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 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 文献标识码:A 文章编号:2095－0845(2013)03－386－07
Nomenclatural Novelities in Swertia (Gentianaceae)
HE Ting-Nong，LIU Shang-Wu，CHEN Shi-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
Swertia is one of the large genera in the family
Gentianaceae (Struwe and Albert，2002)． This ge-
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-
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-
lyses based on molecular data suggested that this ge-
nus is not monophyletic and some of the paralleling
clades from the radiative diversification of some spe-
cies clustered with the multiple related genera
(Yuan and Küpfer，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-
lutions due to the radiative diversifications or repre-
sented the ancestral states of the total family (Hagen
and Kadereit，2002)． As inferred from molecular
phylogenies (e． g． Yuan and Küpfer，1995)and
embryological studies (Xue et al．，1999a，b) ，we
found that sect． Heteranthos of this genus can be ex-
植 物 分 类 与 资 源 学 报 2013，35 (3):386 ～ 392
Plant Diversity and Resources 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-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-
tus based on its recognizable morphological distinct-
ness:dimorphic plants and flowers and a cleistoga-
mous pollination mode． However，up to now，mor-
phological traits，which can be used to distinguish
the other paralleling clades，need further examina-
tions． Therefore，we tentatively follow the traditional
generic circumscription of Swertia for the further in-
frageneric classification． Through examining thou-
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-
ries． Among them，sections Montana and Echinula-
ta，and series Repentes，Kilimandscharicae，Corym-
bosae，Japonicae，Swertopsis，Pumilae，and Abys-
sinicae were newly proposed for the first time． In ad-
dition，some taxonomic entities established previous-
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-
ograph，phylogenetic analyses and resource surveys
related to this genus．
1 Subgen． Swertia
1． 1 Sect． Apterae V． V． Pissjaukova
1． 1． 1 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-
neata by flowers large:corolla 20－30 mm in diam．，
lobes 15－20 mm． Var． cuneata has small flowers:co-
rolla 8－10 mm in diam．，lobes 7． 5－10(－12)mm． The
species Swertia cuneata is characterized by calyx lobes
oblong-spathulate to elliptic， base narrowed， apex
round and nectaries linear-oblong，radially elongated．
Distribution:China (SW Xizang)．
2 Subgen． Poephila C． B． Clarke
2． 1 Sect． Frasera (Walter)Knoblauch
2． 1． 1 Swertia albicaulis var． modocensis (St．
John)T． N． He，comb． nov．
Swertia modocensis St． John，Amer． Midl． Natu-
ralist 26:18 (1941)． Type:United States，Califor-
nia，Lassen County，Madeline，gentle northeast fa-
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-
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-
mer． Midl． Naturalist 26:19 (1941)． Type:Uni-
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-
list 26:16 (1941)． Type:United States，Califor-
nia，Lassen County，Madeline， about two miles
north of，gentle northeast facing slope，in rocky vol-
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-
as Creek Ranger Station，Lake County，1 570 m，
June 29－30，1911，W． W． Eggleston 5098 (NY!，
US!) ;Stein’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 HE Ting-Nong et al．:Nomenclatural Novelities in Swertia (Gentianaceae)
Leiberg 53 (US!)．
Swertia shastaensis St． John，Amer． Midl． Na-
turalist 26:19 (1941)． Type:United States，Cali-
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-
cludes six varieties，in having opposite stem leaves
with white membranous margin，a narrow interrupted
spike-like panicle，and oblong boat-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-
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-
sickii plants glabrous except for leaves puberulent at
base and sometimes along midvein of abaxial sur-
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-
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-
fornia)．
2．2 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-，raro 4-meri． Nectarium 1 in
unoquoque lobo corollae． Stylus brevis usque ab-
sens． Semina longitudinaliter，dense et tenuiter cor-
rugata vel corrugato-porcata． Pollinis granula admo-
dum striata (sparse et tenuiter striata，striato-perfora-
ta，striato-microreticulata usque striato-reticulata)．
Perennials，with slightly fleshy to semiwoody
tap-root and some rootlets，or tap-root absent but
stems trailing or stoloniferous with adventitious
