Table Of Content植 物 分 类 与 资 源 学 报 ,35 ( ):
摇 2013 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
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 Montana 和 Echinulata 和七个新系 Repentes Kilimandscharicae
(sect. sect. ) (ser. , ser. , ser.
Corymbosae Japonicae Swertopsis Pumilae和 Abyssinicae 同时对獐牙菜属中的一些类群
, ser. ,ser. ,ser. ser. )。
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: 文献标识码: 文章编号:
Q949摇 摇 摇 摇 摇 摇 摇 A摇 摇 摇 摇 摇 摇 摇 2095-0845(2013)03-386-07
Nomenclatural Novelities in Swertia Gentianaceae
( )
**
HE Ting鄄Nong, LIU Shang鄄Wu, CHEN Shi鄄Long
KeyLaboratoryofAdaptationandEvolutionofPlateauBiota NorthwestInstituteofPlateauBiology
( , ,
ChineseAcademyofSciences
,Xining810001,China)
Abstract Swertia
: is one large genus of Gentianaceae with a wide distribution in Asia,North America,Europe and
Montana Echinulata Repentes
Africa. In this study, we reported two new sections ( and ) and seven new series ( ,
Kilimandscharicae Corymbosae Japonicae Swertopsis Pumilae Abyssinicae
, , , , ,and ). Inaddition,somespecieswere
reduced, recombined and newly established. These new taxonomic synopses will be used for a further worldwide
monograph of this genus.
Key words Swertia
: Taxonomic synopses; ; Gentianaceae
Swertia
摇 is one of the large genera in the family cies clustered with the multiple related genera
et al
Gentianaceae (Struwe and Albert, 2002). This ge鄄 (Yuan and K俟pfer, 1995; Chassot ., 2001;
et al
nus occurs from Asia, Europe to Africa and North Liu .,2001; Hagen and Kadereit,2002). This
et al
America (He ., 1994). In the past century, suggested that the traits, the rotate corolla and
numerous new species were described under this ge鄄 fringed nectaries, which were used to circumscribe
nus or transferred from the other genera especially the genus, may have derived from homoplastic evo鄄
from China (He and Liu, 1980; He, 1988, 1994; lutions due to the radiative diversifications or repre鄄
He and Pringle, 1995). The circumscription of this sented the ancestral states of the total family (Hagen
genus is highly difficult because all phylogenetic ana鄄 and Kadereit, 2002). As inferred from molecular
lyses based on molecular data suggested that this ge鄄 phylogenies (e. g. Yuan and K俟pfer, 1995) and
et al
nus is not monophyletic and some of the paralleling embryological studies (Xue ., 1999a, b), we
Heteranthos
clades from the radiative diversification of some spe鄄 found that sect. of this genus can be ex鄄
*Foundation tiems:KeyProjectoftheNationalScienceFoundationofChina(31270270) andtheNationalBasicResearchProgramofChina
(2007FY110100)
**Authorforcorrespondence;E鄄mail:[email protected]
Receiveddate: 2013-03-18, Accepteddate: 2013-03-25
作者简介 何廷农 女 已故中国龙胆科植物分类学专家
: (1938-2011) , 。
期 et al Swertia
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387
cluded from the genus and elevated to a generic sta鄄 oblong鄄spathulate to elliptic, base narrowed, apex
tus based on its recognizable morphological distinct鄄 round and nectaries linear鄄oblong, radially elongated.
ness: dimorphic plants and flowers and a cleistoga鄄 Distribution: China (SW Xizang).
mous pollination mode. However, up to now, mor鄄
2摇 Subgen. Poephila
phological traits, which can be used to distinguish C.B. Clarke
2 1摇 Sect. Frasera
the other paralleling clades, need further examina鄄 . (Walter) Knoblauch
2 1 1摇 Swertia albicaulis var. modocensis
tions. Therefore, we tentatively follow the traditional . . (St.
