$¢PhytoKeys

PhytoKeys 251: 257-270 (2025)
DOI: 10.3897/phytokeys.251.142179

Research Article

Carex yangchunensis, a new species of Cyperaceae from
the limestone regions of Guangdong, South China

Yi-Fei Lu'®, De-Chang Meng, Xiao-Feng Jin'®

1 School of Forestry and Biotechnology, Zhejiang A&F University, Hangzhou, 311300, Zhejiang, China
2 College of Horticulture and Landscape Architecture, Zhongkai University of Agriculture and Engineering, Guangzhou 510225, Guangdong, China
Corresponding author: Xiao-Feng Jin (docxfjin@163.com)

OPEN Qaceess

Academic editor: Pedro Jiménez-Mejias
Received: 19 November 2024
Accepted: 28 December 2024
Published: 29 January 2025

Citation: Lu Y-F, Meng D-C, Jin X-F
(2025) Carex yangchunensis, a new
species of Cyperaceae from the
limestone regions of Guangdong,
South China. PhytoKeys 251:
20/-270. https://doi.org/10.3897/
phytokeys.251.142179

Copyright: © Yi-Fei Lu et al.
This is an open access article distributed under
terms of the Creative Commons Attribution

License (Attribution 4.0 International - CC BY 4.0).

Abstract

Carex yangchunensis (Cyperaceae), a new species of Carex sect. Cryptostachyae in
limestone regions of Guangdong, China, is described and illustrated. Both morpholog-
ical observation and molecular analysis revealed that the new species was similar to
C. cryptostachys, but differs in having inflorescence with 4-8 spikes, ovoid or nearly
globose, 3-8 mm long, utricles (2.5-3.5 mm long) and nutlets (2—2.2 mm long) shorter,
style base thickened, leaves narrower, 3-6 mm wide and culms 8-25 cm tall. Scanning
electron micromorphology of utricles and nutlets of the new species and the related
species C. cryptostachys are provided.

Key words: Carex sect. Cryptostachyae, Carex yangchunensis, Cyperaceae, micromor-
phology, phylogeny, taxonomy

Introduction

The Cyperaceae, a family of grass-like plants containing 5600+ species in 95
genera, is the third largest family amongst monocots following the Orchidace-
ae and Poaceae (Larridon et al. 2021). Carex L., with over 2000 species, is the
largest genus within the Cyperaceae and is one of the most species-rich genera
amongst angiosperms. The genus Carex is distributed worldwide, except in Ant-
arctica, forming centres of diversity in temperate regions (Roalson et al. 2021).

Kuikenthal (1909) divided the genus Carex into four subgenera: subg. Primo-
carex, subg. Indocarex, subg. Vignea and subg. Eucarex. Kiikenthal’s classifi-
cation of Carex has been widely adopted (Ohwi 1936; Nelmes 1951; Akiyama
1955; Dai et al. 2010; Katsuyama 2015). Egorova (1999) re-circumscribed these
subgenera and proposed new nomenclatural names: subgenera Psyllophora,
Vigneastra, Vignea and Carex. Additionally, Egorova established a new subge-
nus, subg. Kreczetoviczia Egor., for the species previously placed in subg. Carex
but with two styles.

Phylogenetic analyses, based on DNA sequences, have been increasing-
ly applied in this century, as well as to the systematics of Carex, leading
to a profound re-evaluation of its classification and subgeneric divisions.

257

Yi-Fei Lu et al.: Anew sedge from the limestone regions of Guangdong

The phylogenetic studies revealed that all the other subgenera within Carex
are polyphyletic, except subg. Vignea, which indicated that the traditional
definition of the genus is polyphyletic and several smaller genera (Cymophyl-
lus Mack., Kobresia Willd., Schoenoxiphium Nees and Uncinia Pers.) within
trib. Carieae are nested within the branches of Carex (Starr et al. 1999; Yen
and Olmstead 2000; Roalson et al. 2001). The “Global Carex Group” pro-
posed a broader circumscription of Carex which completely involved trib.
Carieae (Global Carex Group 2015; Jiménez-Mejias et al. 2016; Martin-Bravo
et al. 2019; Roalson et al. 2021). Furthermore, Villaverde et al. (2020) de-
veloped a systematic framework for Carex using HybSegq and introduced a
classification system consisting of six subgenera: subg. Siderosticta M.J.
Waterway, subg. Carex, subg. Euthyceras Peterm., subg. Psyllophorae (De-
gland) Peterm., subg. Uncinia (Pers.) Peterm. and subg. Vignea (P. Beauv. ex
T. Lestib.) Heer.

Carex sect. Cryptostachyae comprises only one species, C. cryptostachys,
which was established by Franchet (1898) during his studies of Carex from
East Asia. Kukenthal (1909) placed it in sect. Mitratae, and Ohwi (1936) treat-
ed sect. Mitratae as a subsection, a view also accepted by Akiyama (1955).
Nelmes (1951) conducted a taxonomic study on Carex from the Malay Archi-
pelago and recognised sect. Cryptostachyae, based on the traits such as having
many androgynous spikes and considered the shapes of utricles and nutlets.
A recent phylogenetic study revealed sect. Mitratae represents an independent
group situated between the Indica and Decora clades.

During a botanical exploration in the limestone regions of Guangdong Prov-
ince in southern China, we collected a plant closely resembling Carex crypto-
stachys, which grows on limestone and has ovoid or nearly globose spikes.
Based on morphological comparison and phylogenetic analysis, we confirm it
as a new species and described it below.

