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Making Carex monophyletic (Cyperaceae, tribe Cariceae): A new broader circumscription

July 2015
Botanical Journal of the Linnean Society 179(1)

DOI:10.1111/boj.12298

Authors:

Kerry A. Ford

Manaaki Whenua - Landcare Research

Modesto Luceño Garcés

Pablo de Olavide University

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Carex (Cyperaceae), with an estimated 2000 species, nearly cosmopolitan distribution and broad range of habitats, is one of the largest angiosperm genera and the largest in the temperate zone. In this article, we provide argument and evidence for a broader circumscription of Carex to add all species currently classified in Cymophyllus (monotypic), Kobresia (c. 60 species), Schoenoxiphium (c. 15 species) and Uncinia (c. 70 species) to those currently classified as Carex. Carex and these genera comprise tribe Cariceae (subfamily Cyperoideae, Cyperaceae) and form a well-supported monophyletic group in all molecular phylogenetic studies to date. Carex as defined here in the broad sense currently comprises at least four clades. Three are strongly supported (Siderostictae, core Vignea and core Carex), whereas the caricoid clade, which includes all the segregate genera, receives only weak to moderate support. The caricoid clade is most commonly split into two clades, one including a monophyletic Schoenoxiphium and two small clades of species of Carex s.s., and the other comprising Kobresia, Uncinia and mostly unispicate species of Carex s.s. Morphological variation is high in all but the Vignea clade, making it extremely difficult to define consistent synapomorphies for most clades. However, Carex s.l. as newly circumscribed here is clearly differentiated from the sister groups in tribe Scirpeae by the transition from bisexual flowers with a bristle perianth in the sister group to unisexual flowers without a perianth in Carex. The naked female flowers of Carex s.l. are at least partially enclosed in a flask-shaped prophyll, termed a perigynium. Carex s.s. is not only by far the largest genus in the group, but also the earliest published name. As a result, only 72 new combinations and 58 replacement names are required to treat all of tribe Cariceae as a single genus Carex. We present the required transfers here, with synonymy, and we argue that this broader monophyletic circumscription of Carex reflects the close evolutionary relationships in the group and serves the goal of nomenclatural stability better than other possible treatments.

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Making Carex monophyletic (Cyperaceae, tribe

Cariceae): a new broader circumscription

GLOBAL CAREX GROUP

Received 17 March 2015; revised 4 May 2015; accepted for publication 13 May 2015

Carex (Cyperaceae), with an estimated 2000 species, nearly cosmopolitan distribution and broad range of habitats,

is one of the largest angiosperm genera and the largest in the temperate zone. In this article, we provide argument

and evidence for a broader circumscription of Carex to add all species currently classiﬁed in Cymophyllus (monotypic),

Kobresia (c. 60 species), Schoenoxiphium (c. 15 species) and Uncinia (c. 70 species) to those currently classiﬁed as

Carex.Carex and these genera comprise tribe Cariceae (subfamily Cyperoideae, Cyperaceae) and form a well-

supported monophyletic group in all molecular phylogenetic studies to date. Carex as deﬁned here in the broad sense

currently comprises at least four clades. Three are strongly supported (Siderostictae, core Vignea and core Carex),

whereas the caricoid clade, which includes all the segregate genera, receives only weak to moderate support. The

caricoid clade is most commonly split into two clades, one including a monophyletic Schoenoxiphium and two small

clades of species of Carex s.s., and the other comprising Kobresia,Uncinia and mostly unispicate species of Carex s.s.

Morphological variation is high in all but the Vignea clade, making it extremely difficult to deﬁne consistent

synapomorphies for most clades. However, Carex s.l. as newly circumscribed here is clearly differentiated from the

sister groups in tribe Scirpeae by the transition from bisexual ﬂowers with a bristle perianth in the sister group to

unisexual ﬂowers without a perianth in Carex. The naked female ﬂowers of Carex s.l. are at least partially enclosed

in a ﬂask-shaped prophyll, termed a perigynium. Carex s.s. is not only by far the largest genus in the group, but also

the earliest published name. As a result, only 72 new combinations and 58 replacement names are required to treat

all of tribe Cariceae as a single genus Carex. We present the required transfers here, with synonymy, and we argue

that this broader monophyletic circumscription of Carex reﬂects the close evolutionary relationships in the group and

London, Botanical Journal of the Linnean Society, 2015, ••, ••–••.

*Corresponding author. Marcia J. Waterway. Current address: Plant Science Department, McGill University, 21 111

Lakeshore Road, Ste-Anne-de-Bellevue, QC, Canada, H9X 3V9. E-mail: marcia.waterway@mcgill.ca

†This paper was written and compiled by members of the Global Carex Group who all contributed in various ways: M. J.

Waterway wrote the draft text based on formal and informal discussions among the group, managed the revisions and prepared

the ﬁgures; K. A. Ford, M. Luceño, S. Martín-Bravo, J. R. Starr, K. L. Wilson, O. Yano and S. R. Zhang (listed alphabetically)

participated in the discussions, provided advice and editorial comments on the manuscript, did the nomenclatural research and

proposed the new names and new combinations, coordinated by E. H. Roalson; W. S. Alverson, L. P. Bruederle, J. J. Bruhl, K.-S.

Chung, T. S. Cochrane, M. Escudero, B. A. Ford, S. Gebauer, B. Gehrke, M. Hahn, A. L. Hipp, M. H. Hoffmann, T. Hoshino, P.

Jiménez-Mejías, X.-F. Jin, J. Jung, S. Kim, E. Maguilla, T. Masaki, M. Míguez, A. Molina, R. F. C. Naczi, A. A. Reznicek, P. E.

Rothrock, D. A. Simpson, D. Spalink, W. W. Thomas and T. Villaverde (listed alphabetically) participated in the discussions and

provided advice and editorial comments on the manuscript. Affiliations, listed alphabetically by country: J. J. Bruhl, University

of New England, Armidale, NSW, Australia; K. L. Wilson, National Herbarium of New South Wales, Sydney, NSW, Australia; B.

