全 文 ：龙胆科獐牙菜属新分类纲要*
何廷农 , 刘尚武, 陈世龙**
(中国科学院高原生物适应与进化重点实验室, 中国科学院西北高原生物研究所, 青海 西宁摇 810001)
摘要: 獐牙菜属是龙胆科中的一个大属, 广泛分布于亚洲、 北美洲、 欧洲和非洲。 本文报道獐牙菜属下的
两个新组 ( sect. Montana 和 sect. Echinulata) 和七个新系 ( ser. Repentes, ser. Kilimandscharicae, ser.
Corymbosae, ser. Japonicae, ser. Swertopsis, ser. Pumilae和 ser. Abyssinicae)。 同时对獐牙菜属中的一些类群
进行了分类修订, 本分类纲要将被獐牙菜属世界专著采用。
关键词: 分类纲要; 獐牙菜属; 龙胆科
中图分类号: Q 949摇 摇 摇 摇 摇 摇 摇 文献标识码: A摇 摇 摇 摇 摇 摇 摇 文章编号: 2095-0845(2013)03-386-07
Nomenclatural Novelities in Swertia (Gentianaceae)
HE Ting鄄Nong , LIU Shang鄄Wu, CHEN Shi鄄Long**
(Key Laboratory of Adaptation and Evolution of Plateau Biota, Northwest Institute of Plateau Biology,
Chinese Academy of Sciences, Xining 810001, China)
Abstract: Swertia is one large genus of Gentianaceae with a wide distribution in Asia, North America, Europe and
Africa. In this study, we reported two new sections (Montana and Echinulata) and seven new series (Repentes,
Kilimandscharicae, Corymbosae, Japonicae, Swertopsis, Pumilae, and Abyssinicae). In addition, some species were
reduced, recombined and newly established. These new taxonomic synopses will be used for a further worldwide
monograph of this genus.
Key words: Taxonomic synopses; Swertia; Gentianaceae
摇 Swertia is one of the large genera in the family
Gentianaceae (Struwe and Albert, 2002). This ge鄄
nus occurs from Asia, Europe to Africa and North
America (He et al., 1994). In the past century,
numerous new species were described under this ge鄄
nus or transferred from the other genera especially
from China (He and Liu, 1980; He, 1988, 1994;
He and Pringle, 1995). The circumscription of this
genus is highly difficult because all phylogenetic ana鄄
lyses based on molecular data suggested that this ge鄄
nus is not monophyletic and some of the paralleling
clades from the radiative diversification of some spe鄄
cies clustered with the multiple related genera
(Yuan and K俟pfer, 1995; Chassot et al., 2001;
Liu et al., 2001; Hagen and Kadereit, 2002). This
suggested that the traits, the rotate corolla and
fringed nectaries, which were used to circumscribe
the genus, may have derived from homoplastic evo鄄
lutions due to the radiative diversifications or repre鄄
sented the ancestral states of the total family (Hagen
and Kadereit, 2002). As inferred from molecular
phylogenies ( e. g. Yuan and K俟pfer, 1995) and
embryological studies (Xue et al., 1999a, b), we
found that sect. Heteranthos of this genus can be ex鄄
植 物 分 类 与 资 源 学 报摇 2013, 35 (3): 386 ~ 392
Plant Diversity and Resources摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 DOI: 10. 7677 / ynzwyj201313041
*
**
Foundation tiems: Key Project of the National Science Foundation of China (31270270) and the National Basic Research Program of China
(2007FY110100)
Author for correspondence; E鄄mail: slchen@ nwipb. ac. cn
Received date: 2013-03-18, Accepted date: 2013-03-25
作者简介: 何廷农 (1938-2011) 女, 已故中国龙胆科植物分类学专家。
cluded from the genus and elevated to a generic sta鄄
tus based on its recognizable morphological distinct鄄
ness: dimorphic plants and flowers and a cleistoga鄄
mous pollination mode. However, up to now, mor鄄
phological traits, which can be used to distinguish
the other paralleling clades, need further examina鄄
tions. Therefore, we tentatively follow the traditional
generic circumscription of Swertia for the further in鄄
frageneric classification. Through examining thou鄄
sands of specimens kept in the major herbaria within
the past three decades, we recognized 168 species
for this genus over the world. We classified these
species into three subgenera, 11 sections and 11 se鄄
ries. Among them, sections Montana and Echinula鄄
ta, and series Repentes, Kilimandscharicae, Corym鄄
bosae, Japonicae, Swertopsis, Pumilae, and Abys鄄
sinicae were newly proposed for the first time. In ad鄄
dition, some taxonomic entities established previous鄄
ly for this genus or related genera were reduced or
recombined. Although the currently circumscribed
genus is still polyphyletic, these infrageneric entities
seem to be monophyletic if assessed only based on
the morphological similarity. Here we report these
new taxonomic synopses established for Swertia.