Material and methods
Observation and comparison of morphological characters

Morphological characteristics of the new species were based both on stud-
ies of specimens and field trips. We focused on the shape of utricles and
nutlets, inflorescence and spikes, which demonstrated that these speci-
mens were similar to Carex cryptostachys. Herbarium specimens (including
all type specimens) of sections Cryptostachyae, Lageniformes and Mitratae
were examined at the following herbaria: BM, E, IBK, IBSC, K, KUN, KYO, P
PE, Tl and ZJFC.

SEM observation of utricles and nutlets

Scanning electron microscope (SEM) observations of utricles and nutlets of
the new species and the similar species Carex cryptostachys were conduct-
ed. Mature utricles and nutlets were gathered from specimens, including X. F.
Jin & Y.F. Lu 5196 for the new species and X. F. Jin & al. 2992 for C. crypto-
stachys. The utricles were submerged in 50% ethanol to clean them for 2 h.

PhytoKeys 251: 257-270 (2025), DOI: 10.3897/phytokeys.251.142179 958

Yi-Fei Lu et al.: Anew sedge from the limestone regions of Guangdong

The nutlets were initially soaked in a solution of concentrated sulphuric acid
and acetic anhydride (volume ratio = 1:9) for 12 hours, then rinsed in acetic
acid for 10 min and water for 5 min; next, they were placed in a bath-type
ultrasonic cleaner for 7 min with 70% ethanol to remove the cuticle and out-
er periclinal wall of the epidermis. After air drying, the cleaned utricles and
nutlets were mounted on stubs by doubled-sided adhesive tape and coated
with gold; next, they were observed and photographed under a Gemini-300
scanning electron microscope (Zeiss, Jena, Germany) (Jin and Zheng 2013;
Lu et al. 2021).

Taxon sampling for phylogenetic analyses

A total of 146 species (including infraspecies) representing 58 clades or sec-
tions proposed by Roalson et al. (2021) were used for phylogenetic analyses.
Of these, the sequences for two species (one sample of C. cryptostachys
and three samples of the new species) were newly generated, with the cor-
responding voucher specimens deposited in ZJUFC. The remaining sequences
were sourced from GenBank with voucher and GenBank accession information
provided in Appendix 1.

DNA extraction, PCR amplification and sequencing

Total genomic DNA was extracted from silica-dried leaves using Plant Genomic
DNA Kit (Tiangen Biotech Co., Ltd., Beijing, China). Two nuclear DNA markers
(ETS and ITS) and three plastid DNA markers (matK, rpl32-trnL® and trnL-F)
were selected to amplification. Sequence amplification was conducted follow-
ing Lu et al. (2021) and sequencing was performed on an ABI 3730 automated
sequencer (Applied Biosystems, Foster City, CA, USA).

Phylogenetic analyses

Sequences were assembled using SeqMan in DNASTAR Lasergene
v.8.1.3 (Burland 2000), followed by alignment with MAFFT software, on-
line version (https://mafft.cbrce.jp/alignment/server/) using the strate-
gy of L-INS-i. Then the aligned sequences were trimmed and manually
edited in MEGA v.7 (Kumar et al. 2016). The best nucleotide substitu-
tion model was determined for each gene region in jModelTest v.2.1.10
(Guindon and Gascuel 2003; Darriba et al. 2012;) using Akaike Informa-
tion Criterion. Finally, five DNA regions were concatenated in PhyloSuite
v.1.2.3 (Zhang et al. 2020). Two strategies were used for phylogenetic
analyses. A Maximum Likelihood (ML) tree was constructed using RAx-
ML-HPC BlackBox v.8.2.12 in the CIPRES Science Gateway (https://www.
phylo.org/); the number of bootstraps was set as 1000 with the GTR+I+G
model. Bayesian Inference (BI) was conducted in MrBayes v.3.2.7a; two
runs and four chains were carried out for 107 simultaneous generations
with sampling of one tree every 1000 generations. A 50% majority-rule
consensus tree was obtained after discarding the first 25% of all trees as
burn-in. The phylogenetic trees were visualised in tvBOT (Xie et al. 2023).

PhytoKeys 251: 257-270 (2025), DOI: 10.3897/phytokeys.251.142179 959

Yi-Fei Lu et al.: Anew sedge from the limestone regions of Guangdong

Results
Comparison of morphological characters

A total of 149 sheets/specimens of Carex cryptostachys from south China, South-
east Asia, South Asia to Australia were deposited in the above-mentioned herbaria
and were available for measurement and a comparison of morphological charac-
ters. The morphological characters of C. cryptostachys showed considerable sta-
bility, but quite different from the new species in the characters listed in Table 1.

Micromorphology of utricles and nutlets

Under a SEM, the utricles of the new species are broadly rhombic-obovoid, obtusely
compressed-trigonous, dorsally glabrous and densely pubescent at the margin, with
many longitudinally veins and beak orifice 2-lobed with minute teeth (Fig. 1A-C).
The utricles of Carex cryptostachys are narrowly rhombic-obovoid, obtusely com-
pressed-trigonous, dorsally glabrous and densely pubescent at the margin, with
many longitudinal veins and beak orifice 2-lobed with sharp teeth (Fig. 1F—H).

The nutlets of the new species and C. cryptostachys are both rhom-
bic-obovoid, dorsally flat and ventrally concaved above and below. The epider-
mal cells of the nutlets of the new species are slightly concaved, while those of
C. cryptostachys are convex with one silica body (Fig. 1D, E, |, J).

