全 文 :中国唇形科草糙苏属分类学研究杂记∗
向春雷1ꎬ 董洪进1ꎬ 胡国雄1ꎬ2ꎬ 周 静3ꎬ 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 云南 昆明 650201ꎻ 2 中国科学院大学ꎬ
北京 100049ꎻ 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ꎬ 云南 昆明 650500)
摘要: 最近的研究结果表明ꎬ 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)ꎮ 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ꎮ 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ꎬ 但仍遗漏了 11个物种名称ꎮ 为方便使用ꎬ 本文对这 11个名称进行了新
组合ꎬ 同时提供了新界定的草糙苏属国产物种名录及检索表ꎬ 共 50种 9变种ꎮ
关键词: 糙苏属ꎻ 糙苏族ꎻ 草糙苏属ꎻ 唇形科ꎻ 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 ̄Lei1ꎬ DONG Hong ̄Jin1ꎬ HU Guo ̄Xiong1ꎬ2ꎬ ZHOU Jing3ꎬ 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 University of Chinese Academy of Sciencesꎬ Beijing 100049ꎬ Chinaꎻ
3 School of Pharmaceutical Science & Yunnan Key Laboratory of Pharmacology for Nature Productsꎬ
Kunming Medical Universityꎬ Kunming 650500ꎬ China)
Abstract: Recent phylogenetic studies have investigated the tribe Phlomideae andꎬ in particularꎬ the degree of “in ̄
clusiveness” of the genus Phlomoides. As a resultꎬ Eremostachysꎬ Lamiophlomisꎬ Notochaete and all Chinese Phlo ̄
misꎬ excepted for cultivated P fruticosaꎬ 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 elsewhereꎬ unfortunatelyꎬ eleven names were o ̄
mitted by previous researchers. Thereforeꎬ this paper proposes 11 new nomenclatural combinations in Phlomoidesꎬ
and provides a checklist and a key to all Chinese Phlomoides.
Key words: Phlomisꎻ Tribe Phlomideaeꎻ Phlomoidesꎻ Lamiaceaeꎻ New combinations
The genus Phlomoides Moench ( Phlomideaeꎬ
Lamioideaeꎬ Lamiaceae) is the species ̄rich genus
with about 150-170 speciesꎬ extending from central
Europe to the Russian Far East (Mathiesen et al.ꎬ
2011ꎻ Salmaki et al.ꎬ 2012aꎬ b). The genus was
separated from Phlomis L. by Moench (1794) who
placed Phlomis tuberosa L. in his monotypic genus
Phlomoides. Howeverꎬ in the past 200 yearsꎬ most
taxonomists (Benthamꎬ 1832-1836ꎻ Boissierꎬ 1879ꎻ
Briquetꎬ 1895-1897ꎻ Wu and Liꎬ 1977) had treated
植 物 分 类 与 资 源 学 报 2014ꎬ 36 (5): 551~560
Plant Diversity and Resources DOI: 10.7677 / ynzwyj201413228
∗
∗∗
Funding: The National Natural Science Foundation of China (31110103911ꎬ 31100164)ꎻ Knowledge Innovation Project of the Chinese Acade ̄
my of Sciences (KSCX2 ̄EW ̄Z ̄1)ꎻ the Ministry of Science and Technology Chinaꎬ Basic Research Project (2013FY112100)
Author for correspondenceꎻ E ̄mail: hpeng@mail kib ac cn
Received date: 2013-11-25ꎬ Accepted date: 2014-07-14
作者简介: 向春雷 (1984-) 男ꎬ 博士ꎬ 副研究员ꎬ 主要从事植物分类与系统发育研究ꎮ 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.ꎬ
1986ꎻ Adylov and Machmedovꎬ 1987ꎻ Kamelin and
Machmedovꎬ 1990aꎬb). The reinstated Phlomoides
includes some species with the upper corolla lip
not laterally compressed and with tuberous lateral
roots ( Salmakiꎬ 2012a). Meanwhileꎬ some other
genera such as Eremostachys Bungeꎬ Notochaete
Benth.ꎬ Paraeremostachys Adylovꎬ Kamelin & Ma ̄
khm.ꎬ Lamiophlomis Kudô were established or sepa ̄
rated from Phlomis (Benthamꎬ 1829ꎻ Bungeꎬ 1830ꎻ
Kudôꎬ 1929ꎻ Adylov et al.ꎬ 1986)ꎬ and all of them
were considered closely related to Phlomis. Based
on pericarp structureꎬ Ryding ( 2008) agreed that
Phlomoides should be treated as a separate genusꎬ
including the species from Lamiophlomis and No ̄
tochaeteꎬ but he didn’ t make normal taxonomical
treatments.