These taxa will be used for a further worldwide mon鄄
ograph, phylogenetic analyses and resource surveys
related to this genus.
1摇 Subgen. Swertia
1. 1摇 Sect. Apterae V. V. Pissjaukova
1. 1. 1 摇 Swertia cuneata var. asterocalyx (T. N.
He & S. W. Liu) T. N. He & S. W. Liu, comb. nov.
Swertia asterocalyx T. N. He & S. W. Liu, Acta
Phytotax. Sin. 18 (1): 77, f. 2 (1980). Type:
China, Xizang, Nyalam, 3 800 -4 000 m, 5 Sept.
1972, Xizang Medic. Herb. Exped. 1825 (HNWP!
—holotype; PE!—isotype).
Var. asterocalyx is distinguished from var. cu鄄
neata by flowers large: corolla 20-30 mm in diam.,
lobes 15-20 mm. Var. cuneata has small flowers: co鄄
rolla 8-10 mm in diam., lobes 7. 5-10(-12) mm. The
species Swertia cuneata is characterized by calyx lobes
oblong鄄spathulate to elliptic, base narrowed, apex
round and nectaries linear鄄oblong, radially elongated.
Distribution: China (SW Xizang).
2摇 Subgen. Poephila C. B. Clarke
2. 1摇 Sect. Frasera (Walter) Knoblauch
2. 1. 1 摇 Swertia albicaulis var. modocensis ( St.
John) T. N. He, comb. nov.
Swertia modocensis St. John, Amer. Midl. Natu鄄
ralist 26: 18 (1941). Type: United States, Califor鄄
nia, Lassen County, Madeline, gentle northeast fa鄄
cing slope, in rocky volcanic soil, among Artemisia
and scattered Juniperus occidentalis, 4 500 ft., 8
June 1936, E. B. Babcock & G. L. Stebbins 1790
(UC! —holotype; GH! —isotype). Paratypes: Ca鄄
lifornia, Modoc County, in 1907, M. H. Manning
(UC!), Shasta County, 17 May 1923, E. Bethel
(SF, not seen); Oregon, July 1898, R. M. Austin
& Bruce 2153 (UC!), Southern Oregon, between
Merrill and Cottonwood, June 1901, H. W. Furlong,
W. B. Greeley, M. Wilson et al. s. n. (UC!).
Swertia modocensis var. adglabra St. John, A鄄
mer. Midl. Naturalist 26: 19 (1941). Type: Uni鄄
ted States, Oregon, Klamath County, June 1901,
H. W. Furlong, W. B. Greeley, M. Wilson et al.
s. n. (UC! —holotype).
Swertia sierrae St. John, Amer. Midl. Natura鄄
list 26: 16 (1941). Type: United States, Califor鄄
nia, Lassen County, Madeline, about two miles
north of, gentle northeast facing slope, in rocky vol鄄
canic soil, among Artemisia and scattered Juniperus
occidentalis, 4 500 ft., 8 June 1936, E. B. Babcock
& G. L. Stebbins 1790 (GH!, UC! —isotypes).
Paratypes: United States, N California, Willow
Creek, Devils Garden, July 1898, R. M. Austin
418 (US!); Dry Lake, Modoc County, 31 May,
1897, E. I. Applegate 877 (US!); Oregon: Thom鄄
as Creek Ranger Station, Lake County, 1 570 m,
June 29-30, 1911, W. W. Eggleston 5098 (NY!,
US!); Stein 爷s Mountain, 1 June, 1885, J. T.
Howell (US!); near Warm Range, 1 500 m, Lake
County, 26 July, 1896, F. V. Coville & J. B.
