全 文 ：植物科学学报 2014，32(4) :329 ～335
Plant Science Journal
DOI:10. 3724 /SP. J. 1142. 2014. 40329
国产唇形科香茶菜属一新记录种———暗红香茶菜
陈亚萍1，2，胡国雄1，2，向春雷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，昆明 650201;2. 中国科学院大学，北京 100049)
摘 要:报道了采自西藏错那县的唇形科香茶菜属的一个中国新记录种，即暗红香茶菜 Isodon atroruber Ｒ. A.
Clement。本研究对该种进行了描述，并应用扫描电子显微镜和光学显微镜对其叶表皮、花粉及小坚果的微形态
特征进行了观察和测量。此外，还编制了西藏产 19 种香茶菜属植物的检索表。
关键词:暗红香茶菜;唇形科;香茶菜亚族;新记录种;中国
中图分类号:Q949. 777. 6 文献标识码:A 文章编号:2095-0837(2014)04-0329-07
收稿日期:2013-11-29，退修日期:2014-01-24。
基金项目:国家自然科学基金(31270245，31110103911) ;中国科学院知识创新工程重要方向性项目(KSCX2-EW-J-24，KSCX2-
EW-Z-1) ;中国科学院植物研究所系统与进化植物学国家重点实验室开放课题(LSEB2012-08)。
作者简介:陈亚萍(1990－ ) ，女，硕士研究生，研究方向为植物分类学(E-mail:chenyaping@mail. kib. ac. cn)。
* 通讯作者(Author for correspondence． E-mail:xiangchunlei@mail. kib. ac. cn)。
Isodon atroruber (Lamiaceae) :ANewly Ｒecorded
Species from China
CHEN Ya-Ping1，2，HU Guo-Xiong1，2，XIANG Chun-Lei1*
(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)
Abstract:Isodon atroruber Ｒ. A. Clement，collected from Cuona County in Tibet， is a new
record for China. The species is described and illustrated here. Scanning electron microscopy
and light microscopy are carried out to examine the micromorphology of the leaf epidermis，
pollen，and mericarp of the species. A key to 19 taxa of Isodon distributed in Tibet is also
provided．
Key words:Isodon atroruber;Lamiaceae;Subtribe Isodoninae;Newly recorded species;Chi-
na
Isodon (Schrad. ex Benth. )Spach (Lami-
aceae: Nepetoideae: Ocimeae ) ，comprising
approximately 100 species，is a morphologically
diverse genus distributed mainly throughout tropi-
cal and subtropical Asia，with two representa-
tives in tropical Africa［1－6］. In China，79 species
and 11 varieties are currently recognized，with
southwest China regarded as the center of mor-
phological diversity of the genus［3，4，7－9］. Isodon
species are mainly shrubs and perennial herbs，
and most are economically or medicinally impor-
tant［1，10］. The genus is characterized by pedun-
culate and bracteolate cymes，equally to sube-
qually 5-toothed or 3 /2-bilabiate calyces，strongly
2-lipped (4 /1)corollas，as well as free filaments
inserted at the base of the corolla tubes［3，11］.
During our preparation of an account for Flo-
ra of Pan-Himalayas，we discovered a herbarium
collection held at the Herbarium of Kunming Insti-
tute of Botany，Chinese Academy of Sciences
(KUN)． It was collected from the Cuona County
of Tibet，China，in August 1975 by the Compre-
hensive Scientific Expedition to the Qinghai-Tibet
Plateau，Chinese Academy of Sciences. This
specimen was identified as Ｒabdosia phulcho-
kiensis Murata［= I. phulchokiensis (Murata)H.
Hara］ in August 1978 by Professor Hsi-Wen Li，
a renowned Chinese expert on the taxonomy of
Lamiaceae. The species was later recorded in
Flora Xizangica［12］. However，a label annotated
as“not Ｒabdosia phulchokiensis，maybe (a)
new species”in September 1982 by Gen Murata，
who first described Ｒ. phulchokiensis［13］，was
later attached to the specimen sheet.