roots． Flowers 5-，rarely 4-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-
prolate;medium-，rarely small- or large-sized，size
range 23． 3－43． 8 ×19． 0－33． 7 μm;colpi long or me-
dium． The sexine ornamentation is essentially striate，
varying from thinly，undulately striate ＆ perforate to
imperforate． The lira width is 0． 17 － 0． 30 μm．
Chromosome number 2n=26，rarely 2n=20．
This section is characterized by perennials，
with slightly fleshy to semiwoody tap-root or with ad-
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-
rugate- ridged and flowers usually 5-merous whereas
the latter having seed coats almost smooth and some-
times winged at one or both end，flowers always 4-
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-
ferent from sect． Montana by having an abbreviated
rhizome，2 nectaries per corolla lobe，and seeds lon-
gitudinally few (8 －12)corrugate-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 ser． Repentes T． N． He ＆ S．W． Liu，ser． nov．
Caulis repens vel stoloniformis，radices adven-
titias emittens． Flores solitarii vel in inflorescentiam
parco-florigeram dispositi． Nectarium planum，mac-
883 植 物 分 类 与 资 源 学 报 第 35 卷
uliforme，absque marginibus et fimbriis．
Stems trailing or stoloniferous with adventitious
roots． Basal leaves crowded into a rosette． Flowers
5-merous (but 4-merous in S． subnivalis and S． sar-
mentosa) ，solitary on long terminal pedicels or in
few-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 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 Léveillé
(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-
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 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-forme，fimbriatum．
Stems erect，without stolons and adventitious
roots． Inflorescences a many-flowered panicle of cymes．
Nectary V-shaped，fimbriate．
Type species:Swertia kilimandscharica Engler
Distribution: from North Africa to Northeast
and Tropical Africa．
5 species．
3 Subgen． Ophelia(D． Don ex Grisebach)C．B． Clarke
3．1 Sect． Ophelia (D． Don ex Grisebach)Bentham
et Hooker f． ex Gilg
3． 1． 1 Ser． Corymbosae T． N． He ＆ S． W． Liu，
ser． nov．
Inflorescentia corymbosa． Nectarium 1 in uno-
quoque lobo corollae，prope basin lobi corollae，par-
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-
rolla lobe，near the base of corolla lobe，smaller，or-
bicular to suborbicular，dish-shaped without or with
short raised scale，the bound with many long papil-
lose fimbriae round margin，or watch-packet-shaped
with an orbicular covered scale and many short papil-
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-
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 Sect． Swertopsis (Makino)Satake
3． 2． 1 Ser． Japonicae T． N． He ＆ S． W． Liu
(subgen． Ophelia sect． Swertopsis) ，ser． nov．
Herba annua． Inflorescentia cymoso-panicula．
Flores 5-，minus 4-meri． Nectaria 2，raro 1 in uno-
quoque lobo corollae，oblongum，naviculare vel sac-
ciforme． Semina tenuiter reticulata．
Annuals． Inflorescences a panicle of cymes．
Flowers 5-，less 4-merous． Nectaries 2，rarely 1 per
corolla lobe，oblong，boat-shaped，or pocket-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-
9833 期 HE Ting-Nong et al．:Nomenclatural Novelities in Swertia (Gentianaceae)
kim，Bhutan，Bengal，almost throughout China，My-
anmar，Thailand，Korea，Japan，and E New Guinea．
31 species．
3． 2． 1． 1 Swertia tetrapetala Pallas
1a． Leaves linear to oblong，10－20(－30)×(2－)3－5(－10)
mm． Apex of corolla lobes obtuse，rarely acuminate
109a． var． tetrapetala………………………………
1b． Leaves ovate-triangular，20 －35 ×(12 －)15 －20(－23)
mm． Apex of corolla lobes acute and mucronate
109b． var． wilfordii…………………………………
109a． var． tetrapetala
Distribution:E Russia (Sachalin，Kamchat-
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-
east Region (Mandschuria) ，44° － 45° 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-triangular，20－35×(12－)15－20
(－23)mm． Apex of corolla lobes acute and mucro-
nate． Flowering and fruiting:August—September．
Distribution:China (Northeast Region) ，NE
Korea． ca． 670 m．
3． 2． 2 Ser． Swertopsis T． N． He ＆ S． W． Liu，
ser． nov．
Herba annua． Folia ovato-elliptica usque rhom-
biformia，apice acuminata caudataque． Inflorescen-
tia in terminales，axillares fasciculatas cymas vel
umbellas disposita． Flores 5-meri． Nectaria 2 in un-
oquoque lobo corollae，sacciformia． Semina tenuiter
reticulata．
Annuals． Leaves ovate-elliptic to rhombic，a-
pex acuminate and caudate． Inflorescences terminal
and axillary fasciculate cymes or umbels． Flowers 5-
merous． Nectaries 2 per corolla lobe， pocket-
shaped． Seeds finely reticulate．
This series is more special in sect． Swertopsis