7833 期摇 摇 摇 摇 摇 摇 摇 HE Ting鄄Nong et al. : Nomenclatural Novelities in Swertia (Gentianaceae) 摇 摇 摇 摇 摇 摇 摇 摇 摇
Leiberg 53 (US!).
Swertia shastaensis St. John, Amer. Midl. Na鄄
turalist 26: 19 (1941). Type: United States, Cali鄄
fornia, Mount Shasta, near Sheep Rock, 5 000 ft.,
June 1903, H. M. Hall & E. B. Babcock 4113
(UC! —holotype).
Frasera albicaulis var. modocensis (St. John) N.
Holmgren in A. Cronquist et al., Intermountain Fl.,
Vasc. Pl. Intermountain West, U. S. A., 4: 22 (1984).
Swertia modocensis and its variety var. adglabra
are close to species group S. albicaulis, which in鄄
cludes six varieties, in having opposite stem leaves
with white membranous margin, a narrow interrupted
spike鄄like panicle, and oblong boat鄄shaped shortly
fimbriate nectaries. Especially it is more similar to
var. albicaulis and var. cusickii, but differs from
both in corolla pale lavender, with dark purple
streak, and also from the former in coronal scales
entire to shallowly lacerate (in var. albicaulis corol鄄
la pale blue, usually not darker mottled; coronal
scales deeply laciniate to near base into numerous
setae), from the latter in whole plants puberulent
and coronal scales short, 1. 5-2. 5 mm (in var. cu鄄
sickii plants glabrous except for leaves puberulent at
base and sometimes along midvein of abaxial sur鄄
face; coronal scales long, 3 - 4 mm; corolla pure
pale blue). However, the coronal scales ( crown
scales) of S. albicaulis group are deltoid to linea鄄
roblong, margin extremely variable from entire to
deeply laciniate. Hence Swertia modocensis and var.
adglabra are not necessary to maintain as a distinct
species but as a variety.
Distribution: W United States (Oregon and Cali鄄
fornia).
2.2摇 Sect. Montana T. N. He & S.W. Liu, sect. nov.
Herba perennis. Radix leviter carosa usque
semilignosa, vel caulis inferior radices adventitias
emittens. Flores 5鄄, raro 4鄄meri. Nectarium 1 in
unoquoque lobo corollae. Stylus brevis usque ab鄄
sens. Semina longitudinaliter, dense et tenuiter cor鄄
rugata vel corrugato鄄porcata. Pollinis granula admo鄄
dum striata (sparse et tenuiter striata, striato鄄perfora鄄
ta, striato鄄microreticulata usque striato鄄reticulata).
Perennials, with slightly fleshy to semiwoody
tap鄄root and some rootlets, or tap鄄root absent but
stems trailing or stoloniferous with adventitious
roots. Flowers 5鄄, rarely 4鄄merous. Nectary 1 per
corolla lobe. Style short to absent. Seeds either many
corrugate and wingless, with longitudinal, thin and
dense corrugations or many corrugate and ridged,
with irregular narrow membranous reticulate ridges.
Pollen grains are subspheroidal, rarely prolate or sub鄄
prolate; medium鄄, rarely small鄄 or large鄄sized, size
range 23. 3-43. 8 伊19. 0-33. 7 滋m; colpi long or me鄄
dium. The sexine ornamentation is essentially striate,
varying from thinly, undulately striate & perforate to
imperforate. The lira width is 0. 17 - 0. 30 滋m.
Chromosome number 2n=26, rarely 2n=20.
This section is characterized by perennials,
with slightly fleshy to semiwoody tap鄄root or with ad鄄
ventitious roots from nodes of trailing or stoloniferous
stems, nectary 1 per corolla lobe so that it conforms
to the essential features of subgen. Poephila. In this
subgenus it is similar to sect. Poephila, but differs by
having seeds longitudinally densely and minutely cor鄄
rugate鄄 ridged and flowers usually 5鄄merous whereas
the latter having seed coats almost smooth and some鄄
times winged at one or both end, flowers always 4鄄
merous. Pissjaukova (1979 ) put some species of
this section (S. volkensii, S. subnivalis, S. scandens,
S. macrosepala and S. keniensis) in sect. Apterae of
subgen. Swertia. However, sect. Aptera is very dif鄄
ferent from sect. Montana by having an abbreviated
rhizome, 2 nectaries per corolla lobe, and seeds lon鄄
gitudinally few (8 -12) corrugate鄄ridged with high
irregular membranous and reticulate ridges.
