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Guihaia Jan. 2016，36 (1)：30-43
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DOI: 10.11931/guihaia.gxzw201510002
岑佳梦，金水虎，金孝锋. 中国薹草属（莎草科）研究随记（IV）：隐匿薹草组的分类修订［J］. 广西植物，2016，36(1)：30-43
CEN JM，JIN SH，JIN XF. Notes on Carex (Cyperaceae) from China (IV): the identity and revision of sect. Infossae［J］. Guihaia，2016，36(1)：30-43

Notes on Carex (Cyperaceae) from China
(IV): the identity and revision of sect. Infossae
CEN Jia-Meng1， JIN Shui-Hu2， JIN Xiao-Feng1*
( 1. College of Life & Environmental Sciences，Hangzhou Normal University，Hangzhou 310036，China; 2. School of
Foresrty and Bio-technology，Zhejiang Agricultural and Forestry University，Lin’an 311300，China )

Abstract: Carex sect. Infossae S. W. Su was recently described from Anhui and Jiangsu provinces of East China，
containing two species with one variety： C. infossa var. infossa，C. infossa var. extensa and C. minuticulmis. According
to the sectional description，fifteen additional species and two varieties should be placed in this section. Based upon lit-
erature survey，specimen examination，field work，and SEM observation of perigynia and achenes，a concise table of
different treatments between Florae Reipublicae Popularis Sinicae and Flora of China was provided，as well as the
morphological characters distinguishing these taxa and micromorphology of perigynia and achenes for the selected taxa
were compared. Among the diagnostic characters，arrangement of spikes and perigynium indumentum are inseparable
from each other，and two species in sect. Infossae are consequently recognized in the present paper. Carex elmeri，C.
granifera，C. minuticulmis，C. shanghaiensis，C. blinii subsp. shanghaiensis，C. sublateralis，C. taihokuensis and C.
tatsutakensis were synonymized to C. blinii. Carex canaliculata，C. changmuensis，C. distantiflora，C. infossa，C. infossa
var. extensa，C. lateralis，C. loheri，C. longerostrata var. exaristata，C. longerostrata var. hoi，C. lyi，C. macrandrolepis，
C. sharyotensis were reduced as synonyms to C. oxyphylla.
Key words： Carex sect. Infossae，scanning electron microscopy，morphological character，new synonym，China
CLC number： Q949.71 Document code： A Article ID: 1000-3142(2016)01-0030-14

中国薹草属（莎草科）研究随记
（IV）：隐匿薹草组的分类修订
岑佳梦 1, 金水虎 2, 金孝锋 1*
（1. 杭州师范大学 生命与环境科学学院，杭州 310036; 2. 浙江农林大学 林业与生物技术学院，浙江 临安 311300）

摘 要：隐匿薹草组(Carex sect. Infossae)为最近建立的薹草属(Carex)的组，当时报导分布于我国安徽和江苏，
有 2种和 1变种，即隐匿薹草(C. infossa)、显穗薹草(变种) (C. infossa var. extensa)和矮秆薹草(C. minuticulmis)。
根据描述，另有 15种和 2变种应归属本组，这些分类群具有较为一致的特征：果囊椭圆球状卵球形或卵球形，
通常疏被短毛或近无毛，先端渐狭成中等长的喙，喙口具 2 小齿；瘦果卵球形，先端钝或微凹。通过文献考
证、标本查阅、野外采集，并结合果囊和瘦果的扫描电镜观察，对隐匿薹草组所涉及的种（含种下类群）进
行了分类修订。该组仅有 2种：百里薹草（C. blinii）和尖叶薹草（C. oxyphylla）。将 Carex elmeri、C. granifera、

收稿日期：2015-10-06 修回日期：2015-12-23
基金项目：国家自然科学基金（31170183，31470304）；国家科技部科技基础性工作专项（2013FY112100）；国家自然科学基金委重大国际合作项
目（31110103911）[Support by the National Natural Science Foundation of China (31170183，31470304); Science and Technology Basic Work
(2013FY112100); Major International Cooperation Project of the National Natural Science Foundation of China (31110103911)]。
作者简介：岑佳梦(1994-)，女，浙江慈溪人，主要从事植物分类学研究，（E-mail）1455722086@qq.com。
*通讯作者：金孝锋，博士，教授，从事植物分类和系统进化方面的研究，(E-mail) docxfjin@163.com。
1期 岑佳梦等：中国薹草属（莎草科）研究随记（IV）：隐匿薹草组的分类修订 31


矮秆薹草（C. minuticulmis）、上海薹草（C. shanghaiensis C. blinii subsp. shanghaiensis）、C. sublateralis、C.
taihokuensis 和锐果薹草 (C. tatsutakensis)并入百里薹草；将沟囊薹草 (C. canaliculata)、樟木薹草 (C.
changmuensis)、C. distantiflora、隐匿薹草(C. infossa)、显穗薹草(C. infossa var. extensa)、C. lateralis、C. loheri、
无芒长嘴薹草(C. longerostrata var. exaristata)、城湾薹草(C. longerostrata var. hoi)、C. lyi、和平菱果薹草(C.
macrandrolepis)和 C. sharyotensis并入尖叶薹草。
关键词：隐匿薹草组，扫描电镜，形态学性状，新异名，中国

