植 物 分 类 学 报 45 (6): 841–848(2007)
Acta Phytotaxonomica Sinica
doi:10.1360/aps07030
http://www.plantsystematics.com
Two newly recorded species of Indigofera
(Fabaceae) in China
GAO Xin-Fen*
(Chengdu Institute of Biology, the Chinese Academy of Sciences, Chengdu 610041, China)
Abstract This paper reports two newly recorded species from China, namely Indigofera
caloneura with 1-foliolate leaves from Yunnan and I. cordifolia with simple leaves from
Guangdong. There are another three species with simple leaves of Indigofera in China.
Based on the microscopic observation of specimens, morphological characters which include
habit, stem, leaf shape and size, indumentum, stipule and stipel, raceme, corolla color,
standard shape and indumentum, anther, pod shape and size, and seed number per pod etc. in
the five species are described and compared among one another. The five most important
characters are chosen to discuss for taxonomic treatment.
Key words Indigofera, Indigofera caloneura, Indigofera cordifolia, Fabaceae, new record,
Guangdong, Yunnan, China.
The genus Indigofera L. of the legume family (Fabaceae) is widely distributed in tropical
and subtropical regions. In the most recent treatment 81 species are distributed in China (Fang
& Zheng, 1994), of which three have simple or 1-foliolate leaves. For Southeast Asia there
are totally eight species with simple or 1-foliolate leaves recorded, namely I. brunoniana
Grah. ex Wall., I. caloneura Kurz, I. cordifolia Heyne ex Roth., I. linifolia (L. f.) Retz., I.
maymyoensis Sanjappa, I. minbuensis Gage, I. nummularifolia (L.) Livera ex Alston, and I.
squalida Prain (de Kort & Thijsse, 1984; Sanjappa, 1985). Among them, I. linifolia, I.
nummularifolia, and I. squalida have been reported in China before (Fang & Zheng, 1994).
When studying this genus for Flora of China, a specimen from Xishuangbanna of Yunnan and
two additional ones from Guangdong with simple leaves or 1-foliolate leaves are found
belonging to I. caloneura and I. cordifolia respectively. These two species have not been
recorded in China before. In order to easily distinguish these species, morphological
characters in habit, stem, leaf size and shape, leaf glands, indumentum, petiole, stipule, stipel,
inflorescence length, flower size and hairs, and fruit shape are observed.
1
Material and methods
Morphological features are based on more than 50 herbarium specimens from CDBI,
HITBC, IBSC, KUN, and PE. Microscopic characters were obtained by using stereo
microscope (Zeiss Stemi 2000, Germany). Each species was represented by more than one
specimens in the microscopic observations. The materials used for microscopic study are
listed in Table 1.
2
Results and discussion
Main morphological characters observed are listed in Table 2. The five most important
characters are discussed below.
———————————
Received: 27 February 2007 Accepted: 26 June 2007
Supported by the National Natural Science Foundation of China, Grant Nos. 30499340, 39899400; and the International
Cooperation Fundation of the Chinese Academy of Sciences, Grant No. KSCX-SW-122.
* E-mail: <xfgao@cib.ac.cn>.
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Table 1
Acta Phytotaxonomica Sinica
Vol. 45
Materials used for microscopic study and their voucher information
Species name
Indigofera caloneura
I. cordifolia
I. linifolia
I. nummularifolia
I. squalida
Locality
Mengla, Yunnan (云南勐腊)
Lian Xian, Guangdong (广东连县)
Huili, Sichuan (四 会理)
Ya Xian, Hainan (海南崖县)
Ya Xian, Hainan (海南崖县)
Anlong, Guizhou (贵 安龙)
Menglian, Yunnan (云南孟连)
Voucher
Exp. Team (考察队) 35312
Veget. Exp. (山地植被组) s.n.