Type species: Swertia volkensii Gilg
Distribution: from North Africa to Northeast
and Tropical Africa.
10 species, may be divided into 2 series.
2.2.1摇 ser. Repentes T. N. He & S.W. Liu, ser. nov.
Caulis repens vel stoloniformis, radices adven鄄
titias emittens. Flores solitarii vel in inflorescentiam
parco鄄florigeram dispositi. Nectarium planum, mac鄄
883摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 35 卷
uliforme, absque marginibus et fimbriis.
Stems trailing or stoloniferous with adventitious
roots. Basal leaves crowded into a rosette. Flowers
5鄄merous (but 4鄄merous in S. subnivalis and S. sar鄄
mentosa), solitary on long terminal pedicels or in
few鄄flowered terminal cymes. Nectary reduced into a
flat gland patch without raised margin and fimbriae.
Type species: Swertia volkensii Gilg
Distribution: East and Central Africa.
5 species.
2. 2. 1. 1 摇 Swertia sarmentosa T. N. He & S. W.
Liu, nom. nov.
Swertia scandens T. C. E. Fries, Notizbl. Bot.
Gart. Berlin 8: 516, 3c ( 1923 ), non L佴veill佴
(1913). Syntypes: W Kenya, in rain forest, ca.
2 350 m, 3 Jan. 1922, E. A. Robenson & Th. C.
E. Fries 691 (BM!, K!, S!, UPS!); same locali鄄
ty, 30 Jan. 1922, E. A. Robinson & Th. C. E.
Fries 1277 (K!). Lectotype: Rob. E. & Th. C.
E. Fries 691 (UPS! —lectotype; BM!, K!, S! —
isolectotypes; designated here).
Distribution: East Africa (Kenya).
2. 2. 2摇 Ser. Kilimandscharicae T. N. He & S. W.
Liu, ser. nov.
Monobothrium Hochstetter, Flora 27: 27 (1844).
Type: M. schimperi Hochstetter
Caules erecti, absque stolinibus et radicibus
adventitiis. Cymae in inflorescentiam paniculatam
dispositae. Nectarium V鄄forme, fimbriatum.
Stems erect, without stolons and adventitious
roots. Inflorescences a many鄄flowered panicle of cymes.
Nectary V鄄shaped, fimbriate.
Type species: Swertia kilimandscharica Engler
Distribution: from North Africa to Northeast
and Tropical Africa.
5 species.
3摇 Subgen. Ophelia (D. Don ex Grisebach) C.B. Clarke
3.1摇 Sect. Ophelia (D. Don ex Grisebach) Bentham
et Hooker f. ex Gilg
3. 1. 1摇 Ser. Corymbosae T. N. He & S. W. Liu,
ser. nov.
Inflorescentia corymbosa. Nectarium 1 in uno鄄
quoque lobo corollae, prope basin lobi corollae, par鄄
vum, patelliforme, breviter elevata squama et longe
papillosis marginalibus fimbriisque instructum vel
sacciforme, orbiculari obtegeni squama et breviter
papillosis apicalibus fimbriisque instructum. Semina
tenuiter verrucata, raro corrugata.
Inflorescences corymblike. Nectaries 1 per co鄄
rolla lobe, near the base of corolla lobe, smaller, or鄄
bicular to suborbicular, dish鄄shaped without or with
short raised scale, the bound with many long papil鄄
lose fimbriae round margin, or watch鄄packet鄄shaped
with an orbicular covered scale and many short papil鄄
lose fimbriae round opening margin at apex of pocket.
Seed coats finely warty, rarely thinly corrugate.
This series is similar to ser. Kilimandscharicae
by fimbriate nectaries, but differs by corymblike in鄄
florescences as well as nectaries smaller, near the
base of corolla lobe, orbicular to suborbicular.
Type species: Swertia corymbosa (Grisebach)
C. B. Clarke
Distribution: Restricted to S India and Sri Lanka.