Su (2009) established Carex sect. Infossae S. W. Su，
designating C. infossa Z. P. Wang as the type. The section as
originally described and contained two species and one va-
riety: C. infossa Z. P. Wang，C. infossa var. extensa S. W.
Su，and C. minuticulmis S. W. Su & S. M. Xu (Su，1996，
2009; Su & Xu，1988). According to Su’s delimitation，the
section is morphologically similar to sect. Radicales (Kük.)
Nelmes and sect. Clandestinae G. Don. It differs from the
former in having spikes unisexual. It differs from the latter
in having involucral bracts leaf-like，longer than spikes，
shortly sheathed; achenes emarginate at apex; and styles
thickened at base but caducous (Su，2009).
China is incredibly rich in Carex，with more than 500
species (Dai et al，2000，2010). Because of scattered
preservation of the literature and specimens，the taxonomic
treatment of Carex in China was not in a satisfactory status，
and extensive examination of specimens and field collection
of new material are needed. Morphological characters of
perigynia and achenes under a 10× hand lens can be used in
species identification and traditional taxonomy of Carex
(Ohwi，1936; Nelmes，1951; Dai et al，2000，2010; Jin &
Zheng，2013). Micromorpholoy of perigynia and achenes
under SEM provided evidences for traditional taxonomy in
both interspecific and intraspecific levels，as well as sec-
tional delimitation (Toivonem & Timonen，1976; Menapace
& Wujek，1987; Waterway，1990; Jin & Zheng，2010，
Jin et al, 2013).
Based on specimen examination，we found several
taxa with achenes similar to those of sect. Infossae，most
distributed in China. They were: Carex blinii H. Lév. &
Vaniot，C. canaliculata P. C. Li，C. changmuensis Tang & F.
T. Wang ex Y. C. Yang，C. distantiflora Nakai，C. elmeri
Kük.，C. granifera Dunn，C. lateralis Kük.，C. loheri C. B.
Clarke，C. longirostrata C. A. Meyer var. exaristata X. F.
Jin & C. Z. Zheng，C. longirostrata var. hoi Chü ex S. Yun
Liang，C. lyi H. Lév.，C. macrandrolepsis H. Lév.，C.
oxyphylla Franch.，C. shanghaiensis S. X. Qian & Y. Q.
Liu，C. sharyotensis Hayata，C. sublateralis T. Koyama，
C. taihokuensis Hayata and C. tatsutakensis Hayata
(Franchet，1898; Clarke，1904; Léveillé，1906，1907，
1908; Dunn，1908; Kükenthal，1909，1910; Nakai，1914;
Hayata，1916，1921; Koyama，1956; Yang，1987; Li，
1990; Liang，1998; Liu and Qian，1999; Jin et al，2004). All
these above-mentioned taxa have the consistent characters
of perigynia and achenes: perigynia ellipsoid-obovoid to
ovoid，or nearly globose，obtusely trigonous，sparsely
pubescent or nearly glabrous，apex gradually contracted into
a beak; beaks mid-sized in length; orifice 2-lobed with min-
ute teeth; achenes frequently ovoid，trigonous，excavate or
obtuse at apex，stipitate at base. According to the shapes of
perigynium and achene，these species and varieties should
be placed in sect. Infossae.
Although taxonomic revisions of the above-mentioned
taxa were published in Flora Reipublicae Popularis Sinicae
(Vol. 12) and Flora of China (Vol. 23)，species treatment or
their sectional placement were problematic (Dai et al，2000，
2010). Important differences between the species treatments
of Flora Reipublicae Popularis Sinicae (Vol. 12) and Flora
of China (Vol. 23) are shown in Table 1.
The present study of the taxonomic status of the spe-
cies in sect. Infossae is based on: (i) carefully checking and
critical examination of all available type specimens; (ii)
observations of morphological variation within and among
populations of the related species/variety; (iii) observations
of variation in perigynium indumentum; and (iv) observa-
tions of achene shape and sexine ornamentation of the se-
lected taxa.
1 Materials and Methods
1.1 Examination of type specimens
Type specimens of these taxa were examined and
32 广 西 植 物 36卷

Table 1 Different treatments of the species in Carex sect. Infossae between Flora
Reipublicae Popularis Sinicae and Flora of China
Flora Reipublicae Popularis Sinicae (Dai et al，2000) Flora of China (Dai et al，2010)
sect. Clandestinae G. Don
Carex infossa C. P. Wang
C. infossa var. extensa S. W. Su
sect. Clandestinae G. Don
C. infossa C. P. Wang
C. infossa var. extensa S. W. Su
C. minuticulmis S. W. Su & S. M. Xu
sect. Rhomboidales Kükenthal (1909: 622)
C. oxyphylla Franch.
C. blinii H. Lév. & Vaniot (doubtful)
sect. Rhomboidales Kük.
C. oxyphylla Franch.
C. blinii H. Lév. & Vaniot
C. longirostrata var. exaristata X. F. Jin & C. Z. Zheng
C. longirostrata var. hoi Chü ex S. Yun Liang
C. macrandrolepsis H. Lév. & Vaniot
syn. C. sharyotensis Hayata
C. blinii ssp. shanghaiensis (S. X. Qian & Y. Q. Liu) S. Yun Liang & T.
Koyama
C. tatsutakensis Hayata
syn. C. taihokuensis Hayata
sect. Careyanae Tuckerman ex Kükenthal (1909: 679)
C. longirostrata C. A. Mey. var. hoi Chü ex S. Yun Liang
C. macrandrolepsis H. Lév. & Vaniot
syn. C. sharyotensis Hayata
C. shanghaiensis S. X. Qian & Y. Q. Liu
C. tatsutakensis Hayata
syn. C. taihokuensis Hayata
C. canaliculata P. C. Li
sect. Careyanae Tuckerm. ex Kük.
C. canaliculata P. C. Li
C. changmuensis Tang & F. T. Wang ex Y. C. Yang

species not treated
C. changmuensis Tang & F. T. Wang ex Y. C. Yang
C. granifera Dunn
C. lyi H. Lév. & Vaniot
C. minuticulmis S. W. Su & S. M. Xu
C. sublateralis T. Koyama
species not treated
C. granifera Dunn
C. lyi H. Lév. & Vaniot
C. sublateralis T. Koyama

carefully checked. Although different taxonomists placed
these taxa in different sections，the above-mentioned taxa
are very similar to each another. Our observations of type
specimens focused on all diagnostic characters，such as
bract sheath，arrangement of spikes，and perigynium in-
dumentum (Table 2).
1.2 Observations on morphological characters
To observe morphological character variation within and
among populations both in Carex blinii and C. infossa，
collections from five different populations were checked as
well. Among these populations，specimens of four popula-
tions were collected by the authors and those of the popula-
tion，Chuzhou of Anhui，were checked from the herbarium of
ACM and determined as C. infossa. The collections from the
other four populations were all determined as C. blinii.
Twenty-eight sheets representing as many individuals were
obtained，and six individuals were collected from the popu-
lation at Chuzhou. Of the others，five individuals were from
the population at Hangzhou，five from Libo，and six from
each of Lin’an and Yueqing (Table 3).