A. Ford, University of Manitoba, Winnipeg, MB, Canada; J. R. Starr, University of Ottawa, Ottawa, ON, Canada; X.-F. Jin,

Hangzhou Normal University, Hangzhou, Zhejiang, China; S. R. Zhang, Institute of Botany, Chinese Academy of Sciences, Beijing,

China; S. Gebauer and M. H. Hoffmann, Martin Luther University Halle-Wittenberg, Halle, Germany; B. Gehrke, Johannes

Gutenberg-Universität, Mainz, Germany; O. Yano, T. Hoshino and T. Masaki, Okayama University of Science, Okayama, Japan;

K. A. Ford, Landcare Research, Lincoln, New Zealand; K.-S. Chung, Jungwon University, Goesan, Chungbuk, South Korea; J.

Jung and S. Kim, Sungshin Women’s University, Seoul, South Korea; M. Escudero, Doñana Biological Station CSIC, Seville,

Spain; M. Luceño, E. Maguilla, S. Martín-Bravo, M. Míguez and T. Villaverde, Pablo de Olavide University, Seville, Spain; A.

Molina, University of León, Spain; D. A. Simpson, Royal Botanic Gardens, Kew, Richmond, Surrey, UK; L. P. Bruederle,

University of Colorado-Denver, Denver, CO, USA; M. Hahn and A. L. Hipp, Morton Arboretum, Lisle, IL, USA; P. E. Rothrock,

Indiana University Herbarium, Bloomington, IN, USA; A. A. Reznicek, University of Michigan Herbarium, Ann Arbor, MI, USA;

R. F. C. Naczi and W. W. Thomas, New York Botanical Garden, Bronx, NY, USA; P. Jiménez-Mejías and E. H. Roalson, Washington

State University, Pullman, WA, USA; W. S. Alverson, T. S. Cochrane and D. Spalink, University of Wisconsin, Madison, WI, USA.

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Botanical Journal of the Linnean Society, 2015, ••, ••–••. With 2 ﬁgures

The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, ••, ••–••

This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and

reproduction in any medium, provided the original work is properly cited.

ADDITIONAL KEYWORDS: classiﬁcations – Cymophyllus – Cyperoideae – generic limits – inﬂorescence

morphology – Kobresia – new combinations – nomenclature – phylogenetic relationships – Schoenoxiphium –

taxonomic revision – Uncinia –Vesicarex.

INTRODUCTION

From the initial naming of 29 species of Carex L. in

Species Plantarum (Linnaeus, 1753), the genus has

grown to >1830 accepted species (Govaerts et al.,

2013). Carex is placed in tribe Cariceae, with Kobresia

Willd., Uncinia Pers., Schoenoxiphium Nees and

Cymophyllus Mack.; together they comprise c. 2150

species (Goetghebeur, 1998). Bruhl included a sixth

genus, the monotypic Vesicarex Steyerm., in the tribe

(Bruhl, 1995). In the most comprehensive global mono-

graph of tribe Cariceae, Kükenthal (1909) recognized

four genera [Carex,Kobresia (as ‘Cobresia’), Schoenox-

iphium and Uncinia] and classiﬁed 793 broadly

deﬁned species into 69 sections of Carex distributed

across four subgenera that differed in inﬂorescence

structure, branching, gender distribution and number

of spikes (Kükenthal, 1909). Although Kükenthal’s

classiﬁcation was criticized, particularly for its treat-

ment of unispicate species as a distinct subgenus

(Kreczetovicz, 1936; Ohwi, 1936; Nelmes, 1952; Kern,

1958; Hamlin, 1959; Koyama, 1961), modiﬁcations of

Kükenthal’s classiﬁcation continue to be used to organ-

ize large regional ﬂoristic manuals (Chater, 1980;

Haines & Lye, 1983; Egorova, 1999; Dai & Liang, 2000;

Ball, Reznicek & Murray, 2002; Luceño, Escudero &

Jiménez-Mejías, 2008; Dai et al., 2010; Hoshino,

Masaki & Nishimoto, 2011). With nomenclatural cor-

rections, the four subgenera used explicitly or indi-

rectly to order the sections of Carex s.s. in most modern

ﬂoristic treatments are subgenus Psyllophora (Degl.)

Peterm. (= subgenus Primocarex Kük.), subgenus

Vignea (P.Beauv ex T.Lestib.) Peterm., subgenus

Vigneastra (Tuck.) Kük. [= subgenus Indocarex (Baill.)

Kük.] and subgenus Carex (= subgenus Eucarex

Peterm.). We use these subgeneric names to refer to

groups in the traditional classiﬁcation.

Ninety years after Kükenthal’s monograph, the ﬁrst

molecular phylogenetic analyses of tribe Cariceae were

published (Starr, Bayer & Ford, 1999; Yen & Olmstead,

2000; Roalson, Columbus & Friar, 2001). These early

studies were based on few genes and limited sampling,

but already they suggested that, although Cariceae

was monophyletic, Carex and Kobresia were polyphy-

letic or paraphyletic. Uncinia and Schoenoxiphium

were each apparently monophyletic, but nested in

Carex, as was the monotypic genus Cymophyllus. The

only traditional subgenus of Carex that was largely

monophyletic in any of these early studies was subge-

nus Vignea. Larger studies of phylogenetic relation-

ships in Cyperaceae, incorporating additional gene

regions, also strongly supported a monophyletic tribe

Cariceae. This tribe has been suggested by most

studies to be sister to tribe Scirpeae or nested in it

(Muasya et al., 1998, 2009; Simpson et al., 2007;

Escudero & Hipp, 2013; Hinchliff & Roalson, 2013;

Jung & Choi, 2013; Léveillé-Bourret et al., 2014), as

predicted by evidence from associations with parasitic

smut fungi in the genus Anthracoidea (Kukkonen &

Timonen, 1979), rather than sister to previously sug-

gested tribes having unisexual ﬂowers, such as Scle-

rieae (Haines & Lye, 1972; Smith & Faulkner, 1976) or

Rhynchosporeae (Koyama, 1961).