3. 2摇 Sect. Swertopsis (Makino) Satake
3. 2. 1 摇 Ser. Japonicae T. N. He & S. W. Liu
(subgen. Ophelia sect. Swertopsis), ser. nov.
Herba annua. Inflorescentia cymoso鄄panicula.
Flores 5鄄, minus 4鄄meri. Nectaria 2, raro 1 in uno鄄
quoque lobo corollae, oblongum, naviculare vel sac鄄
ciforme. Semina tenuiter reticulata.
Annuals. Inflorescences a panicle of cymes.
Flowers 5鄄, less 4鄄merous. Nectaries 2, rarely 1 per
corolla lobe, oblong, boat鄄shaped, or pocket鄄shaped.
Seed coats finely reticulate.
This series is somewhat similar to sect. Ophelia
ser. Ramosae in annuals, with inflorescences a
panicle of cymes, but conspicuously distinguished by
having nectaries 2, rarely 1 per corolla lobe and
seed coats finely reticulate.
Type species: Swertia japonica (Schultes) Makino
Distribution: Much Asia except West Asia,
main in Central and East Asia. Mongolia, Russia,
Kashmir, W Pakistan, Himalaya, India, Nepal, Sik鄄
9833 期摇 摇 摇 摇 摇 摇 摇 HE Ting鄄Nong et al. : Nomenclatural Novelities in Swertia (Gentianaceae) 摇 摇 摇 摇 摇 摇 摇 摇 摇
kim, Bhutan, Bengal, almost throughout China, My鄄
anmar, Thailand, Korea, Japan, and E New Guinea.
31 species.
3. 2. 1. 1摇 Swertia tetrapetala Pallas
1a. Leaves linear to oblong, 10-20(-30)伊(2-)3-5(-10)
mm. Apex of corolla lobes obtuse, rarely acuminate
109a. var. tetrapetala………………………………
1b. Leaves ovate鄄triangular, 20 -35 伊(12 -) 15 -20 ( -23)
mm. Apex of corolla lobes acute and mucronate
109b. var. wilfordii…………………………………
109a. var. tetrapetala
Distribution: E Russia ( Sachalin, Kamchat鄄
ka), Korea, and Japan. Meadows; 100-670 m.
109b. var. wilfordii (Kerner) T. N. He, comb. nov.
Swertia wilfordii Kerner, Ber. Naturw. Ver.
Inssbruck. 1: 102 (1870). Type: China, North鄄
east Region ( Mandschuria), 44毅 - 45毅 N, July—
Aug. 1859, C. Wilford ( K! —holotype; GH!,
P! —isotypes).
Ophelia wilfordii Kerner, Ber. Naturw. Ver.
Innsbruck, 1: 102 (1870).
Swertia anomala Nakai, Bot. Mag., Tokyo 28:
331 (1914). Type: NE Korea ( Corea), Musan
(Musang), Aug. 1913, T. Mori 320 (not seen).
Leaves ovate鄄triangular, 20-35伊(12-)15-20
(-23) mm. Apex of corolla lobes acute and mucro鄄
nate. Flowering and fruiting: August—September.
Distribution: China ( Northeast Region), NE
Korea. ca. 670 m.
3. 2. 2 摇 Ser. Swertopsis T. N. He & S. W. Liu,
ser. nov.
Herba annua. Folia ovato鄄elliptica usque rhom鄄
biformia, apice acuminata caudataque. Inflorescen鄄
tia in terminales, axillares fasciculatas cymas vel
umbellas disposita. Flores 5鄄meri. Nectaria 2 in un鄄
oquoque lobo corollae, sacciformia. Semina tenuiter
reticulata.
Annuals. Leaves ovate鄄elliptic to rhombic, a鄄
pex acuminate and caudate. Inflorescences terminal
and axillary fasciculate cymes or umbels. Flowers 5鄄
merous. Nectaries 2 per corolla lobe, pocket鄄
shaped. Seeds finely reticulate.
This series is more special in sect. Swertopsis
by inflorescences terminal and axillary fasciculate
cymes or umbels and leaves ovate鄄elliptic to rhom鄄
boid, apex acuminate and caudate.
Type species: Swertia swertopsis Makino
Distribution: Japan.