1.3 Scanning electron microscopy (SEM) of perigynium
indumentum and achene for selected taxa
Mature perigynia were gathered from 15 specimens
housed in ACM，HTC，HZU，KYO，N，P，PE and TI，
representing eight species and three varieties (Appendix).
Perigynia were submerged in 50% ethanol to clean for 2
h，then air dried. For SEM，cleaned perigynia were
mounted on stubs using doubled-sided adhesive tape and
coated with gold.
Twelve achene samples representing seven species
and two varieties were gathered from specimens deposited
in ACM，HTC，KYO，N and TI (Appendix). Mature
achenes were initially soaked in a solution of concentrated
sulfuric acid and acetic anhydride (volume ratio=1:9) for
18 h，then rinsed in acetic acid for 10 min and water for 5
min，and placed in a bath-type ultrasonic cleaner for 30
min with 70% ethanol to remove the cuticle and outer
periclinal wall of epidermis (Oda et al，2003; Zhang，
2006). After air drying，the achenes were mounted on
stubs using double-sided adhesive tape，and directly
1期 岑佳梦等：中国薹草属（莎草科）研究随记（IV）：隐匿薹草组的分类修订 33

Table 2 Observation on type specimens of the species in Carex sect. Infossae
Species name Specimen Bract sheath Arrangement of spike
Perigynium indumentums
density
Carex blinii J. Esquirol 343 (lectotype) Almost not-sheathed Crowded，nearly capitate Sparsely pubescent (1)
C. canaliculata Qinghai-Xizang Exped. 6584
(holotype)
Sheathed，0.5-1.5 cm long Terminal two crowded，
lowers remote
Sparsely pubescent (1)
Qinghai-Xizang Exped. 6584
(isotype)
Sheathed，0.6-0.9 cm long Terminal two crowded，
lowers remote
Sparsely pubescent (1)
Qinghai-Xizang Exped. 6584
(isotype)
Sheathed，0.7-1.2 cm long Terminal two crowded，
lowers remote
Sparsely pubescent (1)
Qinghai-Xizang Exped. 6584
(isotype)
Sheathed，0.5-1.2 cm long Terminal two crowded，
lowers remote
Sparsely pubescent (1)
C. changmuensis Y. T. Chang & K. Y. Lang 3227
(holotype)
Short-sheathed，0.3-0.5 cm
long
Terminal two crowded，
lowers remote
Sparsely pubescent (1)
C. distantiflora T. Nakai 1332 (lectotype) Sheathed，1-2.5 cm long Remote Sparsely pubescent (1)
T. Nakai 941 (syntype) Sheathed，0.7-3 cm long Terminal two crowded，
lowers remote
Sparsely pubescent (1)
Y. Tsujikuma s. n. (syntype) Short-sheathed，0.4-0.8 cm
long
Remote Sparsely pubescent (1)
C. elmeri A. D. E. Elmer 8444 (holotype) Short-sheathed，ca. 0.3 cm
long
Crowded，nearly capitate Sparsely pubescent (1)
C. granifera HK Herb. 3655 (holotype) Almost not-sheathed Crowded，nearly capitate Nearly glabrous (0)
C. infossa C. P. Wang 76 (holotype) Short-sheathed，0.3-0.7 cm
long
Terminal two crowded，
lowers remote
Sparsely pubescent (2)
C. P. Wang 76 (isotype) Short-sheathed， 0.4-1 cm
long
Terminal two crowded，
lowers remote
Sparsely pubescent (2)
C. infossa var.
extensa
S. W. Su 82005 (isotype) Sheathed，0.7-1.5 cm long Remote Sparsely pubescent (1)
C. lateralis anonymous 11061 (holotype) Short-sheathed，0.4-0.7 cm
long
Terminal two crowded，
lowers remote
Sparsely pubescent (2)
C. loheri A. Loher 701 (holotype) Sheathed，0.5-2 cm long Terminal two crowded，
lowers remote
Sparsely pubescent (2)
C. longirostrata
var. exaristata
Q. C. Chen & J. H. Zhou 1879
(holotype)
Sheathed，1-2 cm long Remote Nearly glabrous (0)
Q. C. Chen & J. H. Zhou 1879
(isotype)
Sheathed，ca. 1.5 cm long Remote Nearly glabrous (0)
C. longirostrata
var. hoi
Y. Y. Ho 943 (holotype) Short-sheathed，0.5-0.8 cm
long
Remote Sparsely pubescent (1)
C. lyi J. Cavalerie 2329 (holotype) Short-sheathed to sheathed，
0.3-2 cm long
Terminal two crowded，
lowers remote
Sparsely pubescent (1)
J. Cavalerie 2329 (isotype) Sheathed，0.6-1.5 cm long Terminal two crowded，
lowers remote
Sparsely pubescent (1)
C. macrandrolepsis U. Faurie 2285 (isotype) Sheathed，0.5-1.5 cm long Remote Sparsely pubescent (1)
C. minuticulmis S. W. Su 2477 (holotype) Almost not-sheathed Crowded，nearly capitate Sparsely pubescent (3)
C. oxyphylla Delavey 2524 (holotype) Short-sheathed to sheathed，
0.3-1.8 cm long
Terminal two crowded，
lowers remote
Sparsely pubescent (1)
Delavey 2524 (isotype) Short-sheathed to sheathed，
0.4-1.5 cm long
Terminal two crowded，
lowers remote
Sparsely pubescent (1)
C. shanghaiensis Y. Q. Liu & S. X. Qian 63
(holotype)
Almost not-sheathed Crowded，nearly capitate Sparsely pubescent (3)
C. sharyotensis B. Hayata s. n. (holotype) Short-sheathed to sheathed，
0.5-1.3 cm long
Remote Nearly glabrous (0)
C. sublateralis K. Kimura s. n. (holotype) Almost not-sheathed Crowded，nearly capitate Sparsely pubescent (3)
C. taihokuensis U. Faurie s. n. (holotype) Almost not-sheathed Crowded，nearly capitate Sparsely pubescent (1)
U. Faurie s. n. (isotype) Almost not-sheathed Crowded，nearly capitate Sparsely pubescent (1)
C. tatsutakensis W. R. Price 797 (holotype) Almost not-sheathed Crowded，nearly capitate Sparsely pubescent (2)
Note: the numbers in brackets denote the gradual status, with 0 expressing nearly glabrous, and 1, 2, 3 expressing gradual status.
34 广 西 植 物 36卷