Phylogenetic relationships

The lengths of five aligned sequences are: ETS 714 bp, ITS 749 bp, matk
805 bp, rpl32trnL*®) 1035 bp and trnL-F 1134 bp. The GTR+I+G model was
the best for ETS and ITS and the GTR model for matK, rpI32-trnL‘© and trnL-F.
The topology between the Bayesian Inference and Maximum Likelihood trees
did not involve incongruences at the node with Bayesian posterior probability
(PP) > 0.75 or bootstrap values (BS) > 50%. Similar to previous studies, the deep
relationships amongst clades remain unresolved, with PP < 0.75 and BS < 50%.
The three samples of the new species, C. yangchunensis, formed a strongly
supported clade (PP = 1, BS = 100%) and showed a sister relationship with the
morphologically closest species, C. cryptostachys (Fig. 2).

Table 1. Comparison of morphological characters of Carex cryptostachys and the new

species.
Character Carex cryptostachys C. yangchunensis sp. nov.
Culm height 10-35 cm 8-25 cm
Leaf width 6-16 mm 3-6mm
Spike
Number 6-10 4-8
Shape Cylindrical Ovoid or globose
Length 9-25 mm 3-8mm
Utricle
Length 3.5-4.5 mm 2.5-3.5 mm
Beak length ca. 0.5mm ca. 0.3 mm
Nutlet
Length 2.5-3 mm 2-2.2 mm
Style base Not thickened Thickened

PhytoKeys 251: 257-270 (2025), DOI: 10.3897/phytokeys.251.142179 260

Yi-Fei Lu et al.: Anew sedge from the limestone regions of Guangdong

Figure 1. Micromorphology of utricles and nutlets of Carex yangchunensis sp. nov, (A-E) and Carex cryptostachys (F-J).
A-C, F-H utricles D, E, I, J nutlets A, D, F, | overview B, G utricle and nutlet surface, respectively C, H beak E, J utricle and
nutlet surface sculpture, respectively. Scale bars: 200 um (A, D, F, 1); 50 um (B, J); 100 um (C, H); 10 um (E)

PhytoKeys 251: 257-270 (2025), DOI: 10.3897/phytokeys.251.142179

Yi-Fei Lu et al.: Anew sedge from the limestone regions of Guangdong

60/0.82¢ C, nodaeana
00 C. =e eirgldiana
shangchen enisis
: & ta ai eee
51/079 1554 = anchengensis
00 : an Ot gshanica
0 & aiMaensis
aofangil
ciformis
00 ey
raminata 2
mira
00 £5 oxyandra
51/0.99 1 $e Btuensis
| -/0.76
o P90 GC jynhuangsla hanensis
chlorocepha : ;
C. daxinensis
00 Cc. umbrosa subsp. sabynensis
90% 0,99 C. alterniflor;
OO breviculmis
8 G breviaristata hi
inensis
83/0.97 C. conica ee
C. pisiformis
. C. polyse! pon, a
80) parse subsp. huetiana
LO0 C. chung
mitrata var. aristata
Cc. dapanshanigg
anhuiensis
R 0,82 , pingleensis
C, . breviscape
C. Aen icol
OO/1 |b tristachya,
9 or C pseu Stristachya
runcatis cet
86 a 63/0.8 C, reinii
9 20 blep yharicarpa
80 . tear rath dica e
oe C. tristachya var. pocilliformis
00 — C. tristis
C. stenocarpa :
94 C. liparocarpos
¢. tangulashanensis
L100 a C. oligocarya
C. alajica
00 C. cruenta
at oftusea ’
0 , angolensis
9 00 ae ; C. chlorosaccus
4 2 C. doisutepensis
C. pulchra
: (. kashmirens nsis
C. asturica
99 on tighardsonii
C. concinnoides 100/0.8
00/1 C, yan inher
1 cy unensis 3
00 Cer He ie c : sis
iL “a ae c
due 83/ ota ‘ptostac 3
00 uPut caucasica ia.
C. aboriginum
0 C, harrysmithii C. chuik
. chuii
00 C. cardiol eis
qpsyangensis
ay ratnerdii C etioas
. ericetorum
00/1 cc digitata
eo
9 = CS Tokjetensts
C. cherokeensis tg oO oan
9 aucodea
P 94/0,98 — (on abscondita
0 = C. collinsii
: emmonti
issuricolé
00/1) i. baghinishantne
0 . egena
C. fastiziat C. tomentosa
. fastigiata -
e 86/, C. borii
Qaematostoma ,
J . Inanis .
Cuivalovins C. lancisquamata
. ivanovie
7 C. herbacoeli
; ort; ae C. agastachys
86 C. pendu a
00/1 rainbowii 4
s ; C. demi C. cretica
gc _ demissa
8 C. flava .
C. strigosa ls,
00 = — C. perraudieriana
Cc. aeth Qpica
— C. lonchocarpa
Pups C. folliculata
C. triquetra
Cc. frigid 4
00 elicata,
C. capil aris subsp. fuscidula ane
C. esquiroliana
00 C. phacota . ae
6 — C. maximowiczii
C. antoniensis
C. bullata
- . macrochieta
C. stylo: ¢
= C. aperta
C, prasina i ike
: C. magellanica subsp. irrigua
C. limosa ‘ R
Cs C. shortiana
= squarrosa
8 ae bigelowli
C augusta ata
ca faeta
00 C. breedlovel 4
C. chordalis
00 Cc. beiie sis
= glaucescens
00 — C. curatorum
C. scirpoidea
as 9 ci ep onica
35/- Cc. mo tidata
9 00 C. ischnostachya
' C. transversa
99 C. vestita
= ; C. globularis
C. sartwelliana _
100/1 C. pe euler
. picta
LOO P C. halleriana
. C. lativena
00 C. complanata

C. aestivalis
00/1)

C. alliiformis

co rubertiaia

ligula

C. makinoensis

Figure 2. Maximum Likelihood phylogenetic tree, based on two nrDNA (ETS, ITS) and three cpDNA (matK, rpl32-trn-
L(UAG), trnL-F) samples. The numbers above the branches are bootstrap values (BS) and Bayesian Posterior Probabili-

ties (PP). ‘’ indicates BS < 50% or PP < 0.75.