3. 2. 3摇 Ser. Dichotomae T. N. He & S. W. Liu
3. 2. 3. 1 摇 Swertia dichotoma var. rubrostriata
(Y. Z. Zhao, Z. Y. Zhu & L. Q. Zhao ) T. N. He,
comb. nov.
Anagallidium rubrostriata Y. Z. Zhao, Z. Y.
Zhu & L. Q. Zhao, Acta Phytotax. Sin. 42 (1):
83-85, f. 1 (2004). Type: China, Nei Mongol,
Hohhot, Horinger, Mt. Manhan, grassland on brook
side, 1 June 1983, Z. Y. Zhu 83鄄011 (HIMC—ho鄄
lotype and isotype, not seen).
Var. rubrostriata is similar to var. dichotoma in
having dichotomously branched stems, nectaries 2
per corolla lobe, brown, each with an orbicular cov鄄
ered scale and a small corner on back of the scale,
but differs by corolla orange鄄colored, with red stria鄄
te, leaves rugose, with crisp margin.
1a. Corolla white, sometimes with purple spots; leaves level
var. dichotoma………………………………………
1b. Corolla orange鄄colored, with red striate; leaves rugose,
with crisp margin var. rubrostriata…………………
摇 摇 Distribution: China (Nei Mongol) .
3. 3摇 Sect. Platynema T. N. He & S. W. Liu
3. 3. 1摇 Swertia paniculata var. gracilescens (H.
Smith) T. N. He, comb. nov.
Swertia gracilescens H. Smith, Bull. Brit.
Mus. (Nat. Hist. ) Bot. 4: 251, pl. 33 (1970).
Type: Nepal, Bhuji Khola, open slope, 2 100 m,
15 Oct. 1954, J. D. A. Stainton, W. R. Sykes &
L. H. J. Williams 9049 ( BM! —holotype; E!,
GH! —isotypes). Paratypes: Nepal, Gurjakhani
2 600 m, exposed grass slope, 17 Sept. 1954 J. D.
A. Stainton, W. R. Sykes & L. H. J. Williams
4470; Kalii Gandaki valley, Taglung, south of
Tukucha, 3 700 m, on grass slopes, 22 Sept. 1954,
J. D. A. Stainton, W. R. Sykes & L. H. J. Williams
7970 (all BM!, GH!).
093摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 植 物 分 类 与 资 源 学 报摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 第 35 卷
Var. gracilescens is distinguished from var.
paniculata by corolla pure white without coloured
spots and band and whole plant slender and small:
stems 8-30 cm tall, leaves 1 -1. 5 cm伊1. 5 -2. 5
mm, calyx lobes 4. 5-5伊0. 5-1 mm, corolla lobes
to 5. 5 伊4 mm. Var. paniculata is characterized by
the corolla with 2 purple or yellow鄄green spots above
each nectary and plants relatively robust: stems 35-
120 cm tall, leaves 2 -6. 5 cm 伊 4 -7 mm, calyx
lobes 5-5. 5伊1. 5-2. 5 mm, and corolla lobes 6-
6. 5伊 4-4. 5 mm.
Swertia paniculata is a more common species in
the Himalayan region and characterized by oblong to
lanceolate leaves, calyx 2 / 3 to longer than corolla,
white corolla, horseshoe鄄shaped and naked necta鄄
ries, and dilated and free filaments.
Distribution: Nepal.
3. 4 摇 Sect. Echinulata T. N. He & S. W. Liu,
sect. nov.
Herba perennis vel annua. Nectaria 2 in uno鄄
quoque lobo corollae, oblonga, navicularia, fimbria鄄
ta. Stylus absens; stigmate orbiculato. Semina par鄄
va, subtiliter reticulata. Pollinis granula echinulato
et saepe perforato exinio ornata.
Perennials or annuals. Nectaries 2 per corolla
lobe, oblong, boat鄄shaped, with a narrow scale and
raised fimbriate margin. Style absent; stigma orbicu鄄
lar. Seeds finely reticulate. Pollen grains are sub鄄
spheroidal, but subprolate and suboblate grains oc鄄
cur in infraspecific variation; medium鄄sized with the
size range of 22. 2 -27. 3 伊19. 6 -29. 3 滋m; colpi
medium, long or short. The sexine ornamentation is
echinulate and perforate but imperforate in S. usam鄄
barensis. All perforations are very small.