To clarify the identity of this collection，we
undertook a field investigation of Cuona County in
September 2012 where we rediscovered plants of
this species. After careful observations of in situ
plants，comparison with herbarium specimens，
as well as reviewing relevant literature［3，4，12，13］，
we concluded that these Cuona County plants are
Isodon atroruber Ｒ. A. Clement，which was first
reported from Bhutan by Clement in 1993［14］ .
Thus，the species is newly recorded from China.
Detailed characters and micromorphological fea-
tures of the leaf epidermis，pollen grains，and
mericarps are described and illustrated in this
paper，and a key to 19 taxa of Isodon distributed
in Tibet is also provided.
Isodon atroruber Ｒ. A. Clement 暗红香茶
菜(新拟) (Fig. 1，Fig. 2)
Isodon atroruber Ｒ. A. Clement，Edinburgh J.
Bot. 50 (1):33 (1993)． ——— Type:Bhutan，
Thimphu District，Dotena，Thimphu Chu，27°35N，
89°38E，moss bank in Quercus semecarpifolia for-
est，erect herb，30－40 cm，leaves unplea-santly
scented， flowers dark crimson，2550 m，5 ix
1984，Sinclari ＆ Long 4843 (Holotype，E，pho-
to!)． ——— I. pulchokiensis auct. non (Murata)H.
Hara:H. W. Li in Fl. Xizangica 4:213 (1985)．
Herbs perennial，60 － 90 cm tall. Ｒhizomes
woody，massive. Stems erect to ascending，4-
angled，puberulent and pilose. Leaves oppo-
site;petiole (1. 5 －)2. 5 － 6 (－8)cm，puberu-
lent and pilose;lamina 3. 5 －10 × 3－7 cm，pa-
pery，broadly ovate，base broadly cuneate to
truncate，margin crenate，apex acute to acumi-
nate，adaxially olive green， sparsely hirsute，
abaxially greenish，hirsute and glandular puber-
ulent on veins，sparsely red-brown glandular on
both sides. Inflorescence of narrow panicles，
terminal and axillary，3－ 8 × 0. 5 －1 cm;cymes
secund，(3 －)5 －7-flowered，densely puberu-
lent，glandular，pedunculate;floral leaves bract-
like，broadly ovate;bracteoles linear，ca. 1 mm;
peduncle and pedicel ca. 3－5 mm. Calyx cam-
panulate，ca. 1. 5 mm，2-lipped to its 1 /3 length，
densely red-brown glandular， 15-veined， pu-
berulent on veins;teeth broadly triangular，apex
obtuse;fruiting calyx elongated，ca. 3 mm，tu-
bular-campanulate，slightly curved. Corolla dark
brownish purple， densely red-brown glandular
outside，tube short，included in calyx;2-lipped，
posterior lip 4-cleft，much recurved， ca. 2 ×
4 mm，with dark spots on middle two lobes，
anterior lip ca. 3. 5 mm， entire， concave，
much recurved. Stamens and style exserted.
Mericarps brownish， elliptic， ca. 1 mm， gla-
brous. Fl. Aug － Sep，fr. Sep － Oct.
Taxonomic notes:This species is morpho-
logically similar to I. lophanthoides (Buch. -Ham.
ex D. Don)H. Hara，but differs in panicles nar-
row (vs. spreading in I. lophanthoides) ，cymes
secund and (3 －)5 － 7-flowered (vs. not se-
cund，11 －13-flowered I. lophanthoides) ，frui-
ting calyx ca. 3 mm (vs. 4. 5－5 mm I. lophan-
thoides) ，corolla dark brownish purple (vs.