PhytoKeys 251: 257-270 (2025), DOI: 10.3897/phytokeys.251.142179

262

Yi-Fei Lu et al.: Anew sedge from the limestone regions of Guangdong

Taxonomic treatment

Carex yangchunensis X.F.Jin, Y.F.Lu & D.C.Meng, sp. nov.
urn:lsid:ipni.org:names:7735591 6-1

Figs 3A-I, 4A-J

Chinese name: yang chin tai cao (SH 7 2t 5)

Diagnosis. This new species is similar to Carex cryptostachys Brongn., but dif-
fers in having spikes 4-8, ovoid or nearly globose, 3-8 mm long (vs. spikes
6-10, cylindrical, 9-25 mm long), utricles 2.5-3.5 mm long (vs. 3.5-4.5 mm
long), nutlets 2-2.2 mm long, with styles thickened at base (vs. 2.5-3 mm long,
style base not thickened), leaves 3-6 mm wide (vs. 6-16 mm wide) and culms
8-25 cm tall (vs. 10-35 cm tall).

Type. CHINA, Guangdong (J 45), Yangchun County (SH#), Kongdongyan
Scenic Spot (illl45 X), from Banshanting to Zhaixingting (“Eli 23 =),
22.18544°N, 111.74557°E, elev. 80 m, 9 April 2024, X. F Jin & Y¥. F. Lu 5196
(holotype: ZJFC!; isotypes: IBK!, PE!, ZJFC!, ZM!).

Description. Perennial herbs. Rhizomes elongate, woody, thick. Culms lateral,
8-25 cm tall, compressed-trigonous, slender, smooth, base with dark-brown or
brown sheaths, sometimes splitting into fibres. Leaves far longer than culms,
apex slender long-caudate; blades 3-6 mm wide, thinly leathery, flat, upper mar-
gins and both surfaces scabrous. Bracts short-setaceous, sheathed; sheaths
3-10 mm long, with the most proximal one longer than the others. Spikes 4-8,
androgynous, ovoid or nearly globose, 3-8 mm long, base with a 3-17 mm long
slender peduncle, staminate part ca. 2 mm long, slightly exserted or hidden in
pistillate flowers, pistillate part 3-6 mm long, densely 4-12-flowered; pedun-
cles exserted from bract sheaths. Staminate glumes ovate, 1.5-2 mm long, yel-
low-white, obtuse at apex, green 3-veined dorsal costa. Pistillate glumes obo-
vate or ovate, 2-2.5 mm long, pale yellow-green, acute or obtuse at apex, green
3-veined dorsal costa. Utricles yellow-green, broadly rhombic-obovoid (excluding
beak), obtusely compressed-trigonous, 2.5-3.5 mm long (including beak), longer
than pistillate glumes, yellow-green, membranous, distinctly thinly veined, sparse-
ly pubescent on upper dorsal surface and margins, densely pubescent on ventral
surface, base cuneate and narrowed into a ca. 0.5 mm long stipe, apex gradually
contracted into a ca. 0.3 mm long beak, orifice 2-lobed with minute teeth. Nutlets
tightly enveloped, rhombic-ovoid, trigonous, yellow, 2—2.2 mm long, with 3 angles
constricted at middle, sides concave above and below, base with a 0.3-0.7 mm
long stipe; style thickened at base, persistent, coiled; stigmas 3.

Etymology. The specific epithet ‘yangchunensis’ refers to the type locality of
this new species in Yangchun County.

Phenology. Flowering and fruiting all the year round.

Distribution and habitat. The new species has been collected from the lime-
stone regions of Kongdongyan Scenic Spot near Yangchun County, in Heshui
and Chunwan townships. It is believed to be distributed in similar limestone
hills near Yangchun County. It is currently Known to grow on limestone land-
scapes at lower elevations of 60-80 m.

Additional specimens examined (paratypes). CHINA. Guangdong (J 4):
Yangchun County (4%), Chunwan Township (4/448), Gaocun Tourist Resort
(in PR FE (LX), 14 October 2023, D. C. Meng s.n. (IBK, ZJFC, ZM); the same

PhytoKeys 251: 257-270 (2025), DOI: 10.3897/phytokeys.251.142179 763

Yi-Fei Lu et al.: Anew sedge from the limestone regions of Guangdong

Figure 3. Photos of Carex yangchunensis sp. nov. (A-l) and Carex cryptostachys (J, K) A, B habitat C, J habit D lower part
of habit, showing rhizome E middle part of habit, showing lateral culm F spike G utricles (above dorsal surface, below
ventral surface) H nutlet (left dorsal surface, right ventral surface) I, K inflorescence.

locality, 2 January 2024, D. C. Meng 2024010201 (ZJFC, ZM); Heshui Township
(47k #8), Matangjiao Village (AK AA), 22.38104°N, 111.92091°E, elev. 76 m,
8 April 2024, X. F. Jin & Y. F. Lu 5192 (ZJFC, ZM); Kongdongyan Scenic Spot (i
iii 25 5¢ IX), Banshanting (241), 22.18571°N, 111.74601°E, elev. 67 m, 9 April
2024, X. F. Jin & Y. F Lu 5198 (PE, ZJFC, ZM).

