variety oreophi-
la because in Phlomoides oreophila var． oreophila，
bracts densely villous or sometime intermixed with
glandular villous hairs，and calyx stellate puberu-
lent，finely villous on veins outside．
Phlomoides speciosa (Ｒupr．)Adylov，Kame-
lin ＆ Makhm． var． viridifolia (Popov)C. L． Xiang
＆ H． Peng，comb． nov．
Basionym:Eremostachys speciosa Ｒupr． var．
viridifolia Popov，Nov． Mem． Moskovsk． Obsc． Isp．
Prir． 19:100 (1940)． SYNETYPES:Kazakhstan，
Semirechenskaya Ｒegion:30th Apr． 1916，Goro-
detskiy 158 (TASH) ;Uzbekistan，Fergana Ｒegion:
Andizhan District，on the way to Kugart Pass，2
Jun． 1911，Knorring ＆ Minkwitz 266 (TASH) ;Sa-
markand Ｒegion:Zimmarkh，16th Jul． 1915，Bala-
baye 9 (TASH)．
Distribution:This variety is recorded from Xin-
jiang．
Note:Variety viridifolia is distributed to Ka-
zakhstan，Kyrgyzstan，Tajikistan，Turkmenistna，and
SW Asia． In addition，because all the Chinese Ere-
mostachys species were transferred to Phlomoides，so
there is no Eremostachys in China henceforth．
Phlomoides tatsienensis (Bureau ＆ Franch．)
Kamelin ＆ Makhm． var． hirticalyx (Hand．-Mazz．)
3555期 XIANG Chun-Lei et al．:Taxonomic Notes on the Genus Phlomoides (Lamiaceae:Lamioideae)from China
C. L． Xiang ＆ H． Peng，comb． nov．
Basionym:Phlomis franchetiana Diels var． hir-
ticalyx Hand．-Mazz． Symb． Sin． 7 (4) :921 (1936)．
TYPE:China，Yunnan:Beyenjing，in silvis，25
Sept． 1919，S． Ten 1380 (WU)．
Distribution:Endemic to Yunnan．
Note:This variety can be distinguished from
variety tatsienensis by its densely stellate pubescent
bracts and calyx (vs． bracts and calyx are stellate
pubescent)．
Phlomoides tibetica (C． Marquand ＆ Airy Sh-
aw)Kamelin ＆ Makhm． var． wardii (C． Marquand
＆ Airy Shaw)C. L． Xiang ＆ H． Peng，comb． nov．
Basionym:Phlomis tibetica C． Marquand ＆
Airy Shaw var． wardii C． Marquand ＆ Airy Shaw，J．
Linn． Soc．，Bot． 48 (321):218－219 (1929)． TYPE:
China，Xizang:on the edges of cultivated fields，Tum-
batse，3 400 m，14 Jul． 1924，F. K． Ward 5937 (K)．
Distribution:Endemic to Xizang．
Note:This variety is distinguished from variety
tibetica by several traits，i. e． leaf blade cordate to
broadly oblong，margin coarsely crenate (vs． leaf
blade ovate-cordate，margin crenate or coarsely cre-
nate in variety tibetica) ;upper corolla lip densely
villous outside (vs． upper corolla lip stellate tomen-
tose outside)．
Phlomoides umbrosa (Turcz．)Kamelin ＆ Mak-
hm． var． australis (Hemsl．)C. L． Xiang ＆ H． Peng，
comb． nov．
Basionym:Phlomis umbrosa Turcz． var． austra-
lis Hemsl．，J． Linn． Soc．，Bot． 26(175):306(1890)．
Distribution:This variety is recorded from An-
hui，Gansu，Guizhou，Hubei，Hunan，Shanxi，Si-
chuan，and Yunnan． SYNTYPES:China，Hubei:
Yichang，Badong，A． Henry 875，1872，2442，
4749，6029，6179，6423，7360 (K)．
Note:There are 4 varieties under Phlomoides