Swertia
generic circumscription of for the further in鄄 John) T.N. He, comb. nov.
Swertia modocensis
frageneric classification. Through examining thou鄄 St. John, Amer. Midl. Natu鄄
26
sands of specimens kept in the major herbaria within ralist :18 (1941). Type: United States, Califor鄄
the past three decades, we recognized 168 species nia, Lassen County, Madeline, gentle northeast fa鄄
Artemisia
for this genus over the world. We classified these cing slope, in rocky volcanic soil, among
Juniperus occidentalis
species into three subgenera,11 sections and 11 se鄄 and scattered , 4 500 ft., 8
Montana Echinula鄄
ries. Among them, sections and June 1936, E. B. Babcock & G. L. Stebbins 1790
ta Repentes Kilimandscharicae Corym鄄
, and series , , (UC!—holotype; GH!—isotype). Paratypes: Ca鄄
bosae Japonicae Swertopsis Pumilae Abys鄄
, , , , and lifornia, Modoc County, in 1907, M. H. Manning
sinicae
were newly proposed for the first time. In ad鄄 (UC!), Shasta County, 17 May 1923, E. Bethel
dition, some taxonomic entities established previous鄄 (SF, not seen); Oregon, July 1898, R.M. Austin
ly for this genus or related genera were reduced or & Bruce 2153 (UC!), Southern Oregon, between
recombined. Although the currently circumscribed Merrill and Cottonwood, June1901, H.W. Furlong,
genus is still polyphyletic, these infrageneric entities W.B. Greeley, M. Wilson et al. s.n. (UC!).
Swertia modocensis adglabra
seem to be monophyletic if assessed only based on var. St. John, A鄄
26
the morphological similarity. Here we report these mer. Midl. Naturalist :19 (1941). Type: Uni鄄
Swertia
new taxonomic synopses established for . ted States, Oregon, Klamath County, June 1901,
These taxa will be used for a further worldwide mon鄄 H.W. Furlong, W. B. Greeley, M. Wilson et al.
ograph, phylogenetic analyses and resource surveys s.n. (UC! —holotype).
Swertia sierrae
related to this genus. St. John, Amer. Midl. Natura鄄
26
list : 16 (1941). Type: United States, Califor鄄
1摇 Subgen. Swertia
nia, Lassen County, Madeline, about two miles
1.1摇 Sect. Apterae
V.V. Pissjaukova north of, gentle northeast facing slope, in rocky vol鄄
1 1 1摇 Swertia cuneata var. asterocalyx Artemisia Juniperus
. . (T. N. canic soil, among and scattered
occidentalis
He & S.W. Liu) T.N. He & S.W. Liu,comb. nov. , 4500ft.,8 June1936, E.B. Babcock
Swertia asterocalyx
T.N. He & S.W. Liu, Acta & G. L. Stebbins 1790 (GH!, UC! —isotypes).
18
Phytotax. Sin. (1): 77, f. 2 (1980). Type: Paratypes: United States, N California, Willow
China, Xizang, Nyalam, 3 800-4 000 m, 5 Sept. Creek, Devils Garden, July 1898, R. M. Austin
1972, Xizang Medic. Herb. Exped. 1825 (HNWP! 418 (US!); Dry Lake, Modoc County, 31 May,
—holotype; PE!—isotype). 1897, E.I. Applegate 877 (US!); Oregon: Thom鄄
asterocalyx cu鄄
Var. is distinguished from var. as Creek Ranger Station, Lake County, 1 570 m,
neata
by flowers large: corolla 20-30 mm in diam., June 29-30, 1911, W.W. Eggleston 5098 (NY!,
cuneata
lobes15-20mm. Var. has small flowers: co鄄 US!); Stein爷s Mountain, 1 June, 1885, J. T.
rolla8-10mm in diam.,lobes7.5-10(-12)mm. The Howell (US!); near Warm Range, 1 500 m, Lake
Swertia cuneata
species is characterized by calyx lobes County, 26 July, 1896, F. V. Coville & J. B.