Conservation status. Least Concern (LC). The new species is a common
sedge and grows on cliffs of the limestone regions around Yangchun Coun-
ty, Guangdong Province. Two known populations are in good status, but are
strongly influenced by tourist activity, so that the species will need attention at
related locations for conservation (IUCN 2022).

PhytoKeys 251: 257-270 (2025), DOI: 10.3897/phytokeys.251.142179 264

Yi-Fei Lu et al.: Anew sedge from the limestone regions of Guangdong

Se

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ie \\ *

Sl)

i,

Figure 4. Carex yangchunensis, sp. nov. A habit B lateral culms C spike D staminate dolunte E, F pistillate glume G utricles
(dorsal surface) H utricles (ventral surface) I nutlet (dorsal surface) J nutlet (ventral surface) (drawn by Xiao-Feng Jin
from the holotype).

Acknowledgements

We are grateful to the curators and staff of the following herbaria, K, KYO, LE,
P PE, Tl, WUK and ZJFC, for their kind help during the authors’ visit and to Mr.
Philip E. Hyatt for improving the English.

PhytoKeys 251: 257-270 (2025), DOI: 10.3897/phytokeys.251.142179 265

Yi-Fei Lu et al.: Anew sedge from the limestone regions of Guangdong

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work was supported by the National Natural Science Foundation of China (Grant
No. 32070218) and the start-up scientific project of Zhejiang A&F University (Grant No.
2022LFRO11).

Author contributions

Yi-Fei Lu and Xiao-Feng Jin conceived the study. Yi-Fei Lu, De-Chang Meng and Xiao-Feng
Jin collected the samples. Yi-Fei Lu and De-Chang Meng conducted the measurements
of morphological characters, data analysis and SEM observation. Yi-Fei Lu and Xiao-
Feng Jin wrote the manuscript. All authors approved the final version of the manuscript.

Author ORCIDs

Yi-Fei Lu © https://orcid.org/0000-0001-6095-3821
De-Chang Meng ® hitps://orcid.org/0000-0002-1049-1232
Xiao-Feng Jin © https://orcid.org/0000-0002-8195-6738

Data availability

All of the data that support the findings of this study are available in the main text.

References

Akiyama S (1955) Carices of the Far Eastern Region of Asia. Hokkaido, Sapporo, 1-257.

Burland TG (2000) DNASTAR’s Lasergene sequence analysis software. Methods in Mo-
lecular Biology (Clifton, N.J.) 132: 71-91. https://doi.org/10.1385/1-59259-192-2:71

Dai LK, Liang SY, Zhang SR, Tang YC, Koyama T, Tucker GC (2010) Carex L. In: Wu ZY,
Raven PH, Hong DY (Eds) Flora of China (Vol. 23). Science Press, Beijing and Missouri
Botanical Garden Press, St. Louis, 285-461.

Darriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest 2: More models, new
heuristics and parallel computing. Nature Methods 9(8): 772-772. https://doi.
org/10.1038/nmeth.2109

Egorova T (1999) The Sedges (Carex L.) of Russia and adjacent States (within the lim-
its of the former URSS). St.-Petersburg State Chemical-Pharmaceutical Academy,
St.-Petersburg and Missouri Botanical Garden Press, St. Louis.

Franchet A (1898) Les Carex de I'Asie orientale III. Nouvelles Archivesdu Muséum d'His-
toire Naturelle. Paris III 10: 39-120.

Global Carex Group (2015) Making Carex monophyletic (Cyperaceae, tribe Cariceae): A
new broader circumscription. Botanical Journal of the Linnean Society 179(1): 1-42.
https://doi.org/10.1111/boj.12298

Guindon S, Gascuel O (2003) A simple, fast, and accurate algorithm to estimate large
phylogenies by maximum likelihood. Systematic Biology 52(5): 696-704. https://doi.
org/10.1080/10635150390235520

PhytoKeys 251: 257-270 (2025), DOI: 10.3897/phytokeys.251.142179 266

Yi-Fei Lu et al.: Anew sedge from the limestone regions of Guangdong

IUCN (2022) Guidelines for using the IUCN Red List Categories and Criteria. Version
15.1. Prepared by the Standards and Petitions Committee of the IUCN Species Surviv-
al Commission. https://www.iucnredlist.org/resources/redlistguidelines [accessed
on 1 February 2024]

Jimeénez-Mejias P Hahn M, Lueders K, Starr JR, Brown BH, Chouinard BN, Chung KS,
Escudero M, Ford BA, Ford KA, Gebauer S, Gehrke B, Hoffmann MH, Jin XF, Jung
J, Kim S, Luceno M, Maguilla E, Martin-Bravo S, Miguez M, Molina A, Naczi RFC,
Pender JE, Reznicek AA, Villaverde T, Waterway MJ, Wilson KL, Yang JC, Zhang
SR, Hipp AL, Roalson EH (2016) Megaphylogenetic specimen-level approach-
es to the Carex (Cyperaceae) phylogeny using ITS, ETS, and matK sequences:
Implications for classification. Systematic Botany 41(3): 500-518, 519. https://doi.
org/10.1600/036364416X692497

Jin XF, Zheng CZ (2013) Taxonomy of Carex sect. Rnomboidales (Cyperaceae). Science
Press, Beijing.

Katsuyama T (2015) Carex of Japan, enlarged and revised edition. Bun-ichi Co., Ltd.,
Tokyo.

Kiikenthal G (1909) Cyperaceae-Caricoideae. In: Engler H (Ed.) Das Pflanzenreich (Vol.
38). W. Engelmann, Leipzig, 1-824.