umbrosa． In this species complex，variety australis is
most similar to variety umbrosa，but differentiated by
following traits． In variety australis，leaves long peti-
olate，leaf blade membranous，margin crenate-ser-
rate， terminal tooth sometimes very long; bracts
soft，linear-lanceolate，slightly shorter than calyx，
while in variety umbrosa，leaves papery，serrate-
dentate or irregularly crenate，terminal tooth not very
long;bracts rigid， linear-subulate，mostly longer
than calyx．
Phlomoides umbrosa (Turcz．)Kamelin ＆ Mak-
hm． var． latibrateata (Y. Z． Sun)C. L． Xiang ＆ H．
Peng，comb． nov．
Basionym:Phlomis umbrosa Turcz． var． lati-
brateata Y. Z． Sun，Acta Phytotax． Sin． 11 (1) :46
(1966)． TYPE:China，Henan:Jigong Shan，hill-
sides，700－2 000 m，27 Jul． 1925，A． N． Steward
1584 (NAS)．
Distribution:Endemic to Henan．
Note:In Phlomoides umbrosa complex， this
variety is easily distinguished from other taxa be-
cause its bracts are linear-oblong to obovate-oblong，
and much shorter than calyx，only 5 －7×1. 8 －2. 5
cm，while in other taxa，bracts are linear-subulate
to linear-lanceolate，longer or rarely slightly shorter
than calyx，about 8－14×1－2 mm．
Phlomoides umbrosa (Turcz．)Kamelin ＆ Mak-
hm． var． ovalifolia (C. Y． Wu)C. L． Xiang ＆ H．
Peng，comb． nov．
Basionym:Phlomis umbrosa Turcz． var． ovali-
folia C. Y． Wu，Fl． Ｒeipubl． Popularis Sin． 65 (2) :
478，601 (1977)． TYPE:China，Anhui:Chuxian，
hillsides，2 Aug． 1951，East-China Station 3132
(PE)．
Distribution:This variety is recorded from An-
hui and Jiangsu．
Note:In Phlomoides umbrosa complex， this
variety is similar to variety australis and type varie-
ty，but differentiated by leaves ovate，subcuneate to
shallowly cordate，stellate pubescent，rarely abaxial-
ly stellate tomentulose;calyx densely stellate pubes-
cent outside (vs． leaves circular-ovate to ovate-ob-
long，sparsely pilose or sometimes abaxially densely
455 植 物 分 类 与 资 源 学 报 第 36卷
pilose and stellate pilose;calyx stellate puberulent
except sometime pilose on veins)．
Phlomoides umbrosa Turcz． var． stenocalyx
(Diels)C. L． Xiang ＆ H． Peng，comb． nov．
Basionym:Phlomis stenocalyx Diels，Bot． Jahrb．
Syst． 29 (3-4) :555 (1900)． TYPE:China，Giral-
dii 1548 (K)．
Distribution:This variety is recorded from Gansu
and Shaanxi．
Note:This variety is easily distinguished from
other taxa by its verticillasters conspicuously peduncu-
late，lax，and flowers pedicellate，calyx ca． 8 × 3 cm．
2． Checklist of Phlomoides Moench from
China
Now，there is only one species of Phlomis，i. e．
P. fruticosa L． cultivated in China (Shaanxi Prov-
ince)． According to the new definition of Phlo-
moides，there are 50 species and 9 varieties in Chi-
na;of which 40 species and 9 varieties are endemic
and marked with asterisk (* )in following list，and