Pan et al. ( 2009) supported the inclusion of
Lamiophlomis in Phlomis based on their molecular
phylogenetic studyꎬ however only fourteen species of
Phlomis were included in their study. Recentlyꎬ u ̄
sing DNA sequences of the plastid trnL intronꎬ trnL ̄
trnF spacer and rps16 intronꎬ Phlomis s l. were split
into two lineagesꎬ i e. “ Phlomoides group ” and
“Phlomis group”ꎬ and therefore Mathiesen et al.
(2011) recognized two generaꎬ i e.ꎬ Phlomis s str.
and Phlomoides s l.ꎬ and the latter included Pseud ̄
eremostachysꎬ Lamiophlomis as well as Notochaete
hamosa Benth. In the following molecular phyloge ̄
netic study on the tribe Phlomideaeꎬ Salmaki et al.
(2012b) confirmed the inclusion of the members of
Eremostachysꎬ and other mono ̄ or oligotypic genera
(e g. Lamiophlomisꎬ Notochaeteꎬ Paraeremostachys)
in Phlomoidesꎬ and some necessary new combina ̄
tions are proposed therein.
Nowꎬ circumscription for Phlomoides has been
clearꎬ 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 Culterꎬ
1982ꎻ Astanovaꎬ 1984ꎻ Brullo et al.ꎬ 1990ꎻ Ghaf ̄
fariꎬ 2006ꎬ Fang et al.ꎬ 2007). According to the
current circumscription of Phlomoidesꎬ there are for ̄
ty ̄nine species and ten varieties in Chinaꎻ of which
forty ̄two species and nine varieties were previously
in Phlomisꎬ four species and one variety were previ ̄
ously in Eremostachysꎬ 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 Makhmedovꎬ 1987ꎻ Matheisen et al.ꎬ
2011ꎻ Salmaki et al.ꎬ 2012b)ꎬ eleven recognized
taxa are omitted by these authors. In order to aid fu ̄
ture systematic work on these taxaꎬ here we propose
eleven new combinationsꎬ and for the first timeꎬ 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 / ortubers at tip Perennial herbs or small shrubs
Petioles 2-5 (-7) cm 1 5-5 cm
Calyx lobe equal or occasionally 3 / 2 bilabiateꎬ abruptly narrowto acute apex equal
Posterior lip of corolla longꎬ shallowly hooded longꎬ hoodedꎬ 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 Glabrousꎬ papillose or stellate hairy
Chromosome number 2n= 22 2n= 12ꎬ 14ꎬ 20ꎬ 22ꎬ 40ꎬ 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. Pengꎬ comb. nov.
Basionym: Phlomis cuneata C Y. Wuꎬ Fl. Rei ̄
publ. Popularis Sin. 65 ( 2): 434ꎬ 594 ( 1977).
TYPE: Chinaꎬ Xizang: Cuijila Shanꎬ 6 Sep. 1952ꎬ
P C. Tsoong 5335 (PE).
Distribution: Endemic to Xizang (Tibet)ꎬ and
currently known only from the type locality (Cuijila
Shan).
Note: The style of this species is equally 2 ̄
cleft at apexꎬ which is very rare in genus Phlomoides
from China.
Phlomoides dentosa (Franch.) Kamelin & Ma ̄
khm. var. glabrscens (Danguy) C L. Xiang & H.
Pengꎬ comb. nov.
Basionym: Phlomis dentosa Franch. var. glabr ̄
scens Danguyꎬ Bull. Mus. Natl. Hist. Nat. 17 (5):
345 (1911). TYPE: Chinaꎬ Gansu: Si ̄Ning ̄Fouꎬ
2 400 mꎬ 18th Jul. 1908ꎬ Anonymous s n. (P).