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Leiberg 53 (US!). ta, striato鄄microreticulata usque striato鄄reticulata).
Swertia shastaensis
St. John, Amer. Midl. Na鄄 Perennials, with slightly fleshy to semiwoody
26
turalist : 19 (1941). Type: United States, Cali鄄 tap鄄root and some rootlets, or tap鄄root absent but
fornia, Mount Shasta, near Sheep Rock, 5 000 ft., stems trailing or stoloniferous with adventitious
June 1903, H. M. Hall & E. B. Babcock 4113 roots. Flowers 5鄄, rarely 4鄄merous. Nectary 1 per
(UC! —holotype). corolla lobe. Style short to absent. Seeds either many
Frasera albicaulis modocensis
var. (St. John) N. corrugate and wingless, with longitudinal, thin and
Holmgren in A. Cronquist et al., Intermountain Fl., dense corrugations or many corrugate and ridged,
4
Vasc. Pl. Intermountain West,U.S.A., :22 (1984). with irregular narrow membranous reticulate ridges.
Swertia modocensis adglabra
and its variety var. Pollen grainsare subspheroidal,rarely prolate or sub鄄
S. albicaulis
are close to species group , which in鄄 prolate; medium鄄, rarely small鄄 or large鄄sized, size
cludes six varieties, in having opposite stem leaves range 23.3-43.8伊19.0-33.7滋m;colpi long or me鄄
with white membranous margin, a narrow interrupted dium. The sexine ornamentation is essentially striate,
spike鄄like panicle, and oblong boat鄄shaped shortly varying from thinly, undulately striate & perforate to
fimbriate nectaries. Especially it is more similar to imperforate. The lira width is 0. 17 -0. 30 滋m.
albicaulis cusickii n n
var. and var. , but differs from Chromosome number 2 =26, rarely 2 =20.
both in corolla pale lavender, with dark purple This section is characterized by perennials,
streak, and also from the former in coronal scales with slightly fleshy to semiwoody tap鄄root or with ad鄄
albicaulis
entire to shallowly lacerate (in var. corol鄄 ventitious roots from nodes of trailing or stoloniferous
la pale blue, usually not darker mottled; coronal stems, nectary 1 per corolla lobe so that it conforms
Poephila
scales deeply laciniate to near base into numerous to the essential features of subgen. . In this
Poephila
setae), from the latter in whole plants puberulent subgenus it issimilar to sect. ,but differsby
cu鄄
and coronal scales short, 1.5-2.5 mm (in var. having seeds longitudinally densely and minutely cor鄄
sickii
plants glabrous except for leaves puberulent at rugate鄄 ridged and flowers usually 5鄄merous whereas
base and sometimes along midvein of abaxial sur鄄 the latter having seed coats almost smooth and some鄄
face; coronal scales long, 3-4 mm; corolla pure times winged at one or both end, flowers always 4鄄
pale blue). However, the coronal scales (crown merous. Pissjaukova (1979) put some species of
S. albicaulis S.volkensii S.subnivalis S.scandens
scales) of group are deltoid to linea鄄 this section ( , , ,
S.macrosepala S. keniensis Apterae
roblong, margin extremely variable from entire to and ) in sect. of
Swertia modocensis Swertia Aptera
deeply laciniate. Hence and var. subgen. . However, sect. is very dif鄄
adglabra Montana
are not necessary to maintain as a distinct ferent from sect. by having an abbreviated
species but as a variety. rhizome,2 nectaries per corolla lobe, and seeds lon鄄
Distribution: W United States (Oregon and Cali鄄 gitudinally few (8-12) corrugate鄄ridged with high
fornia). irregular membranous and reticulate ridges.