distribution area listed in square bracket．
Phlomoides agraria (Bunge)Adylov，Kame-
lin ＆ Makhm． 耕地草糙苏 (新拟) ［Xinjiang;Ka-
zakhstan，Mongolia，Ｒussia］
Phlomoides alpina (Pall．)Adylov． Kamelin ＆
Makhm． 高山草糙苏 (新拟) ［Xinjiang;Kazakh-
stan，Ｒussia］
Phlomoides ambigua (Popov ex Pazij ＆ Vved．)
Adylov，Kamelin ＆ Makhm．* 沧江草糙苏 (新拟)
［Yunnan］
Phlomoides atropurpurea(Dunn．)Adylov． Kam-
elin ＆ Makhm．* 深裂草糙苏 (新拟) ［Yunnan］
Phlomoides betonicoides (Diels)Adylov． Kame-
lin ＆ Makhm．* 假秦艽［Sichuan，Xizang，Yunnan］
Phlomoides chinghoensis (C. Y． Wu)Kame-
lin ＆ Makhm．* 清河草糙苏 (新拟) ［Xinjiang］
Phlomoides congesta (C. Y． Wu)Kamelin ＆
Makhm．* 乾精菜［Sichuan，Yunnan］
Phlomoides cuneata (C. Y． Wu)C. L． Xiang
＆ H． Peng* 楔叶草糙苏 (新拟) ［Xizang］
Phlomoides dentosa (Franch．)Kamelin ＆ Mak-
hm． * 尖齿草糙苏 (新拟)
Phlomoides dentosa var． dentosa* 尖齿草糙苏
原变种［Gansu，Hebei，Inner Mongolia，Qinghai］
Phlomoides dentosa var． glabrescens (Danguy)
C. L． Xiang ＆ H． Peng* 渐光尖齿草糙苏 (新拟)
［Sichuan，Xizang］
Phlomoides desertorum (P. A． Smirn．)Ma-
vrodiev ＆ Sukhor．* 沙生沙穗［Xinjiang］
Phlomoides fimbriata (C. Y． Wu)Kamelin ＆
Makhm．* 裂唇草糙苏 (新拟) ［Yunnan］
Phlomoides forrestii (Diels)Kamelin ＆ Ma-
khm．* 苍山草糙苏 (新拟) ［Yunnan］
Phlomoides franchetiana (Diels)Kamelin ＆
Makhm．* 大理草糙苏 (新拟) ［Yunnan］
Phlomoides fulgens (Bunge)Adylov，Kamelin
＆ Makhm． 光沙穗［Xinjiang;Kyrgyzstan，SW Asia］
Phlomoides hamosa (Benth．)Mathiesen 钩萼
草［Yunnan;Bhutan，India;Myanmar，Nepal］
Phlomoides inaequalisepala (C. Y． Wu)Ka-
melin ＆ Makhm．* 斜萼草糙苏 (新拟) ［Sichuan］
Phlomoides jeholensis (Nakai ＆ Kitag．)Ka-
melin ＆ Makhm．* 口外草糙苏 (新拟) ［Hebei］
Phlomoides kansuensis (C. Y． Wu)Kamelin
＆ Makhm．* 甘肃草糙苏 (新拟) ［Gansu］
Phlomoides koraiensis (Nakai) Kamelin ＆
Makhm． 长白草糙苏 (新拟) ［Jilin;Korea］
Phlomoides likiangensis (C. Y． Wu)Kamelin
＆ Makhm．* 丽江草糙苏 (新拟) ［Yunnan］
Phlomoides longiaristata (C. Y． Wu ＆ H. W．
Li)Salmaki* 长刺钩萼草 ［Xizang，Yunnan］
Phlomoides longicalyx (C. Y． Wu)Kamelin
＆ Makhm．* 长萼草糙苏 (新拟) ［Yunnan］
Phlomoides maximowiczii (Ｒegel)Kamelin ＆
Makhm．* 大叶草糙苏 (新拟) ［Hebei，Jilin，Lia-
oning］
Phlomoides medicinalis (Diels)Kamelin ＆
Makhm．* 萝卜秦艽［Sichuan，Xizang］
Phlomoides megalantha (Diels)Kamelin ＆
Makhm．* 大花草糙苏 (新拟) ［Hubei，Shaanxi，
Shanxi，Sichuan］
Phlomoides melanantha (Diels)Kamelin ＆
5555期 XIANG Chun-Lei et al．:Taxonomic Notes on the Genus Phlomoides (Lamiaceae:Lamioideae)from China
Makhm．* 黑毛草糙苏 (新拟) ［Sichuan，Yunnan］
Phlomoides milingensis (C. Y． Wu ＆ H. W．
Li)Kamelin ＆ Makhm．* 米林草糙苏 (新拟) ［Xi-
zang］
Phlomoides molucelloides (Bunge)Salmaki 沙
穗［Xinjiang;Kazakhstan，Kyrgyzstan，Mongolia，
Ｒussia，Tajikistan;SW Asia，Europe］
Phlomoides mongolica (Turcz．)Kamelin ＆
Makhm． * 串铃草
Phlomoides mongolica var． mongolica* 串铃
草原变种［Gansu，Hebei，Inner Mongolia，Shaanxi，
Shanxi］
Phlomoides mongolica var． macrocephala (C.