A broader and more representative sampling of tribe

Cariceae using DNA from both nuclear and plastid

genomes (Waterway & Starr, 2007) revealed three

major clades that roughly corresponded to: (1) subge-

nus Vignea, hence named the Vignea clade; (2) subgen-

era Carex and Vigneastra, named the core Carex clade;

and (3) subgenus Psyllophora plus Cymophyllus,

Kobresia,Schoenoxiphium and Uncinia, named

the caricoid clade (Fig. 1). The ﬁrst two were strongly

supported in parsimony and Bayesian analyses,

whereas the caricoid clade received only moderate

support. In the caricoid clade, two clades were strongly

supported in the Bayesian analysis: one with Schoe-

noxiphium and a few Carex spp. (the Schoenoxiphium

clade) and one with Kobresia,Uncinia,Cymophyllus

and several members of Carex subgenus Psyllophora

(the core unispicate clade). Starr, Harris & Simpson

(2003, 2004, 2008) further explored the caricoid clade,

noting a major difference between dioecious unispicate

and androgynous species, and providing additional

support for the monophyly of Uncinia. Dioecious

unispicate Carex spp. showed affinities to multispicate

species in either the Vignea clade or the core Carex

clade, and the androgynous species formed part of the

caricoid clade with androgynous species of Cymophyl-

lus,Kobresia,Schoenoxiphium and Carex, in phyloge-

netic trees based on internal transcribed spacer (ITS)

and external transcribed spacer (ETS) data (Starr

et al., 2004). Further detailed study of Schoenox-

iphium and the caricoid clade supported the mono-

phyly of the African genus Schoenoxiphium and

demonstrated sister group relationships of two other

small clades of Carex spp. to Schoenoxiphium (Gehrke

et al., 2010). The rest of the caricoid clade (core unispi-

cate clade) was moderately supported in that analysis,

2GLOBAL CAREX GROUP

The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, ••, ••–••

but there was no support for a closer relationship to the

Schoenoxiphium clade than to the Vignea or core Carex

clades (Gehrke et al., 2010).

The discovery that section Siderostictae Franch.

ex Ohwi, traditionally classiﬁed in subgenus Carex,

formed a clade sister to all other species in tribe

Cariceae conﬁrmed that Carex in the traditional

sense is a paraphyletic group with all other genera

of tribe Cariceae nested in it (Waterway, Hoshino &

Masaki, 2009). This Siderostictae clade was recently

expanded to include species from two sections pre-

viously classiﬁed in Carex subgenus Vigneastra (sec-

tions Hemiscaposae C.B.Clarke and Surculosae

Raymond) based on the analysis of ITS and trnL-

trnF sequences (Yano et al., 2014). Although these

new additions are broad-leaved species, like most

species in section Siderostictae, they have inﬂores-

cences with more complex branching, thus expand-

ing the range of variation found in this clade that is

sister to the rest of tribe Cariceae. Taken together,

these results from molecular systematic studies

show clearly that Carex as traditionally deﬁned is

not monophyletic, nor are any of the traditional sub-

genera except Vignea (Fig. 1). Furthermore, contin-

ued recognition of those genera that appear to be

monophyletic in tribe Cariceae (Uncinia and Schoe-

noxiphium) would leave Carex paraphyletic and

Kobresia polyphyletic.

Caricoid Clade

Schoenoxiphium

C. andina Clade

C. distachya

Clade

Uncinia

Cymophyllus plus Carex

sect. Phyllostachyae

several unispicate

Kobresia species

Carex subg. Psyllophora

Kobresia

Carex sections

Siderostictae,

Hemiscaposae

Surculosae

Schoenoxiphium

plus a few Carex

subg. Carex and

Psyllophora

Cymophyllus, Kobresia,

Uncinia, majority of

Carex subg.

-Psyllophora

most of subg. Vignea

+ a few dioecious

subg. Psyllophora

most subg. Carex and

subg. Vigneastra +

dioecious subg.

Psyllophora

Core Carex Clade Core Vignea Clade

Core Unispicate

Clade

Schoenoxiphium

Clade

Siderostictae

Clade

Caricoid Clade

Tribe

Scirpeae

Outgroup

Figure 1. Generalized phylogenetic tree of Cyperaceae tribe Cariceae based on molecular phylogenetic studies to date.

Full lines show relationships that are supported by all or most studies. Dotted branches show relationships that are

frequently seen but more inconsistent among studies. Branches with consistently high bootstrap support are indicated

with a ﬁlled ellipse.The number of subclades shown within the caricoid clade is arbitrary; resolution and support in this

clade are inconsistent among studies and so a large polytomy is shown with many individual small clades grouping two

or three taxa. Larger triangles in this polytomy indicate clades that comprise more than three taxa and show fairly

consistent support across studies. It should be noted that the newly reported Hypolytroides clade (Starr et al., 2015) is

sister to the Siderostictae clade shown here and that the Siderostictae + Hypolytroides clade has the same sister

relationship to the rest of tribe Cariceae as shown here for the Siderostictae clade.

MAKING CAREX MONOPHYLETIC 3

The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, ••, ••–••

It is apparent that Carex and tribe Cariceae are

overdue for a new classiﬁcation that better reﬂects

evolutionary relationships. The reclassiﬁcation of tribe

Cariceae was discussed at length at an international

gathering of Cyperaceae specialists in 2011 at a

BioSynC meeting in Chicago and again with an even

larger group of Cyperaceae specialists at the Monocots

V meeting in New York in 2013. The consensus at both

meetings was to broaden the circumscription of Carex

to include all species in tribe Cariceae, thus forming a

monophyletic genus Carex with >2000 species. This

approach was chosen as that most likely to provide

nomenclatural stability. There was some question in

2011 whether increased sampling in China and South-

East Asia would reveal new clades that should be

segregated from Carex or help to deﬁne clear groupings

in the caricoid clade. However, even with much more

extensive sampling from China, South-East Asia and

Africa since 2011 (Luceño et al., 2013; Waterway et al.,

2013; Zhang et al., 2013; Yano et al., 2014; Starr,

Jansen & Ford, 2015), including more complete studies

of Kobresia and Schoenoxiphium, the conclusion that a

single monophyletic Carex would be the best classiﬁ-

cation was strengthened rather than weakened. This

article is the ﬁrst in a series of planned contributions

from the Global Carex Group to completely reclassify

this expanded Carex at the sectional level. Our goals in

this paper are to provide a brief background on the

morphology of tribe Cariceae and its classiﬁcation

history, to summarize the molecular and morphological

evidence for treating tribe Cariceae as the single genus

Carex and to make the required nomenclatural

changes.