Table 3 Observation on individuals/specimens from different populations
Population name Specimen Bract sheath Arrangement of spike Perigynium indumentum
Chuzhou of Anhui，
China
S. W. Su 81065 [2]* Sheathed，0.6-1.5 cm long Terminal two crowde，lowers
remote/ or remote
Sparsely pubescent (1)**
Carex infossa Anonymous s. n. [4] Sheathed，0.5-2 cm long Terminal two crowde，lowers
remote/ or remote
Sparsely pubescent (2)
Hangzhou of
Zhejiang，China
H. Wang & X. F. Jin 1444 [2] Almost not-sheathed Crowded，nearly capitate Sparsely pubescent (2)
Carex blinii H. Wang & X. F. Jin 1446 [3] Almost not-sheathed Crowded，nearly capitate Sparsely pubescent (2)
Libo of Guizhou，
China
X. F. Jin et al 2388 [2] Almost not-sheathed Crowded，nearly capitate Sparsely pubescent (1)
Carex blinii X. F. Jin et al 2390 [3] Almost not-sheathed Crowded，nearly capitate Sparsely pubescent (1)
Lin’an of Zhejiang，
China
X. F. Jin & Y. Y. Zhou 3050 [3] Almost not-sheathed Crowded，nearly capitate Sparsely pubescent (2)
Carex blinii X. F. Jin & Y. Y. Zhou 3051 [3] Almost not-sheathed Crowded，nearly capitate Sparsely pubescent (2)
Yueqing of
Zhejiang，China
X. F. Jin & P. J Cao 1529 [3] Almost not-sheathed Crowded，nearly capitate Nearly glabrous (0)
Carex blinii X. F. Jin & P. J Cao 1533 [3] Almost not-sheathed Crowded，nearly capitate Nearly glabrous (0)
Note: * the numbers in square brackets denote observed individuals or specimens; ** the numbers in brackets denote the gradualstatus, with 0 expressing
nearly glabrous, and 1, 2, 3 expressing gradual status.

coated with a layer of gold.
The coated perigynia and achenes were observed and
photographed under a Philips XL-30E scanning electron
microscope.

2 Results and Analysis

2.1 Morphological variations on type specimens
Based on the observation on type specimens (Fig. 1，
2)，sheaths of involucral bracts，arrangement of spikes and
perigynium indumentum are shown in Table 2. The bract
sheaths of these taxa indicate: (I) almost sheathless，e.g.
Carex blinii ， C. granifera ， C. minuticulmis ， C.
shanghaiensis，C. sublateralis，C. taihokuensis and C.
tatsutakensis; (II) short-sheathed，e.g. C. changmuensis，
C. elmeri，C. lateralis，C. infossa and C. longirostrata var.
hoi; (III) sheathed，e.g. C. canaliculata，C. infossa var.
extensa，C. loheri and C. longirostrata var. exaristata. The
bract sheaths of C. distantiflora shows short-sheathed or
sheathed in different type specimens，while those of C. lyi，
C. oxyphylla and C. sharyotensis show short-sheathed to
sheathed in the same specimens.
Spikes of the above-mentioned taxa arranged as the
following patterns: (I) all spikes crowded，nearly capitate，
e.g. Carex blinii，C. elmeri，C. granifera，C. minuticulmis，
C. shanghaiensis，C. sublateralis，C. taihokuensis and C.
tatsutakensis; (II) terminal two spikes crowded，lowers
remote，e.g. C. canaliculata，C. changmuensis，C. infossa，
C. lateralis，C. loheri，C. lyi and C. oxyphylla; (III) all
spikes remote，e.g. C. infossa var. extensa，C. longirostrata
var. exaristata，C. longirostrata var hoi，C. macrandrolepsis
and C. sharyotensis. Among three syntypes of C.
distantiflora，one specimen (T. Nakai 941) shows the distal
two spikes arranged crowded，proximal spikes remote. The
other two specimens show all spikes remote (Table 2).
Most of the taxa have sparsely pubescent perigynia，
but the indumentum ranges in density from nearly gla-
brous，such as Carex granifera，C. longirostrata var.
exaristata and C. sharyotensis，to more densely pubescent
in C. minuticulmis，C. sublateralis and C. shanghaiensis
(Table 2).
Noticeably，the description of the leaves of Carex
minuticulmis is pubescent in Flora of China (Dai et al，
2010)，whereas those in the protologue are glabrous (Su &
Xu，1988). The examination of the type specimen of C.
minuticulmis indicates all leaves are glabrous on both sur-
faces，which is consistent to the description in the
protologue.

1期 岑佳梦等：中国薹草属（莎草科）研究随记（IV）：隐匿薹草组的分类修订 35



Fig. 1 Types of nine taxa in Carex sect. Infossae A. Carex blinii (lectotype，P)； B. C. elmeri (holotype，K)； C. C. granifera (holotype，
K)； D. C. minuticulmis (holotype，AHU tranferred to ACM)； E. C. shanghaiensis (holotype，PE)； F. C. taihokuensis (holotype，TI)； G. C.
tastutakensis (holotype，TI)； H. C. longirostrata var. exaristata (holotype，HZU)； I. C. longirostrata var. hoi (holotype，PE).