Kumar S, Stecher G, Tamura K (2016) MEGA7: Molecular evolutionary genetics analysis
version 7.0 for bigger datasets. Molecular Biology and Evolution 33(7): 1870-1874.
https://doi.org/10.1093/molbev/msw054

Larridon |, Zuntini AR, Léveillé-Bourret E, Barrett RL, Starr JR, Muasya AM, Villaverde
T, Bauters K, Brewer GE, Bruhl JJ, Costa SM, Elliott TL, Epitawalage N, Escudero M,
Fairlie |, Goetghebeur P Hipp AL, Jiménez-Mejias P Sabino Kikuchi IAB, Luceno M,
Marquez-Corro JI, Martin-Bravo S, Maurin O, Pokorny L, Roalson EH, Semmouri |,
Simpson DA, Spalink D, Thomas WW, Wilson KL, Xanthos M, Forest F, Baker WJ (2021)
A new classification of Cyperaceae (Poales) supported by phylogenomic data. Jour-
nal of Systematics and Evolution 59(4): 852-895. https://doi.org/10.1111/jse.12757

Lu YF, Jin XF, Ikeda H, Yano O, Benitez-Benitez C, Chen WJ, Liu YD, Jiménez-Mejias P
Yu MJ (2021) Revisiting of Carex sect. Confertiflorae s.l. (Cyperaceae): New data
from molecular and morphological evidence and first insights on Carex biogeogra-
phy in East Asia. Journal of Systematics and Evolution 59(4): 668-686. https://doi.
org/10.1111/jse.12795

Martin-Bravo S, Jiménez-Mejias P. Villaverde T, Escudero M, Hahn M, et al. (2019) A
tale of worldwide success: Behind the scenes of Carex (Cyperaceae) biogeography
and diversification. Journal of Systematics and Evolution 57(6): 695-718. https://
doi.org/10.1111/jse.12549

Nelmes E (1951) The genus Carex in Malaysia. Reinwardtia 1: 221-450.

Ohwi J (1936) Cyperaceae Japonicae I. A synopsis of the Caricoideae of Japan, includ-
ing the Kuriles, Saghalin, Korea, and Formosa. Memoirs of the College of Science.
Kyoto Imperial University 11: 229-530.

Roalson EH, Columbus JT, Friar EA (2001) Phylogenetic relationships in Cariceae
(Cyperaceae) based on ITS (nrDNA) and trnT-L-F (cpDNA) region sequences: Assess-
ment of subgeneric and sectional relationships in Carex with emphasis on section
Acrocystis. Systematic Botany 26(2): 318-341.

Roalson EH, Jiménez-Mejias P Hipp AL, Benitez-Benitez C, Bruederle LP Chung KS,
Escudero M, Ford BA, Ford K, Gebauer S, Gehrke B, Hahn M, Hayat MQ, Hoffmann
MH, Jin XF, Kim S, Larridon I, Léveillé-Bourret E, Lu YF, Lucefio M, Maguilla E, Marquez-
Corro JI, Martin-Bravo S, Masaki T, Miguez M, Naczi RFC, Reznicek AA, Spalink D,

PhytoKeys 251: 257-270 (2025), DOI: 10.3897/phytokeys.251.142179 267

Yi-Fei Lu et al.: Anew sedge from the limestone regions of Guangdong

Starr JR, Uzma, Villaverde T, Waterway MJ, Wilson KL, Zhang S-R (2021) A framework
infrageneric classification of Carex (Cyperaceae) and its organizing principles. Journal
of Systematics and Evolution 59(4): 726-762. https://doi.org/10.1111/jse.12722

Starr JR, Bayer RJ, Ford BA (1999) The phylogenetic position of Carex section Phyl-
lostachys and its implications for phylogeny and subgeneric circumscription in
Carex (Cyperaceae). American Journal of Botany 86(4): 563-577. https://doi.
org/10.2307/2656818

Villaverde T, Jiménez-Mejias P Luceno M, Waterway MJ, Kim S, Lee B, Rincon-Barrado
M, Hahn M, Maguilla E, Roalson EH, Hipp AL, Wilson KL, Larridon |, Gebauer S, Hoff-
mann MH, Simpson DA, Naczi RFC, Reznicek AA, Ford BA, Starr JR, Park J, Escudero
M, Martin-Bravo S, The Global Carex Group (2020) A new classification of Carex (Cy-
peraceae) subgenera supported by a HybSeq backbone phylogenetic tree. Botanical
Journal of the Linnean Society 194(2): 141-163. https://doi.org/10.1093/botlinne-
an/boaa042

Xie JM, Chen YR, Cai GJ, Cai RL, Hu Z, Wang H (2023) Tree visualization by one table
(tvBOT): A web application for visualizing, modifying and annotating phylogenetic
trees. Nucleic Acids Research 51(W1): W587-W592. https://doi.org/10.1093/nar/
gkad359

Yen AC, Olmstead RG (2000) Molecular systematics of Cyperaceae tribe Cariceae based
on two chloroplast DNA regions: ndhF and trnL intron-intergenic spacer. Systematic
Botany 25(3): 479-494. https://doi.org/10.2307/2666691

Zhang D, Gao FL, Jakovli¢ |, Zou H, Zhang J, Li WX, Wang GT (2020) PhyloSuite: An
integrated and scalable desktop platform for streamlined molecular sequence data
management and evolutionary phylogenetics studies. Molecular Ecology Resources
20(1): 348-355. https://doi.org/10.1111/1755-0998.13096

Appendix 1

GenBank accession numbers of the species are listed below (species name:
ETS, ITS, matK, rpI32trnL*®, trnL-F) and voucher information is provided for
the newly-generated sequences:

C. aboriginum: MN760588, MN761648, GU172441, —, —; C. abscondita:
MN759823, MN761824, GU172447, —, —; C. acaulis: MN760451, MN762256,
MN763688, —, —; C. aestivalis: MN760562, MN761566, GU172459, —, —;
C. aethiopica: MN759949, MN762349, MN763446, —, —; C. agastachys:
MN761179, MN762597, MN763240, —, —; C. alajica: MN761331, MN762415,
MN763374, —, —; C. alliiformis: MT920727, MT920759, MT920790,, MT920867;
C. alterniflora: OR913315, OR668798, OR866295, ORI13383, ORI13257; C. an-
golensis: MN759931, EU288569, MN762971, —, —; C. angustata: MN760929,
MN761518, GU172491, -—, —; C. anhuiensis: OR913316, OR668799, -,
OR913384, OR913258; C. antoniensis: MN760999, MN762074, MN763743, —,
—; C. aperta: MN760847, MN761632, GU172500, —, —; C. asturica: MN761067,
MN762464, MN763498, —, —; C. atrofusca: MN760042, MN761794, GU172555,
—, —; C. baimaensis: MN760558, MN761417, MN763557, —, —; C. baohu-
ashanica: OR913358, OR689467, OR866324, OR913429, OR913292; C. bige-
lowii: MN761315, MN762339, MN763487, —, —; C. blepharicarpa: MN759781,
—, -, -—, —} C. borii; MN761039, MN762187, MN763700, —, —; C. brainerdii:
MN760630, MN761882, GU172612, —, —; C. breedlovei: MN760977, MN762091,

PhytoKeys 251: 257-270 (2025), DOI: 10.3897/phytokeys.251.142179 268

Yi-Fei Lu et al.: Anew sedge from the limestone regions of Guangdong

MN763222, —, —; C. breviaristata:. OR913318, OR668800, OR866296,
OR913385, OR9I13259; C. breviculmis: OR913319, OR668801, OR866297,
OR913386, OR9I13260; C. breviscapa: MN760015, MN762575, MN763013, —,
—; C. bullata: MN761003, MN761622, GU172630, —, —; C. capillaris subsp. fus-
cidula: MN760542, MN762040, MN763516, —, —; C. cardiolepis: MN760171,
MN762377, MN763696, —, —; C. caucasica: MN760600, MN76225/7,
MN763745, —, —; C. chaofangii: MN759907, MN761627, —, —, —; C. chero-
keensis: MN760098, MN761606, GU172677, —, —; C. chinensis: OR913320,
OR668802, OR866298, —, ORI13261; C. chlorocephala: OR913321, OR668803,
OR866299, —, OR913262; C. chlorosaccus: MN759932, EU288577, MN763179,
—, -; C. chordalis: MN761070, MN762558, MN763114, —, —; C. chuiana:
OR913362, OR668841, OR866328, OR913412, ORI13296; C. chuii: MW458996,
MW459027, MW459094, OR913415, MW459058; C. chungii: OR913322,
OR668804, OR866300, OR9I13387, ORI13263; C. collinsii: MN760100,
MN761432, GU172686, —, —; C. complanata: MN760661, MN761557,
GU172701, —, —; C. concinnoides: MN760092, MN761930, GU172704, —, —;
C. conica: OR9I13323, OR668805, OR866301, ORI13388, ORI13264; C. cretica:
MN760085, MN762258, MN763412, —, —; C. cruenta: MN760446, MN762441,
MN763817, —, —; C. cryptostachys: MN760888, MN762869, MN763020, -,
—; C. cryptostachys 1: OR913324, OR668806, —, —, —; C. cryptostachys 3:
PQ539318, PQ535885, PQ539321, PQ539325, PQ539329, Hainan, Mt. Wuzhis-
han, X.F. Jin 2992; C. dapanshanica: OR913325, OR668807, —, —, ORI13265;
C. daxinensis: OR913363, OR668842, OR866329, OR913413, OR913297;
C. deflexa: MN760618, MN762102, GU172760, —, —; C. delicata: MN760541,
MN762542, MN763772, —, —; C. demissa: MN760083, MN761875, GU173786,
—, —; C. digitata: MN759970, MN762307, MN763754, —, —; C. doisutepensis:
MN759938, MN762045, MN763478, —, —; C. durieui: MN760122, MN762351,
MN763474, —, —; C. egena: OR913359, OR668838, OR866325,, ORI13293;
C. ericetorum: MN760026, MN762286, MN763731, —, —; C. esquiroliana:
MN761064, MN762053, MN763585, —, —; C. fastigiata: MN759999, MN762056,
MN763322, —, —; C. fissuricola. MN761373, MN762017, GU172869, —, —;
C. flava: MN760063, JN634689, MN763787, —, —; C. fokienensis: MT920734,
MT920766, MW459079, MT920838, MT920874; C. folliculata: MN760924,
MN761465, GU172882, —, —; C. foraminata 1: —, OR668835, OR866322, —, —;
C. foraminata 2: MN760492, MN761 796, MN763018, —, —; C. frigida: MN759952,
MN762466, MN763740, —, —; C. funhuangshanensis: 0R913364, OR668843,
OR866330, OR913414, OR913298; C. gifuensis: MN760643, MN761423, -,
—, -; C. giraldiana: MN760554, MN761919, MN763526, —, —; C. glaucodea:
MN759818, MN761439, FJ597209, —, —; C. globularis: MN760973, MN762535,
MN763409, —, —; C. haematostoma var. alexeenkoana: MN761040, MN762444,
MN763766, —, —; C. halleriana: MN760532, MN762338, MN763694, -,
—; C. harlandit. MN760527, MN762606, MN763046, —, —; C. harrysmithii:
MT920738, MT920770, MT920803, MT920842, MT920878; C. herbacoeli:
MN761231, MN762157, MN763527, —, —; C. inanis: MN760504, MN762430,
MN763678, —, —; C. ischnostachya: MT920741, MT920773, MT920806,
MT920845, MT920881; C. ivanoviae: MN761038, MN762446, MN763803, —, —;
C. japonica: MT920743, MT920774, MT920808, —, MT920883; C. kashmiren-
sis: MN759944, MN762354, MN763301, —, —; C. lancisquamata: MN760906,
MN762086, MN763314, —, —; C. lativena: MN760528, MN761490, GU173073,