T. T. Yu (俞德浚) 1633
C. Huang (黄志) 34814
K. Z. Hou (侯宽昭) 73843
G. S. Liao (廖国胜) 1468
Menglian Exp. (孟连队) 9425
Figure number
1A, 1B, 14
2A, 2B, 6, 10, 15
3A, 3B, 7, 11
4A, 4B, 8
12
5A, 5B, 9
13
Table 2 Main morphological characters of Indigofera species with simple and 1-foliolate leaves in China
Characters
I. caloneura
I. cordifolia
I. linifolia
I. nummularifolia
I. squalida
Root
straight
straight
straight
straight
tuberous
Habit
shrub
perennial herb
shrublet or herb
perennial herb
perennial herb
Stem
erect
procumbent
erect or prostrate prostrate
erect
Leaf size (mm)
80–100×40–55
4–8×3–6
5–30×2–4
10–20×7–10
25–55×7–20
Leaf or leaflet
oblong or
cordate
linear, narrowly
obovate-elliptic or
narrowly
shape
elliptic
obovate or
obovate
elliptic to
elliptic
obovate
Hair type
spreading
spreading
adpressed
adpressed
adpressed
sub-basifixed
subbasifixed to
medifixed
medifixed
medifixed
biramous
medifixed
biramous
biramous
biramous
biramous
Glands on leaf
absent
absent
absent
dotted
disc-shape
beneath
Lateral nerves
15–20
invisible
invisible
3–5
10–11
(pairs)
Stipel (mm)
1.5–2
absent
absent
absent
absent
Racemes length
7–13,
0.2–0.3,
1–1.5,
1.5–2
1–2, densely
(cm)
multi-flowers
1–4-flowers
multi-flowers
multi-flowers
flowered
Flower colour
white or yellow
bright red
red
red
red or purple
brown to
orange-yellow
Flower length
6–10
2.5–3
4–5
3
4–5
(mm)
Standard shape
orbicular
spathulate
elliptic
broadly ovate
elliptic
Standard hairs
densely, brown,
sparsely, on the
densely, white,
sparsely, on the
densely,
outside
medifixed
apex part, white,
medifixed
apex part, white,
brown,
sub-basifixed
medifixed
medifixed
Keel hairs
margin ciliate,
margin ciliate
glabrous
margin ciliate
densely
densely outside
outside,
medifixed
Anther hairs
at the base
glabrous
glabrous
glabrous
glabrous
Fruit shape
linearly
ellipsoid or
ovoid
falcate, echinated
linearly
cylindric
subglobose
cylindric
Fruit size (mm)
20–50×2–2.5
4–5×1.5
2×1.5
7×4
11–14×1.5–2
Fruit hairs
adpressed,
spreading, white
adpressed
sparse, with
densely
medifixed
sub-basifixed
medifixed
sub-basifixed hairs medifixed
biramous hairs
biramous hairs
biramous hairs
biramous hairs
Seed number
8–11
1 or 2
1
1
3–5
(1) Habit. The five species with simple or 1-foliolate leaves are usually perennial herbs
or small shrublets with erect or procumbent stem except for I. caloneura which is erect and
tall shrub.
(2) Hairs. Hair morphology is an important identification character in this genus (de
Kort & Thijsse, 1984; Schrire, 1995). Biramous hairs have different fixed points with from
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Figs. 1–5. The leaf surfaces of Indigofera. 1. I. caloneura, biramous hairs with very unequally arms. 2. I. cordifolia. 3.
I. linifolia, biramous hairs with equally arms. 4. I. nummularifolia, with brown dotted glands and biramous hairs vertically
along the margin (B). 5. I. squalida, with disc-shaped glands (B) and medifixed hairs. A, upper surface; B, lower surface.
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Figs. 6–9. The dorsal surfaces of standards in Indigofera. 6. I. cordifolia. 7. I. linifolia. 8. I. nummularifolia. 9. I.
squalida. Figs. 10–13. Pods of Indigofera. 10. I. cordifolia. 11. I. linifolia. 12. I. nummularifolia. 13. I. squalida.
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equally to extremely unequally long arms on the two sides of the fixed point (Figs. 1B–5B).
This feature has long been noticed and used as one of the main identification characters at
specific level in the genus (de Kort & Thijsse, 1984; Sanjappa, 1995; Schrire, 1994).