Sect. Echinulata is somewhat similar to sect. swer鄄
topsis in gross morphology, but differs by absent style,
orbicular stigma as well as echinulate pollen grains.
Type species: Swertia abyssinica Hochstetter
Distribution: East and Northeastern tropical
and Northeast Africa ( Congo, Ethiopia, Kenya,
Tanzania, and Uganda) and West Asia (N. Yemen
and SW. Saudia).
18 species, may be divided in 2 series.
3.4.1摇 Ser. Pumilae T. N. He & S.W. Liu, ser. nov.
Herba perennis. Radix crassa, fibrosa usque semil鄄
ignosa. Caules e basi multum ramosi. Folia basalia sa鄄
epe abundantia, persistentia et in rosulam conferta.
Perennials. Taproot stout, fibrous to semiwoody.
Stems much branched from very base. Basal leaves u鄄
sually abundant, persistent and crowded into a rosette
(usually deciduous at anthesis in S. quartiniana).
Type: Swertia pumila Hochstetter
Distribution: East and Northeast Africa ( Con鄄
go, Ethiopia, Kenya, Tanzania, and Uganda) and
West Asia (N. Yemen and SW. Saudia),
9 species.
3. 4. 2摇 Ser. Abyssinicae T. N. He & S. W. Liu,
ser. nov.
Herba annua. Radix tenuis, fibrosa. Caules
simplices vel e basi ramosi. Folia basalia saepe pau鄄
ca, sub anthesi decidua.
Annuals. Roots slender, fibrous. Stem simple
or branched from base. Basal leaves usually few,
deciduous at anthesis.
Type: Swertia abyssinica Hochstetter
Distribution: throughout the tropical Africa in
North, West, East, Central, and South Africa.
9 species.
4摇 African Swertia species
The African Swertia species have been studied
by some botanists such as Fries (1923), Hedberg
(1957), and Nemomissa (1998), which included
ca. 24 species and divided into some informal
groups. We have also studied on gross morphology,
palynology ( using SEM and LE) and known chro鄄
mosome numbers from the whole African and Wes鄄
tern Asian species. The results indicate that there
are ca. 28 species. They may be divided into two
sections, of which each further into two series
(sect. Montana: ser. Repentes and ser. Kilimands鄄
charica, sect. Echinulata: ser. Pumilae and ser.
Abyssinicae) respectively. Both sect. Montana and
sect. Echinulata differ from each other as follows:
1933 期摇 摇 摇 摇 摇 摇 摇 HE Ting鄄Nong et al. : Nomenclatural Novelities in Swertia (Gentianaceae) 摇 摇 摇 摇 摇 摇 摇 摇 摇
1a. Nectary1 per corolla lobe; seeds corrugate; pollen grains essentially striate, varying from irregularly striate鄄reticulate, striate鄄
microreticulate to striate鄄perforate or scattered and thinly striate鄄imperforate (Sect. Montana):
2a. Nectary reduced to a flat viscous gland patch without raised margin and fimbriae; stems trailing or stoloniferous with ad鄄
ventitious roots at nodes; flowers solitary on long terminal pedicels or in few鄄flowered terminal cymes
Ser. Repentes…………………………………………………………………………………………………………
2b. Nectary fimbriate; stems erect, without stolons and adventitious roots; inflorescences a many鄄flowered panicle of cymes
Ser. Kilimandscharicae………………………………………………………………………………………………
1b. Nectaries 2 per corolla lobe; seeds finely reticulate; pollen grains essentially echinulate and small perforate, but imperforate
in one species (Sect. Echinulata):
3a. Perennials with a stout fibrous to semiwoody taproot; basal leaves usually abundant, persistent and crowded into a ro鄄
sette (usually deciduous at anthesis in S. quartiniana); stems much branched from very base Ser. Pumilae………
3b. Annuals with slender fibrous roots; basal leaves usually few, deciduous at anthesis; stems simple or branched from base
Ser. Abyssinicae………………………………………………………………………………………………………
摇 摇 In addition, there is an African species of Lom鄄
atogonium as follows:
Lomatogonium pleurogynoides ( Baker ) S.