Distribution: Endemic to Chinaꎬ known from
Gansuꎬ Hebeiꎬ Inner Mongoliaꎬ and Qinghai.
Note: Morphological characters distinguished
this variety from Phlomoides dentosa var. dentosa are
that stems sparsely minutely hispid to subglabrousꎬ
leave subglabrousꎬ 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. Pengꎬ comb. nov.
Basionym: Phlomis mongolica Turcz. var. ma ̄
crocephala C Y. Wuꎬ Fl. Reipubl. Popularis Sin. 65
(2): 444ꎬ 595 (1977). TYPE: Chinaꎬ Inner Mon ̄
golia: precisely locality unknownꎬ 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 tallꎬ verticillasters solitaryꎬ terminal or axillaryꎬ
and sometimes inserted at base of stem. In contrastꎬ
plants of Phlomoides mongolica var. mongolica are
40- 70 cm tallꎬ verticillasters numerousꎬ axillaryꎬ
inserted on apical part of stems.
Phlomoides oreophila (Kar. & Kir.) Adylov.
Kamelin & Makhm. var. evillosa (C Y. Wu) C L.
Xiang & H. Pengꎬ comb. nov.
Basionym: Phlomis oreophila Kar. & Kir. var.
evillosa C Y. Wuꎬ Fl. Reipubl. Popularis Sin. 65
(2): 437ꎬ 594 (1977). TYPE: Chinaꎬ Xinjiang:
Fuhaiꎬ 2 100 mꎬ 11 Jul. 1957ꎬ 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. oreophilaꎬ
bracts densely villous or sometime intermixed with
glandular villous hairsꎬ and calyx stellate puberu ̄
lentꎬ finely villous on veins outside.
Phlomoides speciosa (Rupr.) Adylovꎬ Kame ̄
lin & Makhm. var. viridifolia (Popov) C L. Xiang
& H. Pengꎬ comb. nov.
Basionym: Eremostachys speciosa Rupr. var.
viridifolia Popovꎬ Nov. Mem. Moskovsk. Obsc. Isp.
Prir. 19: 100 (1940). SYNETYPES: Kazakhstanꎬ
Semirechenskaya Region: 30th Apr. 1916ꎬ Goro ̄
detskiy 158 (TASH)ꎻ Uzbekistanꎬ Fergana Region:
Andizhan Districtꎬ on the way to Kugart Passꎬ 2
Jun. 1911ꎬ Knorring & Minkwitz 266 (TASH)ꎻ Sa ̄
markand Region: Zimmarkhꎬ 16th Jul. 1915ꎬ Bala ̄
baye 9 (TASH).
Distribution: This variety is recorded from Xin ̄
jiang.
Note: Variety viridifolia is distributed to Ka ̄
zakhstanꎬ Kyrgyzstanꎬ Tajikistanꎬ Turkmenistnaꎬ and
SW Asia. In additionꎬ because all the Chinese Ere ̄
mostachys species were transferred to Phlomoidesꎬ 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. Pengꎬ comb. nov.
Basionym: Phlomis franchetiana Diels var. hir ̄
ticalyx Hand. ̄Mazz. Symb. Sin. 7 (4): 921 (1936).
TYPE: Chinaꎬ Yunnan: Beyenjingꎬ in silvisꎬ 25
Sept. 1919ꎬ 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. Pengꎬ comb. nov.
Basionym: Phlomis tibetica C. Marquand &
Airy Shaw var. wardii C. Marquand & Airy Shawꎬ J.
Linn. Soc.ꎬ Bot. 48 (321): 218-219 (1929). TYPE:
Chinaꎬ Xizang: on the edges of cultivated fieldsꎬ Tum ̄
batseꎬ 3 400 mꎬ 14 Jul. 1924ꎬ F K. Ward 5937 (K).
Distribution: Endemic to Xizang.
Note: This variety is distinguished from variety
tibetica by several traitsꎬ i e. leaf blade cordate to
broadly oblongꎬ margin coarsely crenate ( vs. leaf
blade ovate ̄cordateꎬ margin crenate or coarsely cre ̄
nate in variety tibetica)ꎻ 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. Pengꎬ
comb. nov.