Y． Wu)C. L． Xiang ＆ H． Peng* 大头串铃草 (新
拟) ［Inner Mongolia］
Phlomoides muliensis (C. Y． Wu)Kamelin ＆
Makhm．* 木里草糙苏 (新拟) ［Sichuan］
Phlomoides multifurcata Salmaki* 糙苏沙穗
［Xinjiang］
Phlomoides oreophila (Kar． ＆ Kir．)Adylov．
Kamelin ＆ Makhm． 山地草糙苏 (新拟)
Phlomoides oreophila var． oreophila 山地草
糙苏原变种 ［Xinjiang;Kazakhstan，Kyrgyzstan，
Mongolia，Ｒussia，Tajikistan］
Phlomoides oreophila var． evillosa (C. Y． Wu)
C. L． Xiang ＆ H． Peng* 山地草糙苏无长毛变种
［Xinjiang］
Phlomoides ornata (C. Y． Wu)Kamelin ＆
Makhm．* 美观草糙苏 (新拟) ［Sichuan，Yunnan］
Phlomoides paohsingensis (C. Y． Wu)Kame-
lin ＆ Makhm．* 宝兴草糙苏 (新拟) ［Sichuan］
Phlomoides pararotata (Y. Z． Sun)Kamelin
＆ Makhm．* 假轮状草糙苏 (新拟) ［Yunnan］
Phlomoides pedunculata (Y. Z． Sun)Kamelin
＆ Makhm．* 具梗草糙苏 (新拟) ［Sichuan］
Phlomoides pratensis (Kar． ＆ Kir．)Adylov．
Kamelin ＆ Makhm． 草原草糙苏 (新拟) ［Xin-
jiang;Kazakhstan，Kyrgyzstan］
Phlomoides pygmaea (C. Y． Wu)Kamelin ＆
Makhm．* 矮草糙苏 (新拟) ［Xizang］
Phlomoides rotata (Benth． ex Hook． f．)Math-
iesen 独一味［Gansu，Qinghai，Sichuan，Xizang，
Yunnan;Bhutan，India，Nepal］
Phlomoides ruptilis (C. Y． Wu)Kamelin ＆
Makhm．* 裂萼草糙苏 (新拟) ［Yunnan］
Phlomoides setifera (Bureau ＆ Franch．)Ka-
melin ＆ Makhm．* 刺毛草糙苏 (新拟) ［Sichuan，
Xizang，Yunnan］
Phlomoides speciosa (Ｒupr．)Adylov，Kame-
lin ＆ Makhm． var． viridifolia (Popov)C. L． Xiang
＆ H． Peng* 绿叶美丽沙穗［Xinjiang］
Phlomoides strigosa (C. Y． Wu)Kamelin ＆
Makhm．* 糙毛草糙苏 (新拟) ［Yunnan］
Phlomoides szechuanensis (C. Y． Wu)Kame-
lin ＆ Makhm．* 柴续断［Sichuan］
Phlomoides tatsienensis (Bureau ＆ Franch．)
Kamelin ＆ Makhm． * 康定草糙苏 (新拟)
Phlomoides tatsienensis var． tatsienensis* 康
定草糙苏原变种［Sichuan］
Phlomoides tatsienensis var． hirticalyx(Hand．-
Mazz．)C. L． Xiang ＆ H． Peng* 毛萼康定草糙苏
(新拟) ［Yunnan］
Phlomoides tibetica (C． Marquand ＆ Airy Shaw)
Kamelin ＆ Makhm． * 西藏草糙苏 (新拟)
Phlomoides tibetica var． tibetica* 西藏草糙苏
原变种［Xizang］
Phlomoides tibetica var． wardii (C． Marquand
＆ Airy Shaw)C. L． Xiang ＆ H． Peng* 毛盔西藏草
糙苏 (新拟) ［Xizang］
Phlomoides tuberosa (L．)Moench 块根草糙
苏 (新拟) ［Heilongjiang，Inner Mongolia，Xin-
jiang;Kazakhstan，Kyrgyzstan，Mongolia，Ｒussia;
SW Asia，Europe］
Phlomoides umbrosa (Turcz．)Kamelin ＆ Ma-
khm． * 草糙苏
Phlomoides umbrosa var． umbrosa* 草糙苏原
变种［Gansu，Guangdong，Guizhou，Hebei，Hubei，
Liaoning，Inner Mongolia，Shaanxi，Shandong，Shan-
xi，Sichuan］
Phlomoides umbrosa var． australis (Hemsl．)
C. L． Xiang ＆ H． Peng* 南方草糙苏 (新拟) ［An-
hui，Gansu，Guizhou，Hubei，Hunan，Shaanxi，Si-
chuan，Yunnan］
Phlomoides umbrosa var． latibracteata (Y. Z．
655 植 物 分 类 与 资 源 学 报 第 36卷
Sun)C. L． Xiang ＆ H． Peng* 宽苞草糙苏 (新拟)
［Henan］
Phlomoides umbrosa var． ovalifolia (C. Y． Wu)
C. L． Xiang ＆ H． Peng* 卵齿草糙苏 (新拟) ［An-
hui，Jiangsu］
Phlomoides umbrosa var． stenocalyx (Diels)
C. L． Xiang ＆ H． Peng* 狭萼草糙苏 (新拟) ［Gan-
su，Shaanxi］
Phlomoides uniceps (C. Y． Wu)Kamelin ＆
Makhm．* 单头草糙苏 (新拟) ［Gausu］
Phlomoides younghusbandii (S. M． Mukerjee)
Kamelin ＆ Makhm．* 螃蟹甲［Xizang］
3． Key to the species of the genus Phlomoides from China
1． Calyx teeth spinescent，hooked at apex 2………………………………………………………………………………………
1． Calyx teeth not spinescent and hooked at apex 3………………………………………………………………………………
2． Leaf base broadly cuneate to roundish;corolla reddish or yellow;nutlets glabrous P. hamosa…………………………