2.2 Morphological variation within and among popula-
tions in Carex blinii and C. infossa
Bract sheath，arrangement of spikes and perigynium
indumentum were observed from different populations and
shown in Table 3. The bracts of the individuals of Carex
blinii from the populations Hangzhou，Libo，Lin’an and
Yueqing are inconspcuously sheathed，almost sheathless.
The spikes of the individuals in these populations are
crowded，nearly capitate. The perigynium indumentum of
the population Yueqing is nearly glabrous，and those of
Hangzhou，Libo and Lin’an are sparsely pubescent but
with different gradual status of density. The bracts of the
36 广 西 植 物 36卷



Fig. 2 Types of nine species in Carex sect. Infossae A. Carex oxyphylla (holotype，P)； B. C. changmuensis (holotype，PE)； C. C.
distantiflora (lectotype，TI)； D. C. infossa (holotype，N)； E. C. canaliculata (holotype，PE)； F. C. loheri (holotype，K)； G. C. lyi (holotype，
P)； H. C. macrandrolepsis (isotype，KYO)； I. C. sharyotensis (holotype，TI).

individuals of Carex infossa from the population Chuzhou
are sheathed. The lower spikes are remote，but terminal two
spikes sometimes crowded. The perigynium indumentum is
also sparsely pubescent with different gradual status of den-
sity. Two sheets of the specimens (S. W. Su 81065) have the
perigynia with very sparsely pubescence，but four sheets
(anonymous s. n.) are more densely pubescent.
2.3 Perigynium indumentum under SEM
The perigynium indumentum of eight species and three
varieties in Carex sect. Infossae were observed using SEM.
All samples show some degree of pubescence， and
perigynia of C. longirostrata var. exaristata，C. sharyotensis
1期 岑佳梦等：中国薹草属（莎草科）研究随记（IV）：隐匿薹草组的分类修订 37



Fig. 3 Perigynium indumentum of the taxa in Carex sect. Infossae under SEM A. Carex blinii (Sample 1)； B. C. blinii (Sample
2)； C. C. blinii (sample 3)； D. C. minuticulmis； E. C. taihokuensis； F. C. tatsutakensis； G. C. oxyphylla； H. C. infossa var. extensa； I. C.
infossa； J. C. infossa； K. C. longirostrata var. exaristata； L. C. longirostrata var. hoi； M. C. macrandrolepsis (sample 1)； N. C. macrandrolepsis
(sample 2)； O. C. sharyotensis. Scale bar: all=200 μm.

and C. taihokuensis are nearly glabrous. The samples from
different localities/populations for the same species show
sparse pubescence to varying degrees (Fig. 3)，which is
consistent with the observation on types or populations.
2.4 Achene shape and sexine ornamentation under SEM
The achene shape of the taxa in Carex sect. Infossae
isovoid，rarely obovoid or broadly ovoid，trigonous; beaks
absent，excavate rarely obtuse at apex; sides slightly con-
cave. The achene shapes of the three samples of C. infossa
range from obovoid in Jiangsu to ovoid in Anhui and
Zhejiang，(Fig. 4). The periclinal cells of all samples are
straight，with silica bodies generally one per cell，
occasionally two or three (Fig. 4).

3 Taxonmic Treatment

Based on morphological observations on the type
specimens and population collections，the diagnostic
38 广 西 植 物 36卷



Fig. 4 Achene micromorphology of the taxa in Carex sect. Infossae under SEM A，C，D，F，H，I，K–M，O–Q. Achene shape；
B，E，G，J，N，R. Sexine ornamentation. A，B. Carex blinii； C. C. minuticulmis； D，E. C. taihokuensis； F，G. C. tatsutakensis (Sample
1)； H. C. tatsutakensis (Sample 2)； I，J. C. infossa (sample 1)； K. C. infossa (Sample 2)； L. C. infossa (Sample 3)； M，N. C. longirostrata
var. exaristata； O. C. longirostrata var. hoi； P. C. macrandrolepsis (Sample 1)； Q，R. C. macrandrolepsis (Sample 2). Scale bars: A，C，D，
F，H，I，K–M，O–Q=2 mm； B，E，G，J，N，R=100 μm.

characters among these taxa are as follows: bract sheath，
arrangement of spikes，and perigynium indumentums.
These characters were carefully checked and evaluated. We
here treat the taxa in Carex sect. Infossa as two distinct spe-
cies，and use two prior names C. blinii H. Lév. & Vaniot and
C. oxyphylla Franch.，respectively.
1期 岑佳梦等：中国薹草属（莎草科）研究随记（IV）：隐匿薹草组的分类修订 39




Fig. 5 Carex blinii (A–F) and C. oxyphylla (G，H) A，G. Habit； B. Staminate scale； C，D. Pistillate scale； E. Perigynium； F.
Achene； H. Inflorescence. (A–F from X. F. Jin & al. 2390，HTC； G from H. Wang & X. F. Jin 1445，HTC； H from anonymous s. n.，ACM).

Carex sect. Infossae S. W. Su，J. Anhui Agric. Univ. 36:
562. 2009. TYPE: C. infossa Z. P. Wang (=C. oxyphylla
Franch.).
Perennial herbs. Culms central or lateral，trigonous.
Leaves longer than culms，or almost equal in length.
Involucral bract leaf-like or shortly leaf-like，shorter than
inflorescence，almost sheathless or short-sheathed to con-
spicuously sheathed. Inflorescence paniculate，spikes 2–5;
spikes crowded，or distal two spikes crowded，or all remote;
terminal spike staminate; lateral spikes pistillate. Perigynia
ellipsoid-obovoid，obovoid，ovoid or nearly globose (ex-
cluding beaks)，obtusely trigonous，distinctly or
40 广 西 植 物 36卷


Key to the species of Carex sect. Infossae.
1a. Culms lateral，<15 cm; involucral bracts almost not-sheathed; spikes crowded ………….Carex blinii H. Lév. & Vaniot
1b. Culms central，rarely lateral，(10–)20–40 cm tall; involucral bracts short-sheathed to conspicuously sheathed; spikes
remote，or distal two spikes crowded，proximal spikes remote ………….C. oxyphylla Franch.