2 2摇 Sect. Montana Swertia volkensii
. T.N. He &S.W. Liu,sect. nov. Type species: Gilg
Herba perennis. Radix leviter carosa usque Distribution: from North Africa to Northeast
semilignosa, vel caulis inferior radices adventitias and Tropical Africa.
emittens. Flores 5鄄, raro 4鄄meri. Nectarium 1 in 10 species, may be divided into 2 series.
2 2 1摇 ser. Repentes
unoquoque lobo corollae. Stylus brevis usque ab鄄 . . T.N. He &S.W. Liu,ser. nov.
sens. Semina longitudinaliter, dense et tenuiter cor鄄 Caulis repens vel stoloniformis, radices adven鄄
rugata vel corrugato鄄porcata. Pollinis granula admo鄄 titias emittens. Flores solitarii vel in inflorescentiam
dum striata (sparse et tenuiter striata,striato鄄perfora鄄 parco鄄florigeram dispositi. Nectarium planum, mac鄄
期 et al Swertia
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389
uliforme, absque marginibus et fimbriis. Inflorescentia corymbosa. Nectarium 1 in uno鄄
Stems trailing or stoloniferous with adventitious quoque lobo corollae, prope basin lobi corollae, par鄄
roots. Basal leaves crowded into a rosette. Flowers vum, patelliforme, breviter elevata squama et longe
S. subnivalis S. sar鄄
5鄄merous (but 4鄄merous in and papillosis marginalibus fimbriisque instructum vel
mentosa
), solitary on long terminal pedicels or in sacciforme, orbiculari obtegeni squama et breviter
few鄄flowered terminal cymes. Nectary reduced into a papillosis apicalibus fimbriisque instructum. Semina
flat gland patch without raised margin and fimbriae. tenuiter verrucata, raro corrugata.
Swertia volkensii
Type species: Gilg Inflorescences corymblike. Nectaries 1 per co鄄
Distribution: East and Central Africa. rolla lobe, near the base of corolla lobe, smaller, or鄄
5 species. bicular to suborbicular, dish鄄shaped without or with
2 2 1 1摇 Swertia sarmentosa
. . . T. N. He & S. W. short raised scale, the bound with many long papil鄄
Liu, nom. nov. lose fimbriae round margin, or watch鄄packet鄄shaped
Swertia scandens
T. C. E. Fries, Notizbl. Bot. with an orbicular covered scale and many short papil鄄
8
Gart. Berlin : 516, 3c (1923), non L佴veill佴 lose fimbriae round opening margin at apex of pocket.
(1913). Syntypes: W Kenya, in rain forest, ca. Seed coats finely warty, rarely thinly corrugate.
Kilimandscharicae
2350 m, 3 Jan. 1922, E. A. Robenson & Th. C. This series is similar to ser.
E. Fries 691 (BM!, K!, S!, UPS!); same locali鄄 by fimbriate nectaries, but differs by corymblike in鄄
ty, 30 Jan. 1922, E. A. Robinson & Th. C. E. florescences as well as nectaries smaller, near the
Fries 1277 (K!). Lectotype: Rob. E. & Th. C. base of corolla lobe, orbicular to suborbicular.
Swertia corymbosa
E. Fries 691 (UPS!—lectotype; BM!, K!, S! — Type species: (Grisebach)
isolectotypes; designated here). C.B. Clarke
Distribution: East Africa (Kenya). Distribution:Restricted to S India and Sri Lanka.
2 2 2摇 Ser. Kilimandscharicae 3 2摇 Sect. Swertopsis
. . T.N. He & S.W. . (Makino) Satake
3 2 1摇 Ser. Japonicae
Liu, ser. nov. . . T. N. He & S. W. Liu
Monobothrium 27 Ophelia Swertopsis
Hochstetter,Flora :27 (1844). (subgen. sect. ), ser. nov.
M.schimperi
Type: Hochstetter Herba annua. Inflorescentia cymoso鄄panicula.