Although a broader circumscription of Carex is in

the interest of long-term nomenclatural stability, 130

nomenclatural changes are needed at this time to

change the circumscription. Species of Cymophyllus

and Vesicarex already have valid names as Carex spp.,

as do several species of Kobresia,Schoenoxiphium

and Uncinia. Many other needed changes are simply

new combinations, because several speciﬁc epithets

currently used in Kobresia (23), Schoenoxiphium (six)

and Uncinia (27) have never been used in Carex. For

cases in which the speciﬁc epithets are already occu-

pied, 58 are here given replacement names in Carex

and 16 others adopt the speciﬁc, varietal or forma

epithet from a previously published synonym. The

changes are detailed in the taxonomic section below,

including synonymy and notes on geographical distri-

bution and any nomenclatural issues.

MATERIAL AND METHODS

We reviewed the major literature on the classiﬁcation

in Cyperaceae tribe Cariceae, including papers pro-

posing evolutionary theories related to classiﬁcation.

We also reviewed recent work on inﬂorescence mor-

phology and all molecular phylogenetic studies to

date to provide a synthetic view of current evidence

for phylogenetic relationships in the group. An initial

list of names, with geographical distributions, for

currently recognized species of Cymophyllus,Kobre-

sia,Schoenoxiphium and Uncinia was constructed

from the World Checklist of Cyperaceae (Govaerts

et al., 2013) and then modiﬁed by those in our group

most familiar with each genus (S.R.Z. and O.Y. for

Kobresia, K.A.F., J.R.S. and K.L.W. for Uncinia and

M.L. and S.M.-B. for Schoenoxiphium) to create the

ﬁnal taxonomic listing with new combinations and

new names. The names fall into three categories: (1)

species that already have a valid name in Carex; (2)

species with speciﬁc epithets that are available in

Carex; and (3) species with speciﬁc epithets that are

not available in Carex. New combinations are made

where appropriate and new names are created where

a speciﬁc epithet was not available.

DISCUSSION

MORPHOLOGICAL VARIATION IN TRIBE CARICEAE

The inﬂorescence structure is complex and variable in

tribe Cariceae and has long been seen as a rich source

of taxonomic characters. An understanding of the

terminology used to describe inﬂorescence structure

is critical in evaluating theories of relationship that

underpin the various classiﬁcations proposed for the

group. Features of the inﬂorescence have been impor-

tant in constructing classiﬁcations and identiﬁcation

keys, especially because vegetative features are quite

similar across the group, with the exception of the

unusual leaves of Cymophyllus, which lack a midrib

(Reznicek, 1990), and some broad-leaved, pseudo-

petiolate Carex spp. from South-East Asia (Raymond,

1959). The interpretation of the ﬂowers, spikelets and

overall inﬂorescence architecture of Cyperaceae and

tribe Cariceae goes back to the early 19th century

(e.g. Kunth, 1835; Caruel, 1867) and has been a

popular topic since then (e.g. Snell, 1936; Blaser,

1944; Levyns, 1945; Holttum, 1948; Kukkonen, 1967,

1984, 1990; Kern, 1974; Eiten, 1976; Smith &

Faulkner, 1976; Goetghebeur, 1986; Reznicek, 1990;

Bruhl, 1991; Timonen, 1998; Richards, Bruhl &

Wilson, 2006; Prychid & Bruhl, 2013). Detailed new

typological interpretations of inﬂorescence structure

in tribe Cariceae (Vegetti, 2002, 2003; Guarise &

Vegetti, 2008; Molina, Acedo & Llamas, 2012;

Reutemann et al., 2012) and ontogenetic studies of

ﬂoral development in Cariceae using scanning elec-

tron microscopy published during the last decade

(Vrijdaghs et al., 2009, 2010; Gehrke et al., 2012)

demonstrate the similarities in basic architecture of

4GLOBAL CAREX GROUP

The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, ••, ••–••

the inﬂorescence. The terminology used to describe

inﬂorescences in Cariceae has been applied inconsist-

ently (Reznicek, 1990; Kukkonen, 1994; Vegetti, 2002;

Molina et al., 2012; Reutemann et al., 2012) and

efforts to apply typological principles to inﬂorescence

description strictly, whilst demonstrating the similar-

ity in inﬂorescence architecture across Cariceae, have

also resulted in a proliferation of terminology unfa-

miliar to non-specialists. Reconciling this terminology

is an ongoing issue for cyperologists and requires

additional study and discussion. Here, we provide

only enough background to make our arguments for a

broader circumscription of Carex clear.

In tribe Cariceae, the ﬂowers are normally uni-

sexual and lack a perianth. Their structure is simple:

each staminate ﬂower comprises three stamens

(rarely fewer) and each pistillate ﬂower comprises one

uniovulate ovary arising from an annular primor-

dium, with a single style and two or three stigmas

(Vrijdaghs et al., 2009; Reynders et al., 2012). Com-

plications arise in the ways in which these simple

unisexual ﬂowers are arranged into inﬂorescences.

The spikelet has traditionally been considered as

the basic unit of the inﬂorescence in Cyperaceae

(Snell, 1936; Holttum, 1948; Kukkonen, 1967). In

most species of Cyperaceae subfamily Cyperoideae,

perfect or unisexual ﬂowers are arranged spirally or

distichously on a spikelet axis, the rachilla, each

ﬂower being subtended by a scale-like ﬂoral bract,

usually called a glume or a scale (Fig. 2A, B). Spike-

lets in many tribes of Cyperoideae, including those in

Cariceae, are considered polytelic (indeterminate). A

prophyll, which usually encircles the base of the

rachilla, is the ﬁrst adaxial bract produced on each

spikelet. This small prophyll, often called a cladopro-

phyll to emphasize its position on an axis, is in

addition to the larger, often foliose bract that arises

from the main inﬂorescence axis and subtends a

spikelet or group of spikelets.