Basionym: Phlomis umbrosa Turcz. var. austra ̄
lis Hemsl.ꎬ J. Linn. Soc.ꎬ Bot. 26 (175): 306 (1890).
Distribution: This variety is recorded from An ̄
huiꎬ Gansuꎬ Guizhouꎬ Hubeiꎬ Hunanꎬ Shanxiꎬ Si ̄
chuanꎬ and Yunnan. SYNTYPES: Chinaꎬ Hubei:
Yichangꎬ Badongꎬ A. Henry 875ꎬ 1872ꎬ 2442ꎬ
4749ꎬ 6029ꎬ 6179ꎬ 6423ꎬ 7360 (K).
Note: There are 4 varieties under Phlomoides
umbrosa. In this species complexꎬ variety australis is
most similar to variety umbrosaꎬ but differentiated by
following traits. In variety australisꎬ leaves long peti ̄
olateꎬ leaf blade membranousꎬ margin crenate ̄ser ̄
rateꎬ terminal tooth sometimes very longꎻ bracts
softꎬ linear ̄lanceolateꎬ slightly shorter than calyxꎬ
while in variety umbrosaꎬ leaves paperyꎬ serrate ̄
dentate or irregularly crenateꎬ terminal tooth not very
longꎻ bracts rigidꎬ linear ̄subulateꎬ mostly longer
than calyx.
Phlomoides umbrosa (Turcz.) Kamelin & Mak ̄
hm. var. latibrateata (Y Z. Sun) C L. Xiang & H.
Pengꎬ comb. nov.
Basionym: Phlomis umbrosa Turcz. var. lati ̄
brateata Y Z. Sunꎬ Acta Phytotax. Sin. 11 (1): 46
(1966). TYPE: Chinaꎬ Henan: Jigong Shanꎬ hill ̄
sidesꎬ 700-2 000 mꎬ 27 Jul. 1925ꎬ A. N. Steward
1584 (NAS).
Distribution: Endemic to Henan.
Note: In Phlomoides umbrosa complexꎬ this
variety is easily distinguished from other taxa be ̄
cause its bracts are linear ̄oblong to obovate ̄oblongꎬ
and much shorter than calyxꎬ only 5 -7×1 8 -2 5
cmꎬ while in other taxaꎬ bracts are linear ̄subulate
to linear ̄lanceolateꎬ longer or rarely slightly shorter
than calyxꎬ about 8-14×1-2 mm.
Phlomoides umbrosa (Turcz.) Kamelin & Mak ̄
hm. var. ovalifolia (C Y. Wu) C L. Xiang & H.
Pengꎬ comb. nov.
Basionym: Phlomis umbrosa Turcz. var. ovali ̄
folia C Y. Wuꎬ Fl. Reipubl. Popularis Sin. 65 (2):
478ꎬ 601 (1977). TYPE: Chinaꎬ Anhui: Chuxianꎬ
hillsidesꎬ 2 Aug. 1951ꎬ East ̄China Station 3132
(PE).
Distribution: This variety is recorded from An ̄
hui and Jiangsu.
Note: In Phlomoides umbrosa complexꎬ this
variety is similar to variety australis and type varie ̄
tyꎬ but differentiated by leaves ovateꎬ subcuneate to
shallowly cordateꎬ stellate pubescentꎬ rarely abaxial ̄
ly stellate tomentuloseꎻ calyx densely stellate pubes ̄
cent outside ( vs. leaves circular ̄ovate to ovate ̄ob ̄
longꎬ sparsely pilose or sometimes abaxially densely
455 植 物 分 类 与 资 源 学 报 第 36卷
pilose and stellate piloseꎻ calyx stellate puberulent
except sometime pilose on veins).
Phlomoides umbrosa Turcz. var. stenocalyx
(Diels) C L. Xiang & H. Pengꎬ comb. nov.
Basionym: Phlomis stenocalyx Dielsꎬ Bot. Jahrb.
Syst. 29 (3 ̄4): 555 (1900). TYPE: Chinaꎬ 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 ̄
lateꎬ laxꎬ and flowers pedicellateꎬ calyx ca. 8 × 3 cm.