However, it was only rarely well-described in the past (Craib, 1913; de Kort & Thijsse, 1984;
Fang & Zheng, 1994). For the species with simple or 1-foliolate leaves in China, hairs are
either adpressed (Figs. 3A–5B) or spreading (Figs. 1B, 2B), and either sub-basifixed point
with extremely unequally long arms (Figs. 1B, 2A) or medifixed point with more or less
equally long arms (Figs. 2B–5B).
(3) Leaves. Leaf features (shape, size, and indumentum) are very different in the five
species. Indigofera caloneura has large oblong or elliptic leaves with spreading sub-basifixed
biramous hairs beneath (Fig. 1B) and glabrous above (Fig. 1A), I. cordifolia has small and
cordate leaves with spreading biramous hairs sub-basifixed above (Fig. 2A) and sub-medifixed
beneath (Fig. 2B), I. linifolia has linear or narrowly elliptic leaves with medifixed biramous
hairs on both surfaces (Figs. 3A, 3B), I. nummularifolia leaves obovate with medifixed
biramous hairs along the margin on both surfaces (Figs. 4A, 4B) and dotted glands beneath
(Fig. 4B), and I. squalida leaves elliptic with sparsely medifixed biramous hairs on both
surfaces (Figs. 5A, 5B) and disc-shape glands beneath (Fig. 5B).
(4) Corollas. Corolla morphology (color, size, shape, and indumentum) is valuable for
species identification. Corollas are usually red to purple except for that in I. caloneura which
is whitish, yellow-brown to orange-yellow. Standards have very different shapes. They are
spathulate in I. cordifolia, elliptic in I. linifolia and I. squalida, orbicular in I. caloneura, and
broadly ovate in I. nummularifolia. Standards are usually covered with different hairs. Those
of I. linifolia and I. squalida are covered with adpressed biramous hairs with medifixed point
on the dorsal surface (Figs. 7, 9), while those of I. cordifolia and I. nummularifolia have
biramous hairs with sub-basifixed point only on the dorsal surface of the apex part (Figs. 6,
8). Wings are usually glabrous, sometimes with ciliate hairs. Keels only in I. caloneura and I.
squalida are densely covered with hairs, but those in other species are glabrous or only with
ciliate hairs.
(5) Fruits. Fruit shape is very different in the five species. Falcate pods are found in I.
nummularifolia only (Fig. 12), while I. linifolia has globose pods (Fig. 11), I. cordifolia
subglobose pods (Fig. 10), and I. caloneura and I. squalida linearly cylindrical pods (Figs. 13,
14). They are easy to be identified.
3
Taxonomical treatment
The five species with simple and 1-foliolate leaves in China and additional three species
in Southeast Asia belong to different sections, are distinguished as follows:
1. Leaves with minute and dot-shaped glands beneath, medifixed biramous hairs vertically along margins on
both surfaces (Fig. 4B); standard with sub-basifixed biramous hairs towards the apex outside (Fig. 8); pods
falcate, echinated along dorsal suture (Fig. 12).................................................................. I. nummularifolia
1. Leaves without glands or with disc-shaped glands beneath, medifixed biramous hairs not vertically along
margins; pods linearly cylindric, globose or subglobose.
2. The main root tuberous; leaves obovate, with medifixed biramous hairs on both surfaces, and sometimes
sparsely with disc-shaped glands on lower surface (Fig. 5B); pods straight, linearly cylindric (Fig.
13............................…………......... ......................…………..................................................... I. squalida
2. The main root not tuberous; leaves linear, narrowly obovate-lanceolate or elliptic, oblong or cordate,
without any glands; pods linearly cylindric, globose or subglobose.
3. Leaves linear, narrowly obovate-lanceolate, or narrowly elliptic, with adpressed medifixed biramous
hairs on both surfaces (Fig. 3A, 3B); stipels absent.