white or pink I. lophanthoides) ，and corolla tube
1. 5 mm (vs. 3. 7－5 mm I. lophanthoides)．
Leaf epidermis morphology: Fig. 3:A － C
shows the indumentum on both sides and stomas
on the abaxial epidermis of the leaf of I. atroru-
ber. Two kinds of trichomes can be recognized:
glandular and eglandular. Morphologically，eg-
landular trichomes are multicellular and distribute
033 植 物 科 学 学 报 第 32 卷
A. Habit;B. Leaf;C. Part of a narrow panicle;D. Flower in lateral view;E. Flower in frontal view;F. Calyx;G. Pistil;H. Mericarp．
Fig. 1 Isodon atroruber Ｒ. A. Clement
133第 4 期 陈亚萍等:国产唇形科香茶菜属一新记录种———暗红香茶菜(英文)
A. Habitat;B. Habit;C. Leaves;D. Narrow panicle in frontal view;E. Narrow panicle in lateral view．
Fig. 2 Photographs of Isodon atroruber in the wild
sparsely on the adaxial surface，and along veins
of the abaxial side (Fig. 3:A，C)． Both peltate
and capitate glandular trichomes are present.
The former mainly along veins on the abaxial side
(Fig. 3:A) ，whereas the latter spread sparsely
on both leaf surfaces (Fig. 3:A－C)． Stomatal
apparatuses are arranged irregularly in the aba-
xial epidermis (Fig. 3:B)．
Pollen morphology:Fig. 3:D and E demon-
strate the pollen grains of I. atroruber. From the
polar view (Fig. 3:D) ，I. atroruber is charac-
terized with hexacolpate pollen grains，a shared
morphology of species from subfamily Nepe-
toideae［15，16］. The pollen is oblate with polar axis
(P)= 40. 4 ± 2. 25 μm，equatorial axis (E)=
45. 5 ± 1. 90 μm，and P/E = 0. 887 ± 0. 025.
The exine sculpturing is incomplete reticulate and
perforated (Fig. 3:E)．
Mericarp morphology:The mericarp of I. at-
roruber is shown in Fig. 3:F. The surface orna-
mentation of the mericarp is papillate. Length
(L)= 1. 17 ± 0. 055 mm，width (W)= 0. 74 ±
0. 040 mm，the shape is elliptic.
Distribution and habitat:Bhutan，China (Ti-
bet)． It grows mostly on moist cliffs or on the edge
of forests at an elevation of about 2800 m，together
with Anaphalis sp．，Galium sp．，Hedera ne-
palensis var. sinensis (Tobler)Ｒehder，Memo-
rialis hirta (Blume ex Hassk. )Wedd．，Ophio-
pogon sp．，Polypodiodes sp．，Ｒhododendron
sp．，Synotis alata (Wall. ex DC. )C. Jeffrey ＆
Y. L. Chen，Urtica sp．，etc. In China，Isodon
233 植 物 科 学 学 报 第 32 卷
A. Adaxial surface of leaf;B. Glands and stomata on adaxial epidermis of leaf;C. Abaxial surface of leaf;
D. Whole pollen，equatorial view;E. Exine surface of pollen;F. Whole mericarp．
Fig. 3 Micromorphology of leaf epidermis，pollen grain，and mericarp of Isodon atroruber
using scanning electron microscopy
atroruber is known only from one location in Cuona
County，Tibet，and considered as a rare species
before additional localities for the taxon are found．
Specimens examined:Tibet (西藏) ，Cuona
(错那) ，Le Township， near the Hydropower
Station (勒乡水电站附近) ，alt. 2850 m，27°49
34. 25″N，91°4332. 42″E，on the edge of forest，
3 Sep 2012，G. X. Hu，X. C. Zhang ＆ Z. X.
Fu. 1209005 (KUN) ;Tibet (西藏) ，Cuona (错
那) ，on the way from Mama Township to
Chachang (麻玛乡至茶场途中) ，alt. 2800 m，
on the edge of forest，26 Aug 1975，Qinghai-Ti-
bet Plateau Expedition，Chinese Academy of Sci-
ences 751694 (KUN)．
An identification key for the 19 taxa of Isodon
distributed in Tibet is here provided as follows.