inconspicuously veined，sparsely pubescent or nearly gla-
brous，apex gradually contracted into a beak; orifice 2-lobed
with minute teeth. Stigmas 3; style thickened at base.
Achenes tightly enveloped，ovoid， rarely obovoid or
broadly ovoid，trigonous，sides slightly concave，excavate
or obtuse at apex，erect or curved，stipitate at base.
This section is somewhat morphologically similar to
Carex sect. Clandestinae G. Don，but differs in having
involucral bracts leaf-like，longer than spikes，shortly
sheathed; achenes emarginate at apex; styles thickened at
base but caduceus.
Two species are recognized，and the key to these two
species is as following.
3.1 Carex blinii H. Lév. & Vaniot，Bull. Soc. Bot. France
53: 316. 1906. TYPE: China，Kwei-tschou (Guizhou):
Pintong (Pingtang)，no date，J. Esquirol 343 (lectotype，
P!). Fig 5: A-F
C. elmeri Kük.，Repert. Spec. Nov. Regni Veg. 8: 326.
1910. TYPE: Philippines，Luzon Isl.，Baguio，23 Dec 1907，
A. D. E. Elmer 8444 (holotype，K!; isotype，E!).
C. granifera Dunn，J. Linn. Soc.，Bot. 38: 372. 1908.
TYPE: China，Fokien (Fujian)，Tap Long Ho Wood，no
date，HK Herb. 3655 (holotype，K!).
C. minuticulmis S. W. Su & S. M. Xu，Bull. Bot. Res.，
Harbin 8(3): 79，fig. 3. 1988. TYPE: China，Anhui，Taihu，
28 Apr 1979，S. W. Su 2477 (holotype，ANU!，transferred
to ACM!).
C. shanghaiensis S. X. Qian & Y. Q. Liu，Acta
Phytotax. Sin. 37(1): 103，fig. 1. 1999. ——C. blinii H.
Lév. & Vaniot subsp. Shanghaiensis (S. X. Qian & Y. Q.
Liu) S. Yun Liang & T. Koyama in C. Y. Wu & al.，Fl.
China 23: 387. 2010. TYPE: China，Shanghai，Shanghai
Botanical Garden，27 Apr 1995，Y. Q. Liu & S. X. Qian
63 (holotype，PE!).
C. sublateralis T. Koyama，Jap. J. Bot. 15: 180. 1956.
TYPE: China，Kiangsu (Jiangsu)，Suzhou，Mt. Shangfang，
no date，K. Kimure s. n. (holotype，KYO!).
C. taihokuensis Hayata，Icon. Pl. Formosan. 10: 70.
1921. TYPE: China，Formosa (Taiwan)，near Taihoku
(Taipei)，no date，U. Faurie s. n. (holotype，TI!; isotype，
TI!).
C. tatsutakensis Hayata，Icon. Pl. Formosan. 6: 133，fig.
45. 1916. TYPE: China，Formosa (Taiwan)，Musha，
Tatsutake，no date，W. R. Price 797 (holotype，TI!).
Rhizomes ligneous，thick，short. Culms lateral，tufted，
(2–)5–10(–25) cm tall，trigonous，smooth or scabrous on
upper part，with dark purple or purple fibrous sheaths at
base. Leaves much longer than culms; blades flat，2–8 mm
wide，soft or slightly stiff，smooth or scabrous on upper part.
Involucral bracts leaf-like，almost not-sheathed. Spikes
2–4，crowded，nearly capitate; terminal spike staminate，
linear-cylindrical，4–10 mm long，many-flowered; lateral
spikes pistillate，oblong，broadly ellipsoid，ovoid or nearly
globose，5–12×4–5 mm，loosely 2–7-flowered; peduncles
short，enclosed in sheath，or lowermost one slightly exserted.
Staminate scales elliptic-ovate to ovate，3.5–5 mm long，
obtuse or acuminate at apex，green 3-veined costa on dorsal
surface. Pistillate scales elliptic，lanceolate-ovate to broadly
ovate，2–4 mm long，pale green or greenish yellow，reddish
brown laterally，acute or acuminate at apex，green 3-veined
costa excurrent into a short mucro or awn. Perigynia ovoid
to nearly globose (excluding beaks)，obtusely trigonous，
(3–)5–7 mm long，longer than pistillate scales，greenish
yellow or yellowish brown，purple punctate on upper part，
distinctly or inconspicuously veined，nearly glabrous or
sparsely pubescent，gradually contracted into a beak at apex;
orifice 2-lobed with minute teeth. Stigmas 3; style thickened
at base. Achenes tightly enveloped，ovoid or broadly ovoid，
trigonous，yellowish brown or castaneous，2.5–3 mm long，
sides slightly concave，excavate or obtuse at apex，erect or
curved stipitate at base. Fl. and fr. March–May.
Distribution Eest，Central，South and Southwest
China，Philippines and Thailand.