Spikelets in Cariceae differ from those of more

typical species of Cyperoideae in two important ways.

First, the prophyll arising from the rachilla is modi-

ﬁed into an enclosing sac-like or ﬂask-shaped struc-

ture variously called a perigynium, utricle or

utriculiform prophyll (Fig. 2C–G). A similar enclosing

prophyll is found around the proximal ﬂower on

spikelets in Dulichium Pers. (tribe Dulicheae), but

the ﬂowers distal to this one on a Dulichium spikelet

are each subtended only by a glume. In Cariceae, each

female ﬂower is enclosed by a perigynium that is most

often closed except for an apical oriﬁce from which the

style and stigmas emerge. However, the perigynium is

only partially sealed in some Kobresia spp. (Fig. 2E)

and the oriﬁce may be quite wide on some perigynia

in Schoenoxiphium (Fig. 2G). Kükenthal used the

term utricle instead of perigynium, and many authors

of ﬂoristic treatments have followed his example (e.g.

Kern & Noteboom, 1979; Chater, 1980; Egorova, 1999;

Luceño et al., 2008; Dai et al., 2010). However, in a

broader botanical context, the term utricle may be

misleading, because it refers to a type of fruit and

thus to ovary tissue (Davis & Cullen, 1989; Harris &

Harris, 1994; Spjut, 1994), not to a type of bract.

Perigynium has also been used to refer to non-

homologous structures, such as the stem tissue sur-

rounding the ‘perianth’ in some foliose liverworts (e.g.

Hentschel et al., 2006), and in Cyperaceae, to the

cupule, interpreted as the perianth, surrounding the

ovary in some Scleria spp. (Barros, 1960), but it has

been most widely and consistently used, especially in

North America, for the prophyllar bract surrounding

the ovary in species of Cariceae (e.g. Tuckerman,

1843; Bailey, 1886; Holm, 1903; Ivanova, 1939; Bruhl,

1995; Ball & Reznicek, 2002; Vrijdaghs et al., 2010;

Hoshino et al., 2011). Here, we use the term perigy-

nium, but recognize that both terms are widely used

in ﬂoristic treatments and should be considered as

synonyms when applied to Cariceae.

Second, the rachilla is reduced compared with that

of other Cyperaceae, often bearing only a single

female ﬂower that is at least partially enclosed by the

perigynium. The rachilla is vestigial in Cymophyllus

and absent or reduced to a tiny structure in most

species of Carex s.s. (Reznicek, 1990; Vrijdaghs et al.,

2010; Fig. 2C), but elongated to varying extents in

Kobresia,Schoenoxiphium,Uncinia and a few species

of Carex s.s. (Fig. 2D–G). In Schoenoxiphium, stami-

nate ﬂowers, subtended by glumes, may be produced

distally on the rachilla, and these protrude from the

perigynium with the stigmas (Fig. 2F, G). Distal

staminate ﬂowers or vestigial remnants of them or

their glumes can also be found in many Kobresia spp.

(Fig. 2E) and even rarely in Carex s.s. (Jin, Ding &

Zheng, 2005). The rachilla in Uncinia extends beyond

the perigynium as a hook-shaped tip that aids in

dispersal (Fig. 2D), but only rarely bears staminate

ﬂowers (Hamlin, 1959). Thus, most spikelets in

Cariceae are much reduced compared with those in

other Cyperaceae, appearing from the outside as per-

igynia with styles, stigmas and sometimes a rachilla

bearing staminate ﬂowers protruding from the apical

opening.

Much of the confusion surrounding the use of the

terms spikelet and spike in Cariceae arises because

these reduced spikelets of Cariceae, subtended by

glumes, are themselves spirally arranged on lateral

branches into spike-like inﬂorescences that resemble

the spikelets of other genera of Cyperaceae (compare

Fig. 2B with 2H, I, K, L). Reznicek (1990) chose to

abandon the term spikelet altogether, because it is

sometimes used as described above and other times

used incorrectly (notably by Kükenthal, 1909) to refer

MAKING CAREX MONOPHYLETIC 5

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Figure 2. See caption on next page.

6GLOBAL CAREX GROUP

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to a lateral inﬂorescence unit that is a spike of spike-

lets which may also bear male ﬂowers directly on the

same axis. Rather than abandoning the term, we

refer to perigynia and their enclosed ﬂower-bearing

axes as reduced spikelets in tribe Cariceae, thus

maintaining a link to the equivalent structure in

other groups of Cyperaceae (Timonen, 1998).

Most authors subsequent to Kükenthal (1909)

referred to the set of ﬂowers borne at the tip of the

main culm as the terminal spike or spikelet, and to

the aggregations of ﬂowers on ﬁrst-order lateral

branches as lateral spikes. In Carex,Cymophyllus

and Uncinia, these so-called spikes are really spikes

of reduced spikelets or of mixed male ﬂowers and

reduced spikelets, except for the terminal one, which

often bears only male ﬂowers (Fig. 2H, I). Spikes may

be unisexual, androgynous (perigynia proximal and

staminate ﬂowers distal on the spike axis), gynecan-

drous (staminate ﬂowers proximal and perigynia

distal), mesogynous (staminate ﬂowers both proximal

and distal to perigynia), mesandrous (perigynia both

proximal and distal to staminate ﬂowers) or with

alternating staminate ﬂowers and perigynia on the

spike axis (Eiten, 1976). Early choices in identiﬁca-

tion keys for Carex s.s. often distinguish between

unispicate and multispicate inﬂorescences.