2． Leaf base shallowly cordate;corolla white;nutlets stellate at apex P. longiarsitata………………………………………
3． Nutlets prominently hairy at apex;calyx teeth truncate，apiculate at apex 4…………………………………………
4． Calyx funnelform，dilated at apex，especially in fruit，membranous P. molucelloides……………………………
4． Calyx tubular or tubular-campanulate，not dilated at apex，not membranous 5…………………………………
5． Basal leaves pinnatipartite to bipinnatipartite P. speciosa……………………………………………………
5． Basal leaves not pinnatipartite，irregularly shallow serrate or crenate 6………………………………………
6． Bracts leaflike，separated at base;leaves irregularly shallow serrate P. fulgens…………………………
6． Bracts spinescent，united at base in 3s;leaves crenate 7………………………………………………
7． Plants sparsely white hairy;calyx teeth with long soft spines P. multifurcata………………………
7． Plants densely white hairy;calyx teeth with robust rigid spines P. desertorum………………………
3． Nutlets slightly hairy of glabrous;calyx characters not as above 8……………………………………………………
8． Upper corolla lip not fringed or incised;leaf venation fan-shaped P. rotata……………………………………
8． Upper corolla lip always hairy or fringed-incised;leaf venation not fan-shaped 9………………………………
9． Style equally 2-cleft at apex P. cuneata………………………………………………………………………
9． Style unequally 2-cleft at apex 10……………………………………………………………………………
10． Plants with basal rosette of leaves 11……………………………………………………………………
11． Posterior filaments without appendages at base 12…………………………………………………
12． Plants less than 5 cm tall P. pygmaea……………………………………………………………
12． Plants more than 5 cm tall 13……………………………………………………………………
13． Leaf glabrous and calyx margin ciliate P. atropurpurea……………………………………
13． Plant uniformly hairy 14……………………………………………………………………
14． Bracts densely brown ciliate P. tibetica…………………………………………………
14． Bracts densely white villous 15…………………………………………………………
15． Stems retrorse appressed villous;all verticillasters approximate;nutlets stellate pu-
berulent at apex P. oreophila………………………………………………………
15． Stems stellate puberulent;at least basal verticillasters separate;nutlets glabrous
P. chinghoensis……………………………………………………………………
11． Posterior filaments with appendages at base 16………………………………………………………
16． Nutlets hairy 17…………………………………………………………………………………
17． Floral leaves subverticillate，ovate to broadly triangular-ovate;plants ca． 10 cm tall
P. uniceps……………………………………………………………………………………
17． Floral leaves not arranged as above;plants taller 18………………………………………
18． Abaxial leaf surface with unbranched hairs or subglabrous 19…………………………
19． Leaves cordate，crenate，with unbranched hairs on both surfaces P. alpina………
7555期 XIANG Chun-Lei et al．:Taxonomic Notes on the Genus Phlomoides (Lamiaceae:Lamioideae)from China
19． Leaves triangular to ovate，cordate at base，irregularly coarsely crenate，adaxially
sparsely hispid or glabrous，abaxially glabrous or hispid on veins，or hispid on both
surfaces 20…………………………………………………………………………
20． Verticillaster 1，terminal P. kansuensis…………………………………………
20． Verticillaster 4s 3－10，on upper part of stems and branches P. tubrosa………
18． Abaxial leaf surface with stellate hairs 21………………………………………………