1期 岑佳梦等：中国薹草属（莎草科）研究随记（IV）：隐匿薹草组的分类修订 41


Additional specimens examined
CHINA. Guangxi: Hechi，Dongnanwang，L. Q. Chen
92156 (IBK). Guizhou: Libo，Daqikong，X. F. Jin & al.
2388 (HTC)，2390 (HTC). Hainan: Wuzhishan，Mt. Nanle，
Ng Sai Chit 5125 (IBSC). Hubei: Hefeng，Pingxi，Sangzhi
For. Ins. 531 (KUN). Sichuan: Yanyuan，Bailing，
Qing-zang Exped. 12871 (KUN). Taiwan: Chiayi，Mt.
Arisan，anonymous 94 (TI)，U. Faurie 19 (P). Hsingchu，
Chienshih，P. F. Lu 9593 (HAST). Hualien，Hsiulin，Y. Y.
Huang & al. 622 (HAST). Nantou，Jenai，Wushe，W. C.
Leong 2001 (HAST)，2018 (HAST); Mt. Lianhua，T.
Koyama & al. 13787 (TI). Pingtung，Lilungshan，W. C.
Leong 2281 (HAST); Manchou，Laofoshan，C. H. Liu & al.
353 (HAST). Taitung，Peinan，W. C. Leong 2697 (HAST);
without precise locality，J. Ohwi 1388 (KYO). Taipei，
Tatungshan，W. C. Leong 773 (HAST); Wenshan，J. Ohwi
570 (KYO)，617 (KYO)，774 (KYO). Urai，U. Faurie 12
(KYO)，19 (KYO). Yunnan: Funing，Lungmai，C. W.
Wang 88878 (KUN). Zhejiang: Hangzhou，H. Wang & X.
F. Jin 1444 (HTC)，1446 (HTC). Lin’an，Changhua，X. F.
Jin & Y. Y. Zhou 3050 (HTC)，3051 (HTC). Yueqing，Mt.
Yandang，X. F. Jin & P. J. Cao 1529 (HTC)，1533 (HTC).
PHILIPPINES. Canlaon: Volcano，Negros，E. D. Merrill
6972 (BM，K，P). Luzon: Kalinga，Lubuagan，M. Ramos
& E. Edano 37573 (BM，K); Laguna，Mt. Banajao，E. D.
Merrill 8025 (BM，K). Mindoro: Mt. Calavite，M. Ramos
39401 (K， P). TAILAND. Chaiyaphum: Ban Nam
Phrom.，R. Geesink & al. 6920 (KYO); C. F. von Beusekom
& al. 4075 (KYO).
3.2 C. oxyphylla Franch.，Nouv. Arch. Mus. Hist. Nat.，III，
10: 57. 1898. TYPE: China，Yunnan: Langcang (Lancang)，
5 Jun 1886，Delavey 2524 (holotype，P!; isotype，E!). Fig
5: G，H
C. canaliculata P. C. Li，Acta Bot. Yunnan. 12: 148.
1990. TYPE: China，Yunnan: Weixi，Xinhua，19 May
1982，Qinghai-Xizang Exped. 6584 (holotype， PE!;
isotypes，KUN!，PE!).
C. changmuensis Tang & F. T. Wang ex Y. C. Yang，Fl.
Xizang. 5: 410，fig. 234. 1987. TYPE: China，Xizang，
Nielamu，Changmu，6 May 1966，Y. T. Chang & K. Y. Lang
3227 (holotype，PE!)，not 3221.
C. distantiflora Nakai，Repert. Spec. Nov. Regni Veg.
13: 245. 1914. TYPE: Korea，Cheju Isl. 20 May 1913，1332
(lectotype，TI!).
C. infossa C. P. Wang，J. Nanjing Univ. (Biol.) 1962 (2):
52. 1962. TYPE: China，Anhui，Chuzhou，Mt. Langya，2
Jun 1957，C. P. Wang 76 (holotype，N!; isotype，N!).
C. infossa C. P. Wang var. extensa S. W. Su，J. Anhui
Univ.，Nat. Sci. Ed. 20(3): 108. 1996. TYPE: China，Anhui，
Qimen，Chawan，7 May 1982，S. W. Su 82005 (holotype，
ANU; isotype，N!).
C. jackiana Boott f. oxyphylla (Franch.) Kük.，
Pflanzenr. IV，20(38): 638. 1909.
C. lateralis Kük.，Pflanzenr.，IV，20(38): 639. 1909.
TYPE: Ceylon，Obere Bergregion，no date，Thwaites 3198
(holotype，K!).
C. loheri C. B. Clarke，J. Linn. Soc.，Bot. 37: 14. 1904.
TYPE: Philippines，Benguet，no date，A. Loher 701
(holotype，K!).
C. longirostrata C. A. Mey. var. exaristata X. F. Jin &
C. Z. Zheng，Acta Phytotax. Sin. 42(6): 548. 2004. TYPE:
China，Zhejiang，Yueqing，Mt. Yandang，28 Apr 1980，
Q. C. Chen & J. H. Zhou 1879 (holotype，HZU!; isotype，
PE!).
C. longirostrata C. A. Mey. var. hoi Chü ex S. Yun
Liang，Acta Phytotax. Sin. 36(6): 537. 1998. TYPE: China，
Zhejiang，Zhenhai，Chengwan，no date，Y. Y. Ho 943
(holotype，PE!).
C. lyi H. Lév. & Vaniot，Repert. Spec. Nov. Regni Veg.
3: 349. 1907. TYPE: China，Kouy-Tcheou (Guizhou)，Pinfa
(Pingfa)，26 Apr 1909，J. Cavalerie 2329 (holotype，P!;
isotype，E!).
C. macrandrolepis H. Lév. & Vaniot，Repert. Spec.
Nov. Regni Veg.5: 241. 1908. TYPE: Korea，Isl. Quelpaert，
May 1907，U. Faurie 2285 (holotype，P; isotype，KYO!).
C. sharyotensis Hayata，Icon. Pl. Formosan. 10: 69，fig.
46. 1921. TYPE: TYPE: China，Taiwan，Sharyoto，19 Mar
1916，B. Hayata s. n. (holotype，TI!).
Rhizomes ligneous，thick，short，sometimes with
slender stolons. Culms central，rarely lateral，(10–) 20–40
cm tall，trigonous，smooth，with dark brown bladeless
sheaths at base. Leaves slightly longer than culms，or almost
equal in length; blades flat，2–6 mm wide，soft，smooth or
scabrous on upper part. Involucral bracts leaf- like or shortly
42 广 西 植 物 36卷