2. Checklist of Phlomoides Moench from
China
Nowꎬ there is only one species of Phlomisꎬ i e.
P fruticosa L. cultivated in China ( Shaanxi Prov ̄
ince). According to the new definition of Phlo ̄
moidesꎬ there are 50 species and 9 varieties in Chi ̄
naꎻ of which 40 species and 9 varieties are endemic
and marked with asterisk (∗) in following listꎬ and
distribution area listed in square bracket.
Phlomoides agraria (Bunge) Adylovꎬ Kame ̄
lin & Makhm. 耕地草糙苏 (新拟) [Xinjiangꎻ Ka ̄
zakhstanꎬ Mongoliaꎬ Russia]
Phlomoides alpina (Pall.) Adylov. Kamelin &
Makhm. 高山草糙苏 (新拟) [Xinjiangꎻ Kazakh ̄
stanꎬ Russia]
Phlomoides ambigua (Popov ex Pazij & Vved.)
Adylovꎬ 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 [Sichuanꎬ Xizangꎬ 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 [Sichuanꎬ 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 [Gansuꎬ Hebeiꎬ Inner Mongoliaꎬ Qinghai]
Phlomoides dentosa var. glabrescens (Danguy)
C L. Xiang & H. Peng∗渐光尖齿草糙苏 (新拟)
[Sichuanꎬ 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) Adylovꎬ Kamelin
& Makhm. 光沙穗 [Xinjiangꎻ Kyrgyzstanꎬ SW Asia]
Phlomoides hamosa (Benth.) Mathiesen 钩萼
草 [Yunnanꎻ Bhutanꎬ Indiaꎻ Myanmarꎬ 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) [Jilinꎻ 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 [Xizangꎬ 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) [Hebeiꎬ Jilinꎬ Lia ̄
oning]
Phlomoides medicinalis ( Diels ) Kamelin &
Makhm.∗萝卜秦艽 [Sichuanꎬ Xizang]
Phlomoides megalantha ( Diels) Kamelin &
Makhm.∗大花草糙苏 (新拟) [Hubeiꎬ Shaanxiꎬ
Shanxiꎬ 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) [Sichuanꎬ 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 [Xinjiangꎻ Kazakhstanꎬ Kyrgyzstanꎬ Mongoliaꎬ
Russiaꎬ Tajikistanꎻ SW Asiaꎬ Europe]
Phlomoides mongolica ( Turcz.) Kamelin &
Makhm. ∗串铃草
Phlomoides mongolica var. mongolica∗串铃
草原变种 [Gansuꎬ Hebeiꎬ Inner Mongoliaꎬ Shaanxiꎬ
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 [ Xinjiangꎻ Kazakhstanꎬ Kyrgyzstanꎬ
Mongoliaꎬ Russiaꎬ 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) [Sichuanꎬ 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 ̄
jiangꎻ Kazakhstanꎬ 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 [Gansuꎬ Qinghaiꎬ Sichuanꎬ Xizangꎬ
Yunnanꎻ Bhutanꎬ Indiaꎬ 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) [Sichuanꎬ
Xizangꎬ Yunnan]
Phlomoides speciosa (Rupr.) Adylovꎬ 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) [ Heilongjiangꎬ Inner Mongoliaꎬ Xin ̄
jiangꎻ Kazakhstanꎬ Kyrgyzstanꎬ Mongoliaꎬ Russiaꎻ
SW Asiaꎬ Europe]
Phlomoides umbrosa (Turcz.) Kamelin & Ma ̄
khm. ∗草糙苏
Phlomoides umbrosa var. umbrosa∗草糙苏原
变种 [Gansuꎬ Guangdongꎬ Guizhouꎬ Hebeiꎬ Hubeiꎬ
Liaoningꎬ Inner Mongoliaꎬ Shaanxiꎬ Shandongꎬ Shan ̄
xiꎬ Sichuan]
Phlomoides umbrosa var. australis (Hemsl.)