21． Leaves with stellate hispid and simple hairs，or with stellate strigose and simple hairs
22……………………………………………………………………………………
22． Basel leaves lanceolate-oblong to narrowly oblong，adaxially with stellate strigose
and simples hairs，abaxially stellate tomentulose P. younghusbandii…………
22． Basal leaves ovate-triangular to triangular-lanceolate，adaxially stellate hispid with
long central arms and simple hairs，or sparsely hispid to subglabrous，abaxially
sparsely to densely stellate pilose P. mongolica…………………………………
21． Leaves with simple hairs adaxially 23………………………………………………
23． Bracts lanceolate or narrow lanceolate P. maximowiczii………………………
23． Bracts linear，needlelike，or linear-subulate 24………………………………
24． Stems simple or branched;bracts slightly shorter than calyx;calyx densely
stellate puberulent outside P. agraria………………………………………
24． Stems many branched;bracts subequal to calyx，calyx sparsely stellate pu-
berulent to hispid outside P. medicinalis……………………………………
16． Nutlets glabrous 25………………………………………………………………………………
25． Basal leaves early deciduous;stem leaves hastate-ovate P. pararotata……………………
25． Basal leaves persistent;stem leaves narrowly to broadly ovate 26…………………………
26． Basal leaves broadly cordate，sparsely strigose and corrugate adaxially P. koraiensis…
26． Leaves triangular-ovate，cordate to ovate-lanceolate 27…………………………………
27． Corolla reddish pink 28………………………………………………………………
28． Stem hairs with unequal arms;leaves narrowly ovate to oblong-lanceolate
P. betonicoides…………………………………………………………………
28． Stem hairs with longer central arms;leaves triangular to triangular-ovate
P. dentosa………………………………………………………………………
27． Corolla purple-red or dark purple 29………………………………………………
29． Corolla dark purple;Leaves broadly ovate to circular-ovate，apex rounded，base
cordate，sparsely strigose P. fimbriata…………………………………………
29． Corolla purple-red;leaves acute to obtuse at apex，shallowly cordate at base
30………………………………………………………………………………
30． Leaves cordate-ovate or ovate-oblong，stellate pilose and with simple hairs on
both surfaces or abaxially stellate pubescent P. pratensis…………………
30． Leaves ovate or triangular-ovate，adaxially densely hispid，abaxially densely
stellate pilose P. milingensis………………………………………………
10． Plants without basal leaf rosette，only stem leaves present 31…………………………………………
31． Corolla more than 3 cm 32……………………………………………………………………………
32． Corolla yellow P. megalantha……………………………………………………………………
32． Corolla dark purple P. ornata……………………………………………………………………
31． Corolla less than 3 cm 33……………………………………………………………………………
33． Verticillasters conspicuously pedunculate 34……………………………………………………
34． Posterior filaments without appendages at base 35…………………………………………
855 植 物 分 类 与 资 源 学 报 第 36卷
35． Bracts ca． 1 /2 as long as calyx;stems densely tawny stellate pubescent
P. szechuanensis…………………………………………………………………………
35． Bracts as long as or slightly longer than calyx;stems sparsely retrorse minute hispid，
sometimes stellate pubescent at apex P. umbrosa………………………………………
34． Posterior filaments with appendages at base 36………………………………………………
36． Calyx gray stellate pannose outside，bracts leaflike，linear-lanceolate，rarely lanceolate