Caules erecti, absque stolinibus et radicibus Flores 5鄄, minus 4鄄meri. Nectaria 2, raro 1 in uno鄄
adventitiis. Cymae in inflorescentiam paniculatam quoque lobo corollae, oblongum, naviculare vel sac鄄
dispositae. Nectarium V鄄forme, fimbriatum. ciforme. Semina tenuiter reticulata.
Stems erect, without stolons and adventitious Annuals. Inflorescences a panicle of cymes.
roots. Inflorescences a many鄄flowered panicle of cymes. Flowers 5鄄, less4鄄merous. Nectaries2, rarely1 per
Nectary V鄄shaped, fimbriate. corolla lobe,oblong,boat鄄shaped,or pocket鄄shaped.
Swertia kilimandscharica
Type species: Engler Seed coats finely reticulate.
Ophelia
Distribution: from North Africa to Northeast This series is somewhat similar to sect.
Ramosae
and Tropical Africa. ser. in annuals, with inflorescences a
5 species. panicle of cymes, but conspicuously distinguished by
having nectaries 2, rarely 1 per corolla lobe and
3摇 Subgen. Ophelia
(D. DonexGrisebach)C.B. Clarke seed coats finely reticulate.
3 1摇 Sect. Ophelia Swertia japonica
. (D. Don ex Grisebach) Bentham Type species: (Schultes) Makino
et Hooker f. ex Gilg Distribution: Much Asia except West Asia,
3 1 1摇 Ser. Corymbosae
. . T.N. He & S.W. Liu, main in Central and East Asia. Mongolia, Russia,
ser. nov. Kashmir,W Pakistan,Himalaya,India,Nepal,Sik鄄
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Swertopsis
kim, Bhutan, Bengal, almost throughout China, My鄄 This series is more special in sect.
anmar, Thailand,Korea,Japan,and E New Guinea. by inflorescences terminal and axillary fasciculate
31 species. cymes or umbels and leaves ovate鄄elliptic to rhom鄄
3 2 1 1摇 Swertia tetrapetala
. . . Pallas boid, apex acuminate and caudate.
Swertia swertopsis
Type species: Makino
1a. Leaves linear to oblong,10-20(-30)伊(2-)3-5(-10)
mm. Apex of corolla lobes obtuse, rarely acuminate Distribution: Japan.
……………………………… 109a. var. tetrapetala 3 2 3摇 Ser. Dichotomae
. . T.N. He & S.W. Liu
3 2 3 1摇 Swertia dichotoma var. rubrostriata
1b. Leaves ovate鄄triangular, 20-35伊(12-)15-20(-23) . . .
mm. Apex of corolla lobes acute and mucronate (Y.Z. Zhao, Z.Y. Zhu & L.Q. Zhao ) T.N. He,
………………………………… 109b. var. wilfordii
comb. nov.
109a. var. tetrapetala Anagallidium rubrostriata
Y. Z. Zhao, Z. Y.
42
Distribution: E Russia (Sachalin, Kamchat鄄 Zhu & L. Q. Zhao, Acta Phytotax. Sin. (1):
ka), Korea, and Japan. Meadows; 100-670 m. 83-85, f. 1 (2004). Type: China, Nei Mongol,
109b. var. wilfordii
(Kerner) T.N. He,comb. nov. Hohhot, Horinger, Mt. Manhan, grassland on brook
Swertia wilfordii
Kerner, Ber. Naturw. Ver. side,1 June 1983, Z.Y. Zhu 83鄄011 (HIMC—ho鄄
1
Inssbruck. : 102 (1870). Type: China, North鄄 lotype and isotype, not seen).
rubrostriata dichotoma
east Region (Mandschuria), 44毅-45毅N, July— Var. is similar to var. in
Aug. 1859, C. Wilford (K! —holotype; GH!, having dichotomously branched stems, nectaries 2
P! —isotypes). per corolla lobe, brown, each with an orbicular cov鄄
Ophelia wilfordii
Kerner, Ber. Naturw. Ver. ered scale and a small corner on back of the scale,
1
Innsbruck, : 102 (1870). but differs by corolla orange鄄colored, with red stria鄄
Swertia anomala 28
Nakai, Bot. Mag., Tokyo : te, leaves rugose, with crisp margin.