C L. Xiang & H. Peng∗南方草糙苏 (新拟) [An ̄
huiꎬ Gansuꎬ Guizhouꎬ Hubeiꎬ Hunanꎬ Shaanxiꎬ Si ̄
chuanꎬ 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 ̄
huiꎬ Jiangsu]
Phlomoides umbrosa var. stenocalyx (Diels)
C L. Xiang & H. Peng∗狭萼草糙苏 (新拟) [Gan ̄
suꎬ 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 spinescentꎬ hooked at apex 2
1. Calyx teeth not spinescent and hooked at apex 3
2. Leaf base broadly cuneate to roundishꎻ corolla reddish or yellowꎻ nutlets glabrous P hamosa
2. Leaf base shallowly cordateꎻ corolla whiteꎻ nutlets stellate at apex P longiarsitata
3. Nutlets prominently hairy at apexꎻ calyx teeth truncateꎬ apiculate at apex 4
4. Calyx funnelformꎬ dilated at apexꎬ especially in fruitꎬ membranous P molucelloides
4. Calyx tubular or tubular ̄campanulateꎬ not dilated at apexꎬ not membranous 5
5. Basal leaves pinnatipartite to bipinnatipartite P speciosa
5. Basal leaves not pinnatipartiteꎬ irregularly shallow serrate or crenate 6
6. Bracts leaflikeꎬ separated at baseꎻ leaves irregularly shallow serrate P fulgens
6. Bracts spinescentꎬ united at base in 3sꎻ leaves crenate 7
7. Plants sparsely white hairyꎻ calyx teeth with long soft spines P multifurcata
7. Plants densely white hairyꎻ calyx teeth with robust rigid spines P desertorum
3. Nutlets slightly hairy of glabrousꎻ calyx characters not as above 8
8. Upper corolla lip not fringed or incisedꎻ leaf venation fan ̄shaped P rotata
8. Upper corolla lip always hairy or fringed ̄incisedꎻ 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 villousꎻ all verticillasters approximateꎻ nutlets stellate pu ̄
berulent at apex P oreophila
15. Stems stellate puberulentꎻ at least basal verticillasters separateꎻ nutlets glabrous
P chinghoensis
11. Posterior filaments with appendages at base 16
16. Nutlets hairy 17
17. Floral leaves subverticillateꎬ ovate to broadly triangular ̄ovateꎻ plants ca. 10 cm tall
P uniceps
17. Floral leaves not arranged as aboveꎻ plants taller 18
18. Abaxial leaf surface with unbranched hairs or subglabrous 19
19. Leaves cordateꎬ crenateꎬ 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 ovateꎬ cordate at baseꎬ irregularly coarsely crenateꎬ adaxially
sparsely hispid or glabrousꎬ abaxially glabrous or hispid on veinsꎬ or hispid on both
surfaces 20
20. Verticillaster 1ꎬ terminal P kansuensis
20. Verticillaster 4s 3-10ꎬ on upper part of stems and branches P tubrosa
18. Abaxial leaf surface with stellate hairs 21
21. Leaves with stellate hispid and simple hairsꎬ or with stellate strigose and simple hairs
22
22. Basel leaves lanceolate ̄oblong to narrowly oblongꎬ adaxially with stellate strigose
and simples hairsꎬ abaxially stellate tomentulose P younghusbandii
22. Basal leaves ovate ̄triangular to triangular ̄lanceolateꎬ adaxially stellate hispid with
long central arms and simple hairsꎬ or sparsely hispid to subglabrousꎬ 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 linearꎬ needlelikeꎬ or linear ̄subulate 24
24. Stems simple or branchedꎻ bracts slightly shorter than calyxꎻ calyx densely
stellate puberulent outside P agraria
24. Stems many branchedꎻ bracts subequal to calyxꎬ calyx sparsely stellate pu ̄
berulent to hispid outside P medicinalis