Although terms such as spike, unispicate and mul-

tispicate are widely used and relatively easy to under-

stand for the large majority of Carex s.s.,Cymophyllus

and Uncinia spp., they are not technically correct and

can be misleading when trying to interpret homology

in inﬂorescence structure. What is generally called a

spike in Cariceae is actually a spike of spikelets or

stachyodium (Reutemann et al., 2012). This may be

further ramiﬁed in Schoenoxiphium if additional per-

igynia are produced on the primary rachilla axis

emerging from a perigynium (Levyns, 1945; Gehrke

et al., 2010) (Fig. 2G). Molina et al. (2012) used the

term pseudospike rather than spike to indicate that

these aggregations of ﬂowers are not true spikes

because they often include ﬂowers that are at different

branching orders in the inﬂorescence. That is, the axis

Figure 2. Diagrammatic representations, not to scale, of ﬂower and inﬂorescence structure in Cyperaceae tribe Cariceae

with comparison to tribe Scirpeae in A and B. Axes are exaggerated in length and spacing of ﬂowers so that the structure

can be clearly seen. Diagrams above the horizontal line show ﬂower and spikelet structure, whereas those below the line

show the arrangement of the spikelets and staminate ﬂowers into inﬂorescences. It should be noted that, for simplicity,

all ovaries are drawn with three stigmas, although two stigmas are also frequently seen, especially in Carex subgenera

Vignea and Psyllophora. All axes are shown in progressively darker shades of grey indicating higher order branching, with

the highest order branches (rachillae) shown in black. All prophylls (including cladoprophylls and perigynia) are also

drawn in black. Scale-like glumes that subtend staminate ﬂowers or perigynia are shown in medium grey, whereas

inﬂorescence bracts at the base of primary and secondary inﬂorescence axes are shown in light grey. The perianth (only

in A, B) is also shown in light grey. The gynoecium is shown in a slightly darker shade of grey than the androecium. A,

Flower structure of a typical species in tribe Scirpeae. B, Spikelet structure in tribe Scirpeae. C, One-ﬂowered pistillate

spikelet typical of Carex s.s. and Cymophyllus, but also sometimes found in Schoenoxiphium and Kobresia. Here and

elsewhere, the black ellipse represents a ﬂask-like perigynium that is closed except for an apical oriﬁce through which

styles, stigmas and sometimes rachillae emerge. D, One-ﬂowered pistillate spikelet of Uncinia, showing hooked rachilla

protruding from the perigynium. E, Bisexual spikelet with perigynium not fully sealed and containing a fertile pistillate

ﬂower and a small staminate ﬂower on the persistent rachilla, typically found in some Kobresia inﬂorescences. F, Bisexual

spikelet with one fertile pistillate ﬂower and multiple staminate ﬂowers borne on an elongated rachilla protruding from

the perigynium, found in Schoenoxiphium and some Kobresia. The number of staminate ﬂowers varies from one to several.

G, Spike of spikelets resulting from additional branching of the primary rachilla after producing the ﬁrst perigynium at

the base, resulting in two additional spikelets, one pistillate and one bisexual, proximal to the distal staminate ﬂowers

of the primary rachilla (redrawn based on ﬁg. 3 in Gehrke et al., 2012). This proliferation of branching of the rachilla after

producing a fertile perigynium can be found in Schoenoxiphium and blurs the distinction between rachis and rachilla.

Note that what we label as the primary rachilla here is called the rachis of the spike of spikelets in Gehrke et al. (2012).

We show it in black and refer to it as a primary rachilla to emphasize the structural similarity between rachis and rachilla

in this case. H–L, Examples of various types of inﬂorescence or parts thereof, using both traditional and typological

terminology to label them. H, Unispicate inﬂorescence of Uncinia, comprising only the main ﬂorescence, which is an

androgynous terminal spike. This type of unispicate inﬂorescence is also found in Cymophyllus and Carex subgenus

Psyllophora, but with short or vestigial rachillae rather than the hooked rachillae of Uncinia. I, Typical synﬂorescence

found in Carex subgenus Carex, showing the main ﬂorescence (terminal spike) and two paracladia to illustrate terms

describing these structures. J, Typical example of proximal paracladium (lowest ﬁrst-order branch and subtending bract)

in Schoenoxiphium ecklonii (redrawn based on Levyns, 1945). K, Main ﬂorescence of the unispicate Kobresia myosuroides

(redrawn based on Kern, 1958). L, Proximal paracladium of the multispicate Kobresia simpliciuscula (redrawn based on

Kern, 1958).

MAKING CAREX MONOPHYLETIC 7

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of the so-called spike may bear male ﬂowers directly,

but each female ﬂower is borne in a perigynium on a

rachilla that is actually a higher order branch

(Fig. 2H–L). We acknowledge this problem here, but

continue to use the term spike because it is widely used

in nearly all ﬂoristic treatments, and to avoid awkward

terminology, such as uni-pseudospicate and multi-

pseudospicate, in reference to whole inﬂorescences.

Flowering culms without lateral branching can be

called spiciform, those with ﬁrst-order lateral branch-

ing are often referred to as racemose or, more properly,

as racemiform, whereas those with higher order

branching are often called paniculate or, more prop-

erly, paniculiform (Molina et al., 2012).

Recent detailed studies of inﬂorescences in Cyper-

aceae, including those in Cariceae, follow typological

methods and terminology that were originally devel-

oped for dicots by Troll (1964) and Weberling (1989) to

describe sedge inﬂorescences in ways that make it

easier to assess homology. Many of these terms are

familiar only to specialists, and so only essential ones

will be used here. Following recent interpretations

(Guarise & Vegetti, 2008; Molina et al., 2012;

Reutemann et al., 2012), each ﬂowering culm in a

sedge plant is a synﬂorescence that ends in a terminal

aggregation of usually staminate ﬂowers that repre-

sents the main ﬂorescence (Fig. 2H, I, K). Below this

main ﬂorescence (a terminal spike in the older termi-

nology), a ﬂowering culm may have an enrichment

zone (= paracladial zone) where it produces one or

more lateral branches, each of which may also end in

a terminal set of (usually staminate) ﬂowers, termed

a co-ﬂorescence (Fig. 2I, terminus of distal paracla-

dium; Fig. 2J, K, terminus of proximal paracladium).

Each lateral branch, including the subtending bract

on the main axis and prophyll on the new axis, is

called a paracladium (Guarise & Vegetti, 2008;

Molina et al., 2012). Unless they remain dormant,

nodes in inﬂorescences of Cariceae have three

options: to produce lateral branches (axes) of the next

higher order; to produce rachillae (also axes) that

each bear at least one pistillate ﬂower and its sur-

rounding perigynium; or to produce staminate ﬂowers

directly on that branch. This recurring pattern may

be ramiﬁed into second-, third- or even higher order

branching in some species. The three node types have

been called inﬂorescence nodes, female ﬂower nodes

and male ﬂower nodes, respectively (Smith &

Faulkner, 1976). The ﬁrst two of these node types are

intrinsically similar in that the node is producing an

axis (lateral branch or rachilla) that will produce at

least one ﬂower, either on that axis or after branching

again.