331 (1914). Type: NE Korea (Corea), Musan 1a. Corolla white, sometimes with purple spots; leaves level
……………………………………… var. dichotoma
(Musang), Aug. 1913, T. Mori 320 (not seen).
Leaves ovate鄄triangular, 20-35伊(12-)15-20 1b. Corolla orange鄄colored, with red striate; leaves rugose,
………………… var. rubrostriata
with crisp margin
(-23) mm. Apex of corolla lobes acute and mucro鄄
nate. Flowering and fruiting: August—September. 摇 摇 Distribution: China (Nei Mongol).
3 3摇 Sect. Platynema
Distribution: China (Northeast Region), NE . T.N. He & S.W. Liu
3 3 1摇 Swertia paniculata var. gracilescens
Korea. ca. 670 m. . . (H.
3 2 2摇 Ser. Swertopsis
. . T. N. He & S. W. Liu, Smith) T.N. He, comb. nov.
Swertia gracilescens
ser. nov. H. Smith, Bull. Brit.
4
Herba annua. Folia ovato鄄elliptica usque rhom鄄 Mus. (Nat. Hist.) Bot. : 251, pl. 33 (1970).
biformia, apice acuminata caudataque. Inflorescen鄄 Type: Nepal, Bhuji Khola, open slope, 2 100 m,
tia in terminales, axillares fasciculatas cymas vel 15 Oct. 1954, J. D. A. Stainton, W. R. Sykes &
umbellas disposita. Flores5鄄meri. Nectaria2 in un鄄 L. H. J. Williams 9049 (BM! —holotype; E!,
oquoque lobo corollae, sacciformia. Semina tenuiter GH! —isotypes). Paratypes: Nepal, Gurjakhani
reticulata. 2600 m, exposed grass slope, 17 Sept. 1954 J.D.
Annuals. Leaves ovate鄄elliptic to rhombic, a鄄 A. Stainton, W. R. Sykes & L. H. J. Williams
pex acuminate and caudate. Inflorescences terminal 4470; Kalii Gandaki valley, Taglung, south of
and axillary fasciculate cymes or umbels. Flowers5鄄 Tukucha,3700m, on grass slopes,22 Sept. 1954,
merous. Nectaries 2 per corolla lobe, pocket鄄 J.D.A. Stainton, W.R. Sykes & L.H.J. Williams
shaped. Seeds finely reticulate. 7970 (all BM!, GH!).
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391
gracilescens
Var. is distinguished from var. 18 species, may be divided in 2 series.
paniculata 3 4 1摇 Ser. Pumilae
by corolla pure white without coloured . . T.N. He &S.W. Liu,ser. nov.
spots and band and whole plant slender and small: Herba perennis. Radixcrassa,fibrosausquesemil鄄
stems 8-30 cm tall, leaves 1-1. 5 cm伊1. 5-2. 5 ignosa. Caules e basi multum ramosi. Folia basalia sa鄄
mm, calyx lobes 4.5-5伊0.5-1 mm, corolla lobes epe abundantia,persistentia et in rosulam conferta.
paniculata
to 5.5伊4 mm. Var. is characterized by Perennials. Taproot stout, fibrous to semiwoody.
the corolla with2 purple or yellow鄄green spots above Stems much branched from very base. Basal leaves u鄄
each nectary and plants relatively robust: stems35- sually abundant, persistent and crowded into a rosette
S.quartiniana
120 cm tall, leaves 2-6. 5 cm 伊4-7 mm, calyx (usually deciduous at anthesis in ).