P. likiangensis…………………………………………………………………………
36． Calyx with other types of hairs kind;bracts usually smaller，narrower 37……………
37． Verticillasters rather loose，usually 1，rarely 2 at apes of stem or branch;flowers long
pedicellate;calyx indehiscent P. pedunculata………………………………………
37． Verticillasters rather compact，usually 2－5 (－9)at apex of stem or branch;flowers
short pedicellate;calyx dehiscent P. congesta………………………………………
33． Verticillasters sessile 38…………………………………………………………………………
38． Bracts ca． 1 /2 as long as calyx or absent 39………………………………………………
39． Calyx ca． 1. 5 cm P. longicalyx…………………………………………………………
39． Calyx 0. 8－1 cm 40………………………………………………………………………
40． Corolla purple P. franchetiana………………………………………………………
40． Corolla yellow P. ambigua…………………………………………………………
38． Bracts more than 1 /2 as long as calyx 41……………………………………………………
41． Leaves cordate to broadly ovate P. paohsingensis………………………………………
41． Leaves of different shapes 42……………………………………………………………
42． Leaves abaxially densely stellate tomentose P. muliensis……………………………
42． Leaves abaxially without hairs as above 43…………………………………………
43． Posterior filaments with a reflexed slender appendage at base 44………………
44． Leaves strigose adaxially and along margin abaxially，with elevated glands
P. melanantha………………………………………………………………
44． Leaves sparsely minutely hispid adaxially，abaxially stellate pilose，densely
and finely spreading setose along veins P. forrestii…………………………
43． Posterior filaments with an upwardly curved appendage at base 45……………
45． Leaves adaxially sparsely short bristly，abaxially pilose，with simple hairs on
both surfaces P. jeholensis…………………………………………………
45． Leaves adaxially with simple and /or stellate hairs，abaxially with stellate and
simple hairs 46………………………………………………………………
46． Calyx indehiscent，veins elevated 47……………………………………
47． Stems sparsely retrorse minutely hispid;calyx tube densely stellate
puberulent，veins spreading bristly P. setifera……………………
47． Stems stellate pubescent or densely tawny stellate tomentulose;calyx
tube stellate pubescent P. tatsienensis………………………………
46． Calyx irregularly dehiscent，veins not elevated 48……………………
48． Leaves leathery more or less，sparsely strigose or subglabrous，margin
ciliolate，abaxially subglabrous or stellate strigose and hairs with a
long central arm P. strigosa…………………………………………
48． Leaves herbaceous，adaxially sparsely strigose or stellate pubescent，
abaxially stellate pubescent or dense stellate pilose 49……………
49． Stem leaves ovate;bracts linear P. inaequalisepala……………
49． Stem leaves sagittate-oblong;bracts subulate P. ruptilis………
9555期 XIANG Chun-Lei et al．:Taxonomic Notes on the Genus Phlomoides (Lamiaceae:Lamioideae)from China
Acknowledgements:We thank two anonymous reviewers for
their constructive comments that improved the quality of the
paper．
Ｒeferences:
Adylov TA，Kamelin ＲV，Makhmedov AM，1986． Zametkio semei-
stve Lamiaceae 1(Notes on Lamiaceae 1) ［J］． Novosti Sistemati-
ki Vysshikh Ｒastenii，23:110—114
Adylov TA，Makhmedov AM，1987． Phlomoides Moench ［A］． In:
Adylov TA (ed．) ，Conspectus florae Asiae Mediae ［M］． Tash-