4. Leaves 10–35 mm long......................................................................................................... I. linifolia
4. Leaves 40–80 mm long.................................................................................................. I. minbuensis
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3. Leaves oblong or cordate; stipels present or absent.
5. Stipels absent; leaves cordate, with spreading sub-basifixed biramous hairs above and sub-basifixed
to medifixed biramous hairs beneath; flower 2.5–3 mm long; fruit elliptic or subglobose; seeds 2
per fruit...............................................................................................................................I. cordifolia
5. Stipels present; leaflets oblong or elliptic, with sub-basifixed biramous hairs; seeds numerous.
6. Leaf glabrous above, biramous stipules 3–5 mm long.
7. Leaf with adpressed medifixed biramous hairs on lower surface; petioles 2–3 cm long;
flowers sparse in racemes which are shorter than leaves ............................... I. maymyoensis
7. Leaf with spreading sub-basifixed biramous hairs on the lower surface; petioles 0.7–1 cm
long; flowers compact in racemes which are longer than leaves.......................... I. caloneura
6. Leaf with adpressed medifixed biramous hairs on both surfaces; stipules 1 mm long; wings
short-ciliate; keels ciliate and with hairs on dorsal surface of apex ..................... I. brunoniana
The two new records are described here in detail.
1. Indigofera caloneura Kurz in J. Asiat. Soc. Bengal 42: 229. 1873; et in Fl. Brit. Burm. 1:
360. 1877; Baker in Hook. f., Fl. Brit. Ind. 2: 93. 1876; de Kort & Thijsse in Blumea 30: 112.
1984; Sanjappa in Reinwardtia 10 (2): 218. 1985. Type: Burma. Pegu, Kurz s.n. (CAL).
I. oblonga Craib in Kew Bull. 6. 1914; Gagnep., Fl. Gen. Indo-Chine. 2: 440, 1916.
美脉木蓝 新拟 Figs. 1A, 1B, 14
[Sect. Indigofera] Erect shrubs; branches angular, densely adpressed brown subbasifixed biramous hairs. Stipules narrowly triangular, 4–5 mm long. Petioles 10–15 mm long.
1-foliolate, lamina 8–10 cm long and 4–5.5 cm broad, oblong or elliptic, obtuse or rounded at
base and apex; dark green and glabrous above; spreading gray or brown sub-basifixed
biramous hairs on lower surface; secondary nerves pinnate, 15–17 pairs; stipels 2, linearly
subulate, 1.5–2 mm long. Racemes 7–13 cm long, compactly many-flowered; bracts 3.5–4
Fig. 14. Specimens of Indigofera caloneura.
Fig. 15. Specimens of Indigofera cordifolia.
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847
mm long, setaceous, deciduous; pedicels 1–1.5 mm long, 3–5 mm long in fruiting, densely
tomentose. Flowers pink, 8–10 mm long; calyx 2–2.5 mm long, tube short, teeth triangular,
acute; standard 8–10 mm long, ovate, obtuse and mucronulate at apex, adpressed densely
brown pubescent on the dorsal surface; wings 6–8 mm long, ciliate; keels 7–9 mm long, with
thickly brown pubescent on dorsal surface; anthers with hairs at the base; ovary glabrous. Pod
straight, cylindrical, 4–5.3 cm long, spreading, adpressed minute hairs, endocarp red-spotted.
Seeds 8–11, dark brown. Fl. May–Aug. Fr. Nov.–Jan.
China. S Yunnan (云南南部): Mengla (勐腊), 101°41′ E, 21°37′ N, alt. 900 m,
1983-12-07, Exped. Team (考察队) 35312 (HITBC!).
Myanmar. Shan hills, C. B. Collett 706 (K!); Geo. B. Vogt. BU-443, (K!); Amherst, J.
H. Lace 5668 (K!).
Thailand. Doi Nang Keo, Kerr 2524 (BM!, K!); Pahem Park, Kerr 5171 (BM!, K!); Doi
Chiengdao, Kerr 4457(BM!); Bo laung, K. Larsen et al. 1912 (K!).
Distribution: S. China, Laos, Myanmar, Thailand, and Vietnam.