Swertia pumila
lobes 5-5.5伊1.5-2.5 mm, and corolla lobes 6- Type: Hochstetter
6.5伊4-4.5 mm. Distribution: East and Northeast Africa (Con鄄
Swertia paniculata
is a more common species in go, Ethiopia, Kenya, Tanzania, and Uganda) and
the Himalayan region and characterized by oblong to West Asia (N. Yemen and SW. Saudia),
lanceolate leaves, calyx 2/3 to longer than corolla, 9 species.
3 4 2摇 Ser. Abyssinicae
white corolla, horseshoe鄄shaped and naked necta鄄 . . T. N. He & S. W. Liu,
ries, and dilated and free filaments. ser. nov.
Distribution: Nepal. Herba annua. Radix tenuis, fibrosa. Caules
3 4摇 Sect. Echinulata
. T. N. He & S. W. Liu, simplices vel e basi ramosi. Folia basalia saepe pau鄄
sect. nov. ca, sub anthesi decidua.
Herba perennis vel annua. Nectaria 2 in uno鄄 Annuals. Roots slender, fibrous. Stem simple
quoque lobo corollae, oblonga, navicularia, fimbria鄄 or branched from base. Basal leaves usually few,
ta. Stylus absens; stigmate orbiculato. Semina par鄄 deciduous at anthesis.
Swertia abyssinica
va, subtiliter reticulata. Pollinis granula echinulato Type: Hochstetter
et saepe perforato exinio ornata. Distribution: throughout the tropical Africa in
Perennials or annuals. Nectaries 2 per corolla North, West, East, Central, and South Africa.
lobe, oblong, boat鄄shaped, with a narrow scale and 9 species.
raised fimbriate margin. Style absent; stigma orbicu鄄
4摇 African Swertia species
lar. Seeds finely reticulate. Pollen grains are sub鄄
Swertia
spheroidal, but subprolate and suboblate grains oc鄄 The African species have been studied
cur in infraspecific variation; medium鄄sized with the by some botanists such as Fries (1923), Hedberg
size range of 22. 2-27. 3伊19. 6-29. 3 滋m; colpi (1957), and Nemomissa (1998), which included
medium, long or short. The sexine ornamentation is ca. 24 species and divided into some informal
S. usam鄄
echinulate and perforate but imperforate in groups. We have also studied on gross morphology,
barensis
. All perforations are very small. palynology (using SEM and LE) and known chro鄄
Echinulata swer鄄
Sect. is somewhat similar to sect. mosome numbers from the whole African and Wes鄄
topsis
in gross morphology, but differs by absent style, tern Asian species. The results indicate that there
orbicular stigma as well as echinulate pollen grains. are ca. 28 species. They may be divided into two
Swertia abyssinica
Type species: Hochstetter sections, of which each further into two series
Repentes Kilimands鄄
Distribution: East and Northeastern tropical (sect. Montana: ser. and ser.
charica Echinulata Pumilae
and Northeast Africa (Congo, Ethiopia, Kenya, , sect. : ser. and ser.