by inflorescences terminal and axillary fasciculate
cymes or umbels and leaves ovate-elliptic to rhom-
boid，apex acuminate and caudate．
Type species:Swertia swertopsis Makino
Distribution:Japan．
3． 2． 3 Ser． Dichotomae T． N． He ＆ S． W． Liu
3． 2． 3． 1 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 83-011 (HIMC—ho-
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-
ered scale and a small corner on back of the scale，
but differs by corolla orange-colored，with red stria-
te，leaves rugose，with crisp margin．
1a． Corolla white，sometimes with purple spots;leaves level
var． dichotoma………………………………………
1b． Corolla orange-colored，with red striate;leaves rugose，
with crisp margin var． rubrostriata…………………
Distribution:China (Nei Mongol)．
3． 3 Sect． Platynema T． N． He ＆ S． W． Liu
3． 3． 1 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 植 物 分 类 与 资 源 学 报 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×1． 5 －2． 5
mm，calyx lobes 4． 5－5×0． 5－1 mm，corolla lobes
to 5． 5 ×4 mm． Var． paniculata is characterized by
the corolla with 2 purple or yellow-green spots above
each nectary and plants relatively robust:stems 35－
120 cm tall，leaves 2 －6． 5 cm × 4 －7 mm，calyx
lobes 5－5． 5×1． 5－2． 5 mm，and corolla lobes 6－
6． 5× 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-shaped and naked necta-
ries，and dilated and free filaments．
Distribution:Nepal．
3． 4 Sect． Echinulata T． N． He ＆ S． W． Liu，
sect． nov．
Herba perennis vel annua． Nectaria 2 in uno-
quoque lobo corollae，oblonga，navicularia，fimbria-
ta． Stylus absens;stigmate orbiculato． Semina par-
va，subtiliter reticulata． Pollinis granula echinulato
et saepe perforato exinio ornata．
Perennials or annuals． Nectaries 2 per corolla
lobe，oblong，boat-shaped，with a narrow scale and
raised fimbriate margin． Style absent;stigma orbicu-
lar． Seeds finely reticulate． Pollen grains are sub-
spheroidal，but subprolate and suboblate grains oc-
cur in infraspecific variation;medium-sized with the
size range of 22． 2 －27． 3 ×19． 6 －29． 3 μm;colpi
medium，long or short． The sexine ornamentation is
echinulate and perforate but imperforate in S． usam-
barensis． All perforations are very small．
Sect． Echinulata is somewhat similar to sect． swer-
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 Ser． Pumilae T． N． He ＆ S．W． Liu，ser． nov．
Herba perennis． Radix crassa，fibrosa usque semil-
ignosa． Caules e basi multum ramosi． Folia basalia sa-
epe abundantia，persistentia et in rosulam conferta．
Perennials． Taproot stout，fibrous to semiwoody．
Stems much branched from very base． Basal leaves u-
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-
go，Ethiopia，Kenya，Tanzania，and Uganda)and
West Asia (N． Yemen and SW． Saudia) ，
9 species．
3． 4． 2 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-
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 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-
mosome numbers from the whole African and Wes-
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-
charica，sect． Echinulata:ser． Pumilae and ser．
Abyssinicae)respectively． Both sect． Montana and
sect． Echinulata differ from each other as follows:
1933 期 HE Ting-Nong et al．:Nomenclatural Novelities in Swertia (Gentianaceae)
1a． Nectary1 per corolla lobe;seeds corrugate;pollen grains essentially striate，varying from irregularly striate-reticulate，striate-
microreticulate to striate-perforate or scattered and thinly striate-imperforate (Sect． Montana) :
2a． Nectary reduced to a flat viscous gland patch without raised margin and fimbriae;stems trailing or stoloniferous with ad-
ventitious roots at nodes;flowers solitary on long terminal pedicels or in few-flowered terminal cymes
Ser． Repentes…………………………………………………………………………………………………………
2b． Nectary fimbriate;stems erect，without stolons and adventitious roots;inflorescences a many-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-
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………………………………………………………………………………………………………
In addition，there is an African species of Lom-
atogonium as follows:
Lomatogonium pleurogynoides (Baker) S．
W． Liu ＆ T． N． He，comb． nov．
Swertia pleurogynoides Baker，Bull． Misc． In-
form．，Kew 1898:158 (1898)． Type:Malawi (Nyasa-
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-
a) ，South Africa (Mozambique，Malawi)． Wet
grassland，shire highland;600－1 800 m．
References:
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Systematics ＆ Evolution，229:1—21
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［J］． Notizblatt des Botanischen Gartens und Museums zu Berlin-
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293 植 物 分 类 与 资 源 学 报 第 35 卷