16. Nutlets glabrous 25
25. Basal leaves early deciduousꎻ stem leaves hastate ̄ovate P pararotata
25. Basal leaves persistentꎻ stem leaves narrowly to broadly ovate 26
26. Basal leaves broadly cordateꎬ sparsely strigose and corrugate adaxially P koraiensis
26. Leaves triangular ̄ovateꎬ cordate to ovate ̄lanceolate 27
27. Corolla reddish pink 28
28. Stem hairs with unequal armsꎻ leaves narrowly ovate to oblong ̄lanceolate
P betonicoides
28. Stem hairs with longer central armsꎻ leaves triangular to triangular ̄ovate
P dentosa
27. Corolla purple ̄red or dark purple 29
29. Corolla dark purpleꎻ Leaves broadly ovate to circular ̄ovateꎬ apex roundedꎬ base
cordateꎬ sparsely strigose P fimbriata
29. Corolla purple ̄redꎻ leaves acute to obtuse at apexꎬ shallowly cordate at base
30
30. Leaves cordate ̄ovate or ovate ̄oblongꎬ stellate pilose and with simple hairs on
both surfaces or abaxially stellate pubescent P pratensis
30. Leaves ovate or triangular ̄ovateꎬ adaxially densely hispidꎬ abaxially densely
stellate pilose P milingensis
10. Plants without basal leaf rosetteꎬ 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 calyxꎻ stems densely tawny stellate pubescent
P szechuanensis
35. Bracts as long as or slightly longer than calyxꎻ stems sparsely retrorse minute hispidꎬ
sometimes stellate pubescent at apex P umbrosa
34. Posterior filaments with appendages at base 36
36. Calyx gray stellate pannose outsideꎬ bracts leaflikeꎬ linear ̄lanceolateꎬ rarely lanceolate
P likiangensis
36. Calyx with other types of hairs kindꎻ bracts usually smallerꎬ narrower 37
37. Verticillasters rather looseꎬ usually 1ꎬ rarely 2 at apes of stem or branchꎻ flowers long
pedicellateꎻ calyx indehiscent P pedunculata
37. Verticillasters rather compactꎬ usually 2-5 (-9) at apex of stem or branchꎻ flowers
short pedicellateꎻ 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 abaxiallyꎬ with elevated glands
P melanantha
44. Leaves sparsely minutely hispid adaxiallyꎬ abaxially stellate piloseꎬ 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 bristlyꎬ abaxially piloseꎬ with simple hairs on
both surfaces P jeholensis
45. Leaves adaxially with simple and / or stellate hairsꎬ abaxially with stellate and
simple hairs 46
46. Calyx indehiscentꎬ veins elevated 47
47. Stems sparsely retrorse minutely hispidꎻ calyx tube densely stellate
puberulentꎬ veins spreading bristly P setifera
47. Stems stellate pubescent or densely tawny stellate tomentuloseꎻ calyx
tube stellate pubescent P tatsienensis
46. Calyx irregularly dehiscentꎬ veins not elevated 48
48. Leaves leathery more or lessꎬ sparsely strigose or subglabrousꎬ margin
ciliolateꎬ abaxially subglabrous or stellate strigose and hairs with a
long central arm P strigosa
48. Leaves herbaceousꎬ adaxially sparsely strigose or stellate pubescentꎬ
abaxially stellate pubescent or dense stellate pilose 49
49. Stem leaves ovateꎻ bracts linear P inaequalisepala
49. Stem leaves sagittate ̄oblongꎻ 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 TAꎬ Kamelin RVꎬ Makhmedov AMꎬ 1986. Zametkio semei ̄
stve Lamiaceae 1 (Notes on Lamiaceae 1) [J] . Novosti Sistemati ̄
ki Vysshikh Rasteniiꎬ 23: 110—114