leaf-like，short-sheathed to conspicuously sheathed; sheaths
3–25 mm long. Spikes 3–5，remote，or distal two crowded
and others remote; terminal spike staminate ，
linear-cylindrical，5–30 mm long，many flowered; lateral
spikes pistillate，oblong or cylindrical，8–20×ca. 4 mm，
loosely 5–10-flowered; upper peduncles enclosed in sheath，
lower ones exserted. Staminate scales ovate to
lanceolate-ovate，5–5.5 mm long，acuminate at apex，green
3-veined costa on dorsal surface. Pistillate scales
lanceolate-ovate，oblong-ovate，ovate to obovate，3.5–4 mm
long，pale green or greenish yellow，reddish brown later-
ally，emarginate or acuminate at apex，green 3-veined costa
excurrent into a short mucro or awn. Perigynia ellip-
soid-obovoid，obovoid or ovoid (excluding beaks)，obtusely
trigonous，4–6 mm long，longer than pistillate scales，
greenish yellow or yellow，purple punctate on upper part，
distinctly or inconspicuously veined，nearly glabrous or
sparsely pubescent，gradually contracted into a beak at apex;
orifice 2-lobed with minute teeth. Stigma 3; style thickened
at base. Achenes tightly enveloped，ovoid，broadly ovoid or
obovoid，trigonous，yellowish brown or castaneous，2.5–3.5
mm long，sides slightly concave，excavate or obtuse at
apex，erect or curved stipitate at base. Fl. and fr. April-June.
Distribution East，Central，Southwest China，Japan，
South Korea，India and Philippines.
Additional specimens examined
CHINA. Anhui: Chuzhou，Mt. Langya，S. W. Su
81065 (ACM); anonymous s. n. (ACM). Guizhou: Pingfa，
J. Cavalerie 2325 (E). Jiangsu: Jurong，Mt. Baohua，C. P.
Wang 72 (NAS). Taiwan: Taipei，Mt. Nankou-taisan，J.
Ohwi 2505 (KYO，SYS); Shangto，J. Ohwi 154 (TI);
without precise locality，J. Ohwi 182 (KYO)，s. n. (KYO).
Yunnan: Fugong，Mt. Biluoxueshan，Qinghai-Xizang
Exped. 7029 (PE). Lushui，Luma，Beijing Hengduan Exped.
245 (PE); Mt. Gaoligongshan，Beijing Hengduan Exped. 47
(PE)，356 (PE). Weixi，Yezhi，Naluo，Qinghai-Xizang
Exped. 6352 (PE). Zhejiang: Hangzhou，Huanglongdong，
H. Wang & X. F. Jin 1445 (HTC). Pan’an，Huaxi，X. F. Jin
2696 (HTC). KOREA. Cheju Isl.，T. Nakai 941 (TI).
JAPAN. Pref. Chiba: Isumi-gun，Misaki-cho，T. Makino
224536 (KYO). Pref. Ehime: without precise locality，J.
Oda 1942 (KYO). Pref. Hyogo: Mihara-gun，Nandan-cho，
N. Fukuoka & N. Kurosaki 10317 (KYO，TI). Pref.
Kagoshima: Satuma，Yamakawa-cho，T. Hatsusima 29935
(KYO，TI). Pref. Kochi: Takaoda-prov.，Kubokawa Town，
T. Miyazaki 605160 (TI)，605180 (TI)，605187 (TI). Pref.
Kumamoto: Kamimatsukuma，K. Mayebara 34 (TI)，207
(KYO). Pref. Nagasaki: Tsushima-shi，Izuhara-cho，T.
Hoshino & al. 21281 (KYO). Pref. Nara: Nara-shi，Mt.
Kasuga，K. Seto 47219 (KYO). Kasugayama，Y. Tsujikuma
s. n. (TI). Pref. Saga: Karatsu-shi，Yobuko-cho，T. Hoshino
& al. 21757 (KYO，TI). Pref. Shizouka: Shizouka-shi，J.
Sugimoto s. n. (KYO，TI). INDIA. Coonoor Lambrock:
Nilgiris，S. S. Hooper & C. Saldanda 18149 (K)，C.
Townsend & T. P. Ramamoorthy 70 (K). PHILIPPINES.
Luzon: Benguet，Baguio，A. D. E. Elemer 8582 (E，K);
Quezon，J. S. Sands s. n. (K).
Acknowledgements We are most grateful to Profes-
sors LIANG Song-Yun and DAI Lun-Kai for their kind help
and continuous encouragement，to the curators of ACM，
BM，E，HTC，HZU，K，KUN，KYO，N，P，PE and
TI for permitting the senior author to visit their herbaria.
Heartfelt thanks are given to Dr. Andrew L. Hipp for his
constructive suggestions and improvement of the manu-
script.
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Appendix.
Voucher specimens which perigynia and achenes gathered.
[Taxa，vouchers (collector，collected number and herbar-
ium)，locality-perigynium-achene]
Carex blinii H. Lév. & Vaniot (sample 1): X. F. Jin & al.
2388，HTC; China，Guizhou，Libo-perigynium-achene. C.
blinii H. Lév. & Vaniot (sample 2): X. F. Jin & al. 2390，HTC;
China，Guizhou，Libo-perigynium. C. minuticulmis S. W. Su &
S. M. Xu: S. W. Su 2477， ACM; China， Anhui，
Taihu-perigynium-achene. C. taihokuensis Hayata: J. Oda
1747，KYO; Japan，Okinawa Pref.，Ishigaki-shi-perigynium-
achene. C. tatsutakensis Hayata: J. Ohwi 169，KYO; China，
Taiwan ， Taipei-perigynium-achene. Carex. tatsutakensis
Hayata (sample 2): anonymous 94，TI; China，Taiwan，Chiayi，
Mt. Arisan-achene. C. infossa Z. P. Wang:Z. P. Wang 72，NAS;
China， Jiangsu， Jurong，Mt. Baohua-perigynium-achene
(sample 1). C. infossa Z. P. Wang (sample 2):S. W. Su 81065，
ACM; China，Anhui，Chuzhou，Mt. Langya-achene. C. infossa
Z. P. Wang (sample 3):H. Wang & X. F. Jin 1445，HTC; China，
Zhejiang，Hangzhou-achene. C. infossa Z. P. Wang var. extensa
S. W. Su: anonymous s. n.， ACM; China， Anhui，
Qimen-perigynium. C. lateralis Kük.: E. D. Merrill 6972，P;
Philippines，Canlaon，Volcano. C. longirostrata C. A. Mey. var.
exaristata X. F. Jin & C. Z. Zheng: Q. C. Chen & J. H. Zhou
1879，HZU; China，Zhejiang，Yueqing，Mt. Yandang-
perigynium-achene. C. longirostrata C. A. Mey. var. hoi S. Yun
Liang: Y. Y. Ho 943， PE; China， Zhejiang， Zhenhai-
perigynium-achene. C. macrandrolepsis H. Lév. (sample 1): J.
Ohwi 2505，KYO; China，Taiwan，Taipei-perigynium-achene.
C. macrandrolepsis H. Lév. (sample 2): T. Hoshino & al.
21757，TI; Japan，Saga Pref.，Karatsu-shi-perigynium-achene.
Carex oxyphylla Franch.: Delavey 2524，P; China，Yunnan，
Lancang-perigynium. C. sharyotensis Hayata: B. Hayata s. n.，
TI; China，Taiwan，Sharyoto-perigynium.