333第 4 期 陈亚萍等:国产唇形科香茶菜属一新记录种———暗红香茶菜(英文)
Key to the Taxa of Isodon from Tibet，China
1. Corolla 1. 2－1. 5 cm，tube 2 × or more longer than limb 2…………………………………………………………………
2. Calyx purplish，lamina sparsely hirtellous or puberulent I. irroratus………………………………………………………
2. Calyx not tinged purplish，lamina glabrous I. wardii………………………………………………………………………
1. Corolla shorter than 1. 2 cm，tube 2 × or less longer than limb 3…………………………………………………………
3. Calyx with 5 equal or subequal teeth or inconspicuously 2-lipped 4………………………………………………………
4. Lamina and calyx red-brown glandular 5…………………………………………………………………………………
5. Corolla dark brown-purple;cymes in narrow panicles，secund I. atroruber………………………………………
5. Corolla white or pink;cymes in spreading panicles，not secund 6…………………………………………………
6. Calyx densely villous，lamina margin serrate I. hispidus…………………………………………………………
6. Calyx sparsely villous basally，lamina margin crenate 7……………………………………………………………
7. Plant 0. 5 － 1 m tall;lamina 1. 5 － 8. 8 × 0. 5 － 5. 3 mm，apex obtuse I. lophanthoides var. lophanthoides…
7. Plant to 1. 5 m tall;lamina 20 × 8. 5 mm，apex acuminate I. lophanthoides var. graciliflorus……………
4. Lamina and calyx not red-brown glandular 8……………………………………………………………………………
8. Branchlet，leaf，and calyx variously dense hairy;lamina mostly corrugate 9……………………………………
9. Branchlet，leaf，and calyx densely stellate tomentose 10…………………………………………………………
10. Calyx teeth ca. 1 /3 as long as tube I. rugosus………………………………………………………………
10. Calyx teeth ca. 1 /2 as long as tube 11…………………………………………………………………………
11. Lamina 3 － 8 cm I. grandifolia var. atuntzensis……………………………………………………………
11. Lamina 0. 7 － 1. 5 cm I. oresbius………………………………………………………………………………
9. Branchlet，leaf，and calyx with different hairs than above 12……………………………………………………
12. Lamina gray abaxially，conspicuously discolored 13…………………………………………………………
13. Lamina 2. 5 －4. 4 mm I. smithianus……………………………………………………………………………
13. Lamina shorter than 2 mm I. parvifolius………………………………………………………………………
12. Lamina pale green abaxially，not discolored I. loxothyrsus…………………………………………………
8. Branchlet，leaf，and calyx sparsely hairy;lamina usually not corrugate 14………………………………………
14. Stem leaf usually shorter than 2 cm I. pharicus……………………………………………………………………
14. Stem leaf longer than 2 cm 15………………………………………………………………………………………
15. Branchlet and petiole densely gray pubescent I. silvaticus……………………………………………………
15. Branchlet and petiole densely glandular pubescent I. pleiophyllus…………………………………………
3. Calyx conspicuously 2-lipped 16………………………………………………………………………………………………
16. Calyx lips extending to middle of calyx 17………………………………………………………………………………
17. Cymes in narrow panicles，corolla purple I. coetsa…………………………………………………………………
17. Cymes in spreading panicles，corolla yellow I. sculponeatus……………………………………………………
16. Calyx lips extending below middle of calyx 18…………………………………………………………………………
18. Corolla whitish yellow，stamens and style long exserted I. scrophularioides……………………………………
18. Corolla white，stamens and style included I. weisiensis……………………………………………………………
Acknowledgements:We would like to express our
warm thanks to Ms. Ling Wang (KUN)for her excellent
drawing，Dr. Tetsuo Ohi-Toma (TI)for providing images
of the type specimens of I. phulchokiensis，Dr. Barry J.
Conn (NSW)for his great help in editing the manuscript.
Thanks are also due to the staff of the following herbaria
for allowing us to study their Isodon specimens:CDBI，
E，IBK，IBSC，K，KUN，L，LE，PE，TI，and W.