2. Indigofera cordifolia Heyne ex Roth, Nov. Plant. Sp. Ind. Orient. 357. 1821; Baker in
Hook., Fl. Brit. Ind. 2: 93. 1876; Ali in Bot. Not. 3: 545. 1958; de Kort & Thijsse in Blumea
30: 116. 1984. Type: India. Or., Heyne s.n. (B)
心叶木蓝 新拟 Figs. 2A, 2B, 6, 10, 15
[Sect. Brachycapae] Herbs, 10–30 cm tall. Branches procumbent or ascending, adpressed
to slightly spreading white sub-basifixed biramous hairs. Leaves simple, petiole 1 mm long;
stipules lanceolate, spreading sub-basifixed biramous hairs outside, 2 mm long, apex
cuspidate; lamina 4–8×3–6 mm, cordate, apex acute, mucro 0.2 mm long; spreading white
sub-basifixed biramous long hairs on upper surface, densely spreading sub-medifixed
biramous long hairs on lower surface; lateral nerves invisible. Racemes 2–3 mm long with 1–4
flowers in the leaf axils; bracts narrowly triangular, 2×0.5 mm, apex cuspidate; pedicels 0.5
mm long or sessile. Calyx with densely spreading white sub-basifixed long hairs, tube 0.5 mm
long, teeth linear-triangular, 1.5–2 mm long; standard spathulate or cuneate, 2.5–3×1–1.5
mm, hairy on dorsal surface of apex, apex obtuse; wings 2.5×0.2 mm, glabrous; keels
2.5×0.5 mm, glabrous, lateral spur 0.25 mm long; Anthers 0.2×0.25 mm, glabrous; ovary
spreading white long hairy; ovules usually 2, rarely 1. Pod ellipsoid or subglobose, 2.5–3×1.6
mm, densely subspreading white sub-basifixed biramous long hairs. seeds 2 or 1, red brown.
Fl. Jul. –Sept. Fr. Sept.–Nov.
China. Guangdong (广东): Lian Xian (连县), Xingzi Qu (星子区), Baitangchong (柏塘
冲), alt. 160 m, 1984–10–20, H. X. Ao (敖惠修) 8401 (IBSC!), the same place, open slope and
exposed hills, 1987–10–23, Vegetation Team (山地植被组) s.n. (IBSC!).
Distribution: S. China, India, Java, Pakistan, Roti, Sawu, Sumba, Timor, Tropical Africa,
and N. Australia.
Acknowledgements I am grateful to the curators of HITBC, IBK, IBSC, KUN, PE for the
loan of specimens, to the staffs of BM, E, K, KUN, LINN, PE, and SZ for their cordially
assistance when I studied specimens in herbaria, to Dr. Sun Hang (Kunming Institute of
Botany, CAS) for his help with literatures, to Dr. B. Schrire (Royal Botanic Garden, Kew,
UK) for his help when I visited at Kew, to Dr. Zhang Libing (Missouri Botanical Garden,
USA) for correcting the manuscript.
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中国木蓝属两个分布新记录
高信芬*
(中国科学院成都生物研究
成都 610041)
摘要 报道并描述了采自云南南部西双版纳具有1小叶的新分布种美脉木蓝Indigofera caloneura Kurz
和采自广东具有单叶的新分布种心叶木蓝I. cordifolia Heyne ex Roth. 在中国还有另外3个具有单叶的
种类, 即单叶木蓝I. linifolia 刺荚木蓝I. nummularifolia和 志木蓝I. squalida 通过从植物习性 茎
叶的形态与大小及毛被 托叶 花序 花冠形状与毛被 花药 子 毛被 果实的形态与大小
含
种子数目等特征的显微观察, 比较了国产单叶类木蓝属Indigofera植物的形态学特征差异, 并针对茎
叶 毛被 花冠旗瓣和果实形状等5个最重要的特征 行了讨论
关键词 木蓝属; 美脉木蓝; 心叶木蓝; 豆科; 新记录; 广东; 云南; 中国