Abyssinicae Montana
Tanzania, and Uganda) and West Asia (N. Yemen ) respectively. Both sect. and
Echinulata
and SW. Saudia). sect. differ from each other as follows:
植 物 分 类 与 资 源 学 报 第 卷
摇392摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 35
1a. Nectary1percorollalobe;seedscorrugate;pollengrainsessentiallystriate,varyingfromirregularlystriate鄄reticulate,striate鄄
Sect. Montana
microreticulate to striate鄄perforate or scattered and thinly striate鄄imperforate ( ):
2a. Nectary reduced to aflatviscousglandpatchwithoutraisedmarginandfimbriae;stemstrailingorstoloniferouswithad鄄
ventitious roots at nodes; flowers solitary on long terminal pedicels or in few鄄flowered terminal cymes
………………………………………………………………………………………………………… Ser. Repentes
2b. Nectary fimbriate; stemserect,withoutstolonsandadventitiousroots;inflorescencesamany鄄floweredpanicleofcymes
……………………………………………………………………………………………… Ser. Kilimandscharicae
1b. Nectaries2 percorollalobe;seedsfinelyreticulate;pollengrainsessentiallyechinulateandsmallperforate,butimperforate
Sect. Echinulata
in one species ( ):
3a. Perennials with a stout fibrous to semiwoody taproot; basal leaves usually abundant, persistent and crowded into a ro鄄
S.quartiniana ……… Ser. Pumilae
sette (usually deciduous at anthesis in ); stems much branched from very base
3b. Annuals with slenderfibrousroots;basalleavesusuallyfew,deciduousatanthesis;stemssimpleorbranchedfrombase
……………………………………………………………………………………………………… Ser. Abyssinicae
Lom鄄 SystematicBotany 27
摇 摇 In addition, there is an African species of ofvariableproportions [J]. , :548—572
atogonium HedbergO,1957. Afroalpine vascular plants - a taxonomic revision
as follows: SymbolaeBotanicaeUpsalienses 15
Lomatogonium pleurogynoides [J]. , :1—411,P1. 1—12
(Baker) S. FloraReipublicaePopularisSini鄄
HeTN,1988. Gentianaceae [A].
W. Liu & T.N. He, comb. nov. cae
Tom. 62 [M]. Beijing:SciencePress
Swertia pleurogynoides Novon 4
Baker, Bull. Misc. In鄄 HeTN,1994. NewtaxaofGentianaceaefromChina [J]. , :
1898
form., Kew :158 (1898). Type:Malawi (Nyasa鄄 369—372
Swertia Ac鄄
HeTN,LiuSW,1980. Newtaxaof L. fromChina [J].
land), between Kondow and Karonga,2000-6000ft.,
taPhytotaxonomicaSinica 18
, :75—85
Whyte s.n. (K! —holotype; BM! —isotype).
Swertia heterosepala HeTN,Pringle JS,1995. Gentianaceae [A]. In Wu CY, Raven
Gilg, Notizbl. Bot. Gart. Flora of China
PH(ed.), [M]. Beijing: Science Press; St.
14
16
Berlin : 115 (1938). Type: Tanzania, Matengo Louis:MissouriBotsnicalGarden, :1—140
highland, WSW from Songea, 1 100-1 200 m, 18 HeTN,XueCY,WangW,1994. The origin, dispersal and forma鄄
Swertia
tion of the distribution pattern of L. (Gentianaceae)
April 1936, H. Zerny 604 (K! —isotype).
ActaPhytotaxonomicaSinica 32
L. rotatum [J]. , :525—537
It is similar to in leaves and calyx
LiuJQ,ChenZD,LuAM,2001. Apreliminaryanalysisofthephy鄄
lobes linear, calyx as long as or rarely longer or
logenyoftheSwertiinae(Gentianaceae) basedonITSdata [J].
IsraelJournalofPlantScience 49
slightly shorter than corolla, but differs by corolla , :301—308
Swertia
smaller, 7-8.5 mm long. NemomissaS,1998. A synopsis of (Gentianaceae) in east
Kew Bulletin 53
Distribution: East Africa (Ethiopia, Tanzani鄄 andnortheast tropical Africa [J]. , : 419—
436,http://www.jstor.org/stable/4114507
a), South Africa (Mozambique, Malawi). Wet Swertiae
PissjaukovaVV, 1979. Generis L. (Gentianaceae) taxa no鄄
grassland, shire highland; 600-1800 m. NovostiSistematikiVysshikhRastenii 15
va,1 [J]. , :202—203
SystematicsandNat鄄
StruweL,AlbertVA,2002. Gentianaceae [A].
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: [M]. Cambridge:CambridgeUniversityPress,652
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[J]. YuanYM,K俟pfer P,1995. Molecular phylogenetics of the subtribe
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