Ｒeferences:
［1］ Wu CY，Li HW. Ｒabdosia (Bl. )Hassk［M］∥Wu
CY ed. Flora Ｒeipublicae Popularis Sinicae. Bei-
jing:Science Press，1977，66:416－534．
433 植 物 科 学 学 报 第 32 卷
［2］ Codd LE. The genus Isodon (Schard. ex
Benth. )Spach in Africa and a new genus Ｒab-
dosiella Codd (Lamiaceae) ［J］． Bothalia，1984，
15 (1－2) :7－10．
［3］ Li HW. Taxonomic review of Isodon (Labiatae)
［J］． J Arnold Arbor，1988，69 (4) :289－400．
［4］ Li HW，Hedge IC. Lamiaceae［M］∥Wu CY，Ｒa-
ven PH，eds. Flora of China. Beijing:Science
Press，Saint Louis:Missouri Botanical Garden
Press，1994，17:269－291．
［5］ Paton AJ，Springate D，Suddee S，Otieno D，
Grayer ＲJ， Harley MM，Willis F， Simmonds
MSJ，Powell MP，Savolainen V. Phylogeny and
evolution of basils and allies (Ocimeae，Labia-
tae)based on three plastid DNA regions［J］． Mol
Phylogenet Evol，2004，31 (1) :277－299．
［6］ Mabberley DJ. Mabberleys Plant-book:A Porta-
ble Dictionary of the Plants，Their Classifications，
and Uses［M］． Cambridge:Cambridge University
Press，2008．
［7］ Paton AJ， Ｒyding O. Hanceola， Siphocranion
and Isodon and their position in the Ocimeae (La-
biatae) ［J］． Kew Bull，1998，53 (3) :723－731．
［8］ Zhong JS，Li J，Li L，Conran JH，Li HW. Phy-
logeny of Isodon (Schard. ex Benth. ) Spach
(Lamiaceae)and related genera inferred from nu-
clear ribosomal ITS，trnL-trnF region，and rps16
intron sequences and morphology［J］． Syst Bot，
2010，35 (1) :207－219．
［9］ Xiang CL，Liu ED. A new species of Isodon
(Lamiaceae，Nepetoideae) from Yunnan Prov-
ince，Southwest China［J］． Syst Bot，2012，37
(3) :811－817．
［10］ Sun HD，Xu YL，Jiang B. Diterpenoids from
Iso-don species［M］． Beijing:Science Press，
2001．
［11］ Harley ＲM，Atkins S，Budantsev AL，Cantino PD，
Conn BJ，Grayer Ｒ，Harley MM，De-Kok Ｒ，
Krestovskaja T，Morales Ｒ，Paton AJ，Ｒyding O，
Upson T. Labiatae［M］∥Kubitzki K ed. The Fami-
lies and Genera of Vascular Plants. Berlin:
Springer，2004，7:167－275．
［12］ Li HW. Labiatae［M］∥Wu CY ed. Flora Xizangi-
ca. Beijing:Science Press，1983，4:98－224．
［13］ Murata G. Labiatae［M］∥Ohashi H ed. The Flora
of Eastern Himalaya. Tokyo:University of Tokyo
Press，1975，3:91－98．
［14］ Clement ＲA. Notes relating to the flora of Bhutan:
ⅩⅧ. New taxa and a new combination in the Labia-
tae［J］． Edinburgh J Bot，1993，50 (1):33－38．
［15］ Erdtman G. Pollen morphology and plant taxono-
my. IV. Labiatae，Verbenaceae and Avicenniace-
ae［J］． Svensk Bot Tidskr，1945，39 (3) :279－
285．
［16］ Cantino PD，Sanders ＲW. Subfamilial classifica-
tion of Labiatae［J］． Syst Bot，1986，11 (1) :
163－185．
(责任编辑:张 平)
533第 4 期 陈亚萍等:国产唇形科香茶菜属一新记录种———暗红香茶菜(英文)