Adylov TAꎬ Makhmedov AMꎬ 1987. Phlomoides Moench [ A]. In:
Adylov TA ( ed.)ꎬ Conspectus florae Asiae Mediae [M]. Tash ̄
kent: Izdatel’stvo Akademii Nauk SSSRꎬ 9: 82—107
Astanova SBꎬ 1984. Chromosome numbers in the species of the fami ̄
lies Alliaceaeꎬ Asteraceaeꎬ Caryophyllaceaeꎬ Ebenaceaeꎬ Linace ̄
aeꎬ Oleaceaeꎬ Lamiaceae from Tadjikistan [ J ] . Botanicˇhnyi
Žurnal (Moscow & Leningrad)ꎬ 69: 1563—1564
Azizian Dꎬ Cutler DFꎬ 1982. Anatomicalꎬ cytological and phytochem ̄
ical studies on Phlomis L. and Eremostachys Bunge ( Labiatae)
[J] . Botanical Journal of the Linnean Societyꎬ 85: 249—281
Bentham Gꎬ 1829. Notochaete [ J] . Edwards’s Botanical Registerꎬ
15: pl. 1289
Bentham Gꎬ 1832-1836. Labiatarum Genera et Species [M]. London:
James Ridgway & Sons
Boissier Eꎬ 1879. Flora orientalisꎬ Vol 4ꎬ pt 1 [M]. Genevaꎬ Ba ̄
selꎬ Leiden: H. Georg
Briquet Jꎬ 1895 - 1897. Labiatae. Eremostachys Bungeꎬ Phlomis L
[A]. In: Engler Aꎬ Prantl K (eds.)ꎬ Die natürlichen Pflanzen ̄
familein [M]. Leipzig: Engelmannꎬ 4 (3aꎬ 134): 246—249
Brullo Sꎬ Guglielmo Aꎬ Pavone P et al.ꎬ 1990. Chromosome counts of
flowering plants from N. Cyrenaica [J] . Candolleaꎬ 45: 65—74
Bunge Aꎬ 1830. Eremostachys Bunge [A]. In: Ledebour CFꎬ Meyer
CAꎬ Bunge A (eds.)ꎬ Flora Altaica [M]. Berlin: G. Reimerꎬ
2: 414—416
Fang LQ (房丽琴)ꎬ Pang YZ (潘跃芝)ꎬ Gong X (龚洵)ꎬ 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)ꎬ 45: 627—632
Ghaffari SMꎬ 2006. New or rare chromosome counts of some angio ̄
sperm species from Iran [ J] . Iranian Journal of Botanyꎬ 11:
185—192
Kamelin RVꎬ Makhmedov AMꎬ 1990a. The system of the genus Phlo ̄
moides (Lamiaceae) [J] . Botanicˇhnyi Žurnal (Moscow & Len ̄
ingrad)ꎬ 75: 241—250
Kamelin RVꎬ Makhmedov AMꎬ 1990b. A new system of the genus
Phlomis (Lamiaceae) [J] . Botanicˇhnyi Žurnal (Moscow & Len ̄
ingrad)ꎬ 75: 1163—1167
Kudö Yꎬ 1929. Labiatarum Sino ̄Japonicarum prodormus [ A ].
Memoirs of the Faculty of Science and Agricultureꎬ Taihoku Im ̄
perial Universityꎬ 2: 1—332
Mathiesen Cꎬ Scheen ACꎬ Lindqvist Cꎬ 2011. Phylogeny and bioge ̄
ography of the lamioid genus Phlomis ( Lamiaceae) [ J] . Kew
Bulletinꎬ 66: 83—99
Moench Cꎬ 1794. Methodus Plantas Horti Botanici et Argi Marburgen ̄
sisꎬ A Staminum Situ Describendi [M]. Marburg: Officina Nova
Libraria Academiae
Pan YZꎬ Fang LQꎬ Hao G et al.ꎬ 2009. Systematic positions of Lami ̄
ophlomis and Paraphlomis ( Lamiaceae) based on nuclear and
chloroplast sequences [J] . Journal of Systematics and Evolutionꎬ
47: 535—542
Ryding Oꎬ 2008. Pericarp structure and phylogeny of the Phlomis
group (Lamiaceae subfam. Lamioideae) [J] . Botanische Jahrbü ̄
cher für Systematikꎬ Pflanzengeschichte und Pflanzengeographieꎬ
127: 299—316
Salmaki Yꎬ Zarres Sꎬ Heubl Gꎬ 2012a. The genus Phlomoides Moench
(Lamiaceaeꎻ Lamioideaeꎻ Phlomideae) in Iran: an updated syn ̄
opsis [J] . Iranian Journal of Botanyꎬ 18: 207—219
Salmaki Yꎬ Zarre Sꎬ Ryding O et al.ꎬ 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] . Taxonꎬ 61 (1): 161—179
Wu CY (吴征镒)ꎬ Li HW (李锡文)ꎬ 1977. Phlomis L. [A]. In:
Wu CY (吴征镒) ( ed.)ꎬ Flora Reipublicae Popularis Sinica
[M]. Beijing: Science Pressꎬ 65 (2): 428—478
065 植 物 分 类 与 资 源 学 报 第 36卷