W. Liu & T. N. He, comb. nov.
Swertia pleurogynoides Baker, Bull. Misc. In鄄
form., Kew 1898: 158 (1898). Type: Malawi (Nyasa鄄
land), between Kondow and Karonga, 2 000-6 000 ft.,
Whyte s. n. (K! —holotype; BM! —isotype).
Swertia heterosepala Gilg, Notizbl. Bot. Gart.
Berlin 14: 115 (1938). Type: Tanzania, Matengo
highland, WSW from Songea, 1 100-1 200 m, 18
April 1936, H. Zerny 604 (K! —isotype).
It is similar to L. rotatum in leaves and calyx
lobes linear, calyx as long as or rarely longer or
slightly shorter than corolla, but differs by corolla
smaller, 7-8. 5 mm long.
Distribution: East Africa ( Ethiopia, Tanzani鄄
a), South Africa ( Mozambique, Malawi ). Wet
grassland, shire highland; 600-1 800 m.
References:
Chassot P, Nemomissa S, Yuan YM et al., 2001. High paraphyly of
Swertia L. (Gentianaceae) in the Gentianella鄄lineage as revealed
by nuclear and chloroplast DNA sequence variation [ J] . Plant
Systematics & Evolution, 229: 1—21
Fries TCE, 1923. Die Swertia鄄arten der afrikanischen Hochgebirge
[J] . Notizblatt des Botanischen Gartens und Museums zu Berlin鄄
Dahlem, 8: 389—423
Hagen KB, Kadereit JW, 2002. Phylogeny and flower evolution of the
Swertiinae (Gentianaceae鄄Gentianeae): homoplasy and the principle
of variable proportions [J]. Systematic Botany, 27: 548—572
Hedberg O, 1957. Afroalpine vascular plants - a taxonomic revision
[J] . Symbolae Botanicae Upsalienses, 15: 1—411, P1. 1—12
He TN, 1988. Gentianaceae [A]. Flora Reipublicae Popularis Sini鄄
cae Tom. 62 [M]. Beijing: Science Press
He TN, 1994. New taxa of Gentianaceae from China [J] . Novon, 4:
369—372
He TN, Liu SW, 1980. New taxa of Swertia L. from China [J] . Ac鄄
ta Phytotaxonomica Sinica, 18: 75—85
He TN, Pringle JS, 1995. Gentianaceae [A]. In Wu CY, Raven
PH (ed.), Flora of China [M]. Beijing: Science Press; St.
Louis: Missouri Botsnical Garden, 16: 1—140
He TN, Xue CY, Wang W, 1994. The origin, dispersal and forma鄄
tion of the distribution pattern of Swertia L. ( Gentianaceae)
[J] . Acta Phytotaxonomica Sinica, 32: 525—537
Liu JQ, Chen ZD, Lu AM, 2001. A preliminary analysis of the phy鄄
logeny of the Swertiinae (Gentianaceae) based on ITS data [J] .
Israel Journal of Plant Science, 49: 301—308
Nemomissa S, 1998. A synopsis of Swertia ( Gentianaceae) in east
and northeast tropical Africa [ J] . Kew Bulletin, 53: 419—
436, http: / / www. jstor. org / stable / 4114507
Pissjaukova VV, 1979. Generis Swertiae L. (Gentianaceae) taxa no鄄
va, 1 [J]. Novosti Sistematiki Vysshikh Rastenii, 15: 202—203
Struwe L, Albert VA, 2002. Gentianaceae [A]. Systematics and Nat鄄
ural History [M]. Cambridge: Cambridge University Press, 652
Xue CY, He TN, Liu JQ, 1999a. Embryology of Swertia tetraptera
Maxim. (Gentianaceae) and its systematic implication [J] . Ac鄄
ta Phytotaxonimica Sinica, 37: 259—263
Xue CY, He TN, Liu JQ, 1999b. Embryology of a Tibetan medicine
Halenia elliptica [J] . Acta Botanica Yunnanica, 21: 212—217
Yuan YM, K俟pfer P, 1995. Molecular phylogenetics of the subtribe
Gentianae (Gentianaceae) inferred from the sequences of inter鄄
nal transcribed spacers (ITS) of nuclear ribosomal DNA [ J] .
Plant Systematics & Evolution, 196: 207—226
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