Molina et al. (2012) applied this typological system

to 110 Carex spp. from all four traditional subgenera.

They treated the lateral branch or paracladium as the

basic unit of the inﬂorescence, similar to Timonen’s

(1998) emphasis on the axis as the basic unit, viewing

the inﬂorescence as a hierarchy of axes with recurring

developmental patterns (Timonen, 1998). Summariz-

ing the results of Molina et al. (2012) provides a

convenient opportunity to demonstrate how the ter-

minology is applied, although we use spike where

they used pseudospike. It is important to note that,

although Molina et al. (2012) indicated that each

reduced spikelet, consisting of perigynium, rachilla or

its vestiges, and unisexual ﬂower(s), and subtended

by a glume, can be thought of as the extreme reduc-

tion of a paracladium, they did not consider the

reduced spikelets as paracladia in their analyses.

Instead, they interpreted the single androgynous

spikes in subgenus Psyllophora as the main ﬂores-

cence on a ﬂowering culm that lacks paracladia and

has no bract subtending the spiciform inﬂorescence

(Fig. 2H, K). The main ﬂorescence in species of sub-

genus Carex was interpreted as the terminal spike,

which is entirely staminate in many species, but

gynecandrous, androgynous or entirely pistillate in

others. One to several ﬁrst-order paracladia, compris-

ing subtending bract, tubular cladoprophyll and a

pistillate or androgynous spike, are found below the

main ﬂorescence, resulting in a racemiform inﬂores-

cence (Fig. 2I), except in dioecious species, which are

unispicate and lack paracladia. Spikes are usually on

relatively long peduncles, may be entirely pistillate,

androgynous, entirely staminate or, much less com-

monly, gynecandrous, and have a tendency for stami-

nate ﬂowers to be found only in distal paracladia. The

few species examined from subgenus Vigneastra were

interpreted to have a paniculiform or racemiform

inﬂorescence with the main ﬂorescence androgynous

and with androgynous spikes on the paracladia,

which exhibit up to third-order branching from the

main axis. Of note in subgenus Vigneastra is that

each spike has a perigynium-like prophyll at its base,

in contrast with the tubular cladoprophyll found at

the base of the ﬁrst-order paracladium. The axis of

the ﬁrst-order paracladium is also usually peduncu-

late, and more than one axis is sometimes produced

at each node of the main axis, particularly from the

lower nodes. In subgenus Vignea, Molina et al. (2012)

interpreted the main ﬂorescence as the terminal

spike, which can be androgynous, gynecandrous or

entirely staminate or pistillate, as can the spikes in

the paracladia. The inﬂorescence can be spiciform to

paniculiform with ﬁrst-, second- or even third-order

paracladia branching below the main ﬂorescence.

Spikes in subgenus Vignea are generally compact and

sessile, subtended by relatively small, non-sheathing

bracts and few species have cladoprophylls.

The inﬂorescence structure in Cariceae is thus

based on a recurring architectural pattern in which

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each new lateral axis is subtended by a bract and

enclosed in a prophyll at its base (Levyns, 1945;

Timonen, 1998). The other recurring pattern is the

production of pistillate ﬂowers proximally and stami-

nate ﬂowers distally on each ﬂowering axis (Fig. 2H–

L), presumably under genetic or environmental

control, mediated through growth regulators (Gehrke

et al., 2010; Vrijdaghs et al., 2010). Ontogenetic

studies in Carex s.s. suggest that each node of a

ﬂowering axis is ﬂexible and can develop into either a

male ﬂower or a new ﬂowering axis bearing a single

female ﬂower enclosed in a perigynium (Vrijdaghs

et al., 2010). Ontogenetic study revealed no evidence

that staminate ﬂowers in Cariceae are highly reduced

spikelets, as assumed by Timonen (1998). Bracts sub-

tending the lateral and higher order spikes tend to

decrease in size from the base to apex of the culm and

vary from foliaceous to small scale-like structures.

Processes such as truncation or axis abortion, homog-

enization, initiation or suppression of paracladia,

elongation or reduction of internodes, increases or

decreases in degree of branching, reduction or sup-

pression of bracts or prophylls and others operate in

different ways and combinations to give the variety of

inﬂorescence forms seen in the tribe (Guarise &

Vegetti, 2008; Reutemann et al., 2012). These recur-

rent patterns are least obvious in subgenus Vignea,in

which the position effects on the production of female

vs. male ﬂowers appear to be minimal and additional

processes must be invoked to derive the various pat-

terns (for examples, see Timonen, 1998).

Inﬂorescence architecture in Cariceae thus differs

in fundamental ways from that of other Cyperaceae:

female ﬂowers are only produced when branching

occurs in the inﬂorescence, and the ﬁrst prophyll on

the new branch encloses the female ﬂower. Another

fundamental difference between Cariceae and other

Cyperaceae is the nature of the transition from a

non-ﬂowering to a ﬂowering branch. In most Cyper-

aceae, the transition occurs at the ﬁnal branching

event, i.e. the one giving rise to the rachilla, which

produces ﬂowers. All nodes above this one produce

either ﬂowers subtended by scale-like bracts, vari-

ously called glumes or scales, or sterile bracts. This

means that the ultimate level of inﬂorescence branch

in most Cyperaceae is the spikelet axis (rachilla).

Cariceae is quite different, because the transition to

ﬂowering in the lateral branches depends on whether

the ﬂower is male or female. Female ﬂowers are

produced only when branching occurs. This occurs on

the spikelet rachilla, i.e. on the rachilla that is

homologous to that in other Cyperaceae, but it can

also occur on both higher and lower order branches.

In Schoenoxiphium, the rachilla can branch after

producing the ﬁrst perigynium and produce one or

more additional perigynia, each enclosing a pistillate

ﬂower or both a pistillate ﬂower and male ﬂowers on

the new rachilla, before it produces the male ﬂowers

distally on the initial rachilla (Levyns, 1945; Gehrke

et al., 2012; Fig. 2G). Thus, the ﬁrst ﬂower-producing

rachilla is not always the ultimate branch in an

inﬂorescence in Cariceae as it is in most Cyperaceae.

Instead, the rachilla will branch each time it produces

another pistillate ﬂower. Ramiﬁcation of the rachilla

can be superposed on the primary inﬂorescence

branching pattern to produce a complex inﬂorescence

that is difficult to describe using traditional terms

(Fig. 2G). Also, in Schoenoxiphium, a perigynium can

be produced on a lower order axis than the spikelet,

e.g. at the base of a ﬁrst- or second-order paracla-

dium. Whether or not this perigynium contains a

pistillate ﬂower appears to be determined by its posi-

tion in the inﬂorescence, the likelihood being greater

in proximal than in distal positions (Levyns, 1945).

The lateral axis surrounded by this perigynium may

give rise to a second-order paracladium which then

produces bisexual spikelets or unisexual reduced

spikelets similar to those in Carex s.s., or it may give

rise directly to bisexual spikelets or reduced uni-

sexual spikelets (Levyns, 1945) (Fig. 2J). Male ﬂowers

are generally produced when the inﬂorescence axis (of

whatever order) stops branching. Thus, male ﬂowers

can be found on the distal tips of the main ﬂorescence,

the paracladia, the spikes and the bisexual spikelets

(of Kobresia and Schoenoxiphium only) (Fig. 2E–L),

unless the axis is truncated. Both male and female

ﬂowers are thus produced at different levels of the

inﬂorescence branching hierarchy, the females

directly connected to branching events and the males

most often produced at the tips of axes that no longer

branch. We return to these fundamental differences

in inﬂorescence structure in our argument for

merging all species of Cariceae into Carex.

OVERVIEW OF PREVIOUS CLASSIFICATIONS

Classiﬁcations of Carex and its closely allied genera

in Cariceae are numerous and conﬂicting, most being

based on particular ideas of natural or evolutionary

relationships. Robertson (1979) discussed this classi-

ﬁcation history from pre-Linnean times until the mid-

20th century, and Kern (1958) and Zhang (2001)

reviewed the early classiﬁcation history of Schoenox-

iphium and Kobresia, respectively. Reznicek (1990)

and Egorova (1999) provided good summaries of post-

Linnean classiﬁcations until the end of the 20th

century, and Starr et al. (2004) summarized classiﬁ-

cation issues related to the segregate genera. Inﬂo-

rescence morphology has played a key role in most

classiﬁcations from Linnaeus’ ﬁrst division of Carex

s.s. into ﬁve groups, deﬁned by the number of spikes

and the arrangement of staminate and pistillate

MAKING CAREX MONOPHYLETIC 9

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ﬂowers in the inﬂorescence. Early attempts to segre-

gate smaller genera from Carex (Raﬁnesque-

Schmaltz, 1840; Heuffel, 1844) were poorly justiﬁed

and not widely accepted. However, much effort was

expended during the 19th and early 20th centuries to

organize species in this large genus into natural

groups as subgenera, sections and series (e.g.

Tuckerman, 1843; Drejer, 1844; Bailey, 1886; Holm,

1903; Kükenthal, 1909; reviewed by Robertson, 1979).

We provide here an overview of previous 20th century

classiﬁcations and associated theories of relation-

ships, using the evolutionary terminology of the

authors to give the ﬂavour of the pre-cladistic think-

ing on which most of these classiﬁcations were based.

Subgeneric classiﬁcation of Carex s.s.

Although Kükenthal’s (1909) division of Carex s.s.

into four subgenera based on inﬂorescence structure

is that most often followed, with some modiﬁcations,

in subsequent ﬂoristic manuals, fewer subgenera

have been recognized by others (Ohwi, 1936; Koyama,

1962). Many rejected Psyllophora as a distinct subge-

nus, dispersing these species among subgenera Carex

and Vignea, or even into Uncinia,Kobresia or Schoe-

noxiphium (see below for details). The recognition of

subgenus Vignea has been almost universal, the most

notable change being the merger of a few dioecious,

unispicate species from subgenus Psyllophora into it

(e.g. section Physoglochin Dumort.). Carex subgenera

Carex and Vigneastra have been treated as a single

subgenus by some (Ohwi, 1936; Koyama, 1962; Kern

& Noteboom, 1979). There have also been suggestions

that subgenus Vigneastra is ancestral to one or more

of the other Carex subgenera (Nelmes, 1951, 1952,

1955; Hamlin, 1959; Nannfeldt, 1977), and possibly

derived from Schoenoxiphium (Haines & Lye, 1972) or

from a ‘primitive Kobresia–Schoenoxiphium stock’

(Smith & Faulkner, 1976) based on the similarity of

inﬂorescence structure in these groups.

New subgenera in Carex s.s. have been proposed by

various 20th century botanists. The monotypic sub-

genus Altericarex H.St.John & C.S.Parker was

described to accommodate Carex concinnoides,a

North American species with four stigmas and

tetragonous nutlets (St. John & Parker, 1925). Sub-

genus Kuekenthalia Savile & Calder was proposed for

those species with more or less inﬂated perigynia and

often persistent styles [sections such as Lupulinae

Tuck. ex J.Carey, Paludosae G.Don, Vesicariae

(Heuff.) J.Carey etc., plus a few unispicate species]

(Savile & Calder, 1953). Another subgenus of Carex,

Kreczetoviczia T.V.Egorova, was segregated from sub-

genus Carex to accommodate species with two

stigmas, but mostly unisexual spikes [e.g. sections

Phacocystis Dumort. s.l.,Graciles (Tuck. ex Kük.)

Ohwi and Abditispicae G.A.Wheeler] (Egorova, 1985).

None of these subgenera has, however, been widely

adopted.

Relationships of allied genera to Carex s.s. and to

each other

Kobresia,Uncinia and Schoenoxiphium, named in the

early 19th century, have always been considered to be

closely related to Carex s.s. and evolutionary sce-

narios forming the basis for classiﬁcations have

included them. Kükenthal’s (