PhytoKeys 251: 257-270 (2025), DOI: 10.3897/phytokeys.251.142179 269

Yi-Fei Lu et al.: Anew sedge from the limestone regions of Guangdong

—, —} C. lemmonii: MN760021, MN761555, GU173090, —, —; C. lianchengensis:
MN760555, MN761654, —, —, —; C. ligulata: MK481321, MK481341, MT920810,
MT920846, MK481361; C. limosa: MN761060, MN761787, GU173120, —, —;
C. liparocarpos: MN760648, MN762401, MN763805, —, —; C. lonchocarpa:
MN760923, MN761429, GU173128, —, —; C. longicolla: OR913333, OR668813,
—, 0R913391, OR913269; C. longirostrata: MN761208, MN762075, MN763305,
—, -; C. macrochaeta: MN760976, MN762098, GU173175, —, —; C. magellan-
ica subsp. irrigua: MN760861, MN762190, MN763818, —, —; C. makinoensis:
MN761145,MN762552, —, —, —;C. manciformis: MN760556, —,MN763629, —, —;
C. maubertiana: MW458988, MW459019, MW459086,, MW459050; C. maximo-
wiczil: ORI13360, OR668839, OR866326, ORI13409, ORI13294; C. microchae-
ta: MN761263, MN761572, GU173216, —, —; C. mitrata var. aristata: OR913334,
OR668814, OR866304, OR913392, OR913270; C. mollicula: MT920746,
MT920777, MT920813, MT920848, MT920887; C. nodaeana: MN760553,
MN762293, MN763800, —, —; C. oligocarya: MN760659, MN762404, MN763608,
—, —;C. ornithopoda: MN759995, MN762308, MN763753, —, —; C. pedunculata:
MN760639, MN761431, GU173371, —, —; C. pendula: MN761195, MN762602,
MN763148, —, —; C. perraudieriana: MN760016, MN762134, MN763400, —, —;
C. phacota: OR913361, OR668840, OR866327, OR913410, OR913295; C. picta:
MN760546, MN761561, GU1 73398, —, —; C. pingleensis: OR913338, OR668817,
OR866307, OR913395, OR913273; C. pisiformis: OR913339, OR668818,
OR866308, OR913396, OR9I13274; C. polyschoena: OR913340, —, OR866309,
OR913397, OR913275; C. prasina: MN760974, MN761582, GU173431, —, —; C.
pseudotristachya: OR913337, —, —, —, —; C. pudica: MN759801, MN762546, —,
—,—;C. pulchra: MN759942, —, MN763695, —, —; C. gingyangensis: MW458981,
MW459012, MW459078, OR913411, MW459043; C. rainbowii: KC122388,
KC122380, MN763804, —, —; C. reinii: MN760674, MN762213, —, —, —; C. rich-
ardsonii: MN759988, MN761635, GU173477, —, —; C. sartwelliana: MN760980,
MN761649, GU173500, —, —; C. shangchengensis: MN759909, MN761578,
MN763099, —, —; C. shortiana: DQ115273, DQ115272, —, —, —; C. sprengelii:
MN760547, MN761413, GU173599, —, —; C. squarrosa: MN760967, MN761 563,
GU173603, —, —; C. stenocarpa: MN760110, MN762363, MN763676, —, —;
C. strigosa: MN761116, MN762211, MN763217, —, —; C. stylosa: MN761010,
MN761414, GU173641, —, —; C. tangulashanensis: MN760545, MN762246,
MN763687, —, —; C. tomentosa: MN759974, MN762344, MN763732, —, —;
C. transversa: MT920755, MT920786, MT920828, MT920863, MT920896;
C. triquetra: MN759951, MN761924, GU173722, —, —; C. tristachya: OR913349,
OR668827, OR866316, OR9I13401, OR913283; C. tristachya var. pocilliform-
is: OR913350, OR668828, —, ORI13402, OR9I13284; C. tristis: MN760116,
MN762438, MN763724, —, —; C. truncatigluma: OR913351, OR668829,
OR866317, OR913403, OR9I13285; C. ulobasis: MN760557, —, MN763109, -—,
—; C. umbrosa: MN759791, MN762384, MN763798, —, —; C. umbrosa subsp.
huetiana: MN760662, MN762420, MN763658, —, —; C. vestita: MN760966,
MN761634, GU173772, —, —; C. yandangshanica: MN760559, MN761541,
MN763047, —, —; C. yangchunensis 1: PQ539317, PQ535884, PQ539320,
PQ539324, PQ539328, Guangdong, Yangchun, X.F. Jin & YF. Lu 5196; C. yangc-
hunensis 2: PQ539315, PQ535882, —, PPQ539322, PQ539326, Guangdong, Yang-
chun, D.C. Meng 2024010201; C. yangchunensis 3: PQ539316, PQ535883,
PQ539319, PQ539323, PQ539327, Guangdong, Yangchun, D.C. Meng s.n.

PhytoKeys 251: 257-270 (2025), DOI: 10.3897/phytokeys.251.142179 270