kent:Izdatel’stvo Akademii Nauk SSSＲ，9:82—107
Astanova SB，1984． Chromosome numbers in the species of the fami-
lies Alliaceae，Asteraceae，Caryophyllaceae，Ebenaceae，Linace-
ae，Oleaceae，Lamiaceae from Tadjikistan ［J］． Botanicˇhnyi
urnal (Moscow ＆ Leningrad) ，69:1563—1564
Azizian D，Cutler DF，1982． Anatomical，cytological and phytochem-
ical studies on Phlomis L． and Eremostachys Bunge (Labiatae)
［J］． Botanical Journal of the Linnean Society，85:249—281
Bentham G，1829． Notochaete ［J］． Edwards’s Botanical Ｒegister，
15:pl． 1289
Bentham G，1832－1836． Labiatarum Genera et Species［M］． London:
James Ｒidgway ＆ Sons
Boissier E，1879． Flora orientalis，Vol. 4，pt. 1［M］． Geneva，Ba-
sel，Leiden:H． Georg
Briquet J，1895 － 1897． Labiatae． Eremostachys Bunge，Phlomis L
［A］． In:Engler A，Prantl K (eds．) ，Die natürlichen Pflanzen-
familein ［M］． Leipzig:Engelmann，4 (3a，134) :246—249
Brullo S，Guglielmo A，Pavone P et al．，1990． Chromosome counts of
flowering plants from N． Cyrenaica ［J］． Candollea，45:65—74
Bunge A，1830． Eremostachys Bunge［A］． In:Ledebour CF，Meyer
CA，Bunge A (eds．) ，Flora Altaica ［M］． Berlin:G． Ｒeimer，
2:414—416
Fang LQ(房丽琴) ，Pang YZ(潘跃芝) ，Gong X(龚洵) ，2007． A
karyomorphological study in the monotypic genus Lamiophlomis
and five species in Phlomis (Lamiaceae) ［J］． Acta Phytotaxo-
nomica Sinica (植物分类学报) ，45:627—632
Ghaffari SM，2006． New or rare chromosome counts of some angio-
sperm species from Iran ［J］． Iranian Journal of Botany，11:
185—192
Kamelin ＲV，Makhmedov AM，1990a． The system of the genus Phlo-
moides (Lamiaceae) ［J］． Botanicˇhnyi urnal (Moscow ＆ Len-
ingrad) ，75:241—250
Kamelin ＲV，Makhmedov AM，1990b． A new system of the genus
Phlomis(Lamiaceae) ［J］． Botanicˇhnyi urnal(Moscow ＆ Len-
ingrad) ，75:1163—1167
Kud Y， 1929． Labiatarum Sino-Japonicarum prodormus ［A］．
Memoirs of the Faculty of Science and Agriculture，Taihoku Im-
perial University，2:1—332
Mathiesen C，Scheen AC，Lindqvist C，2011． Phylogeny and bioge-
ography of the lamioid genus Phlomis (Lamiaceae) ［J］． Kew
Bulletin，66:83—99
Moench C，1794． Methodus Plantas Horti Botanici et Argi Marburgen-
sis，A Staminum Situ Describendi［M］． Marburg:Officina Nova
Libraria Academiae
Pan YZ，Fang LQ，Hao G et al．，2009． Systematic positions of Lami-
ophlomis and Paraphlomis (Lamiaceae)based on nuclear and
chloroplast sequences ［J］． Journal of Systematics and Evolution，
47:535—542
Ｒyding O，2008． Pericarp structure and phylogeny of the Phlomis
group (Lamiaceae subfam． Lamioideae) ［J］． Botanische Jahrbü-
cher für Systematik，Pflanzengeschichte und Pflanzengeographie，
127:299—316
Salmaki Y，Zarres S，Heubl G，2012a． The genus Phlomoides Moench
(Lamiaceae;Lamioideae;Phlomideae)in Iran:an updated syn-
opsis ［J］． Iranian Journal of Botany，18:207—219
Salmaki Y，Zarre S，Ｒyding O et al．，2012b． Phylogeny of the tribe
Phlomideae (Lamioideae:Lamiaceae)with special focus on Ere-
mostachys and Phlomoides:new insights from nuclear and chloro-
plast sequences［J］． Taxon，61 (1) :161—179
Wu CY (吴征镒) ，Li HW (李锡文) ，1977． Phlomis L．［A］． In:
Wu CY (吴征镒) (ed．) ，Flora Ｒeipublicae Popularis Sinica
［M］． Beijing:Science Press，65 (2) :428—478
065 植 物 分 类 与 资 源 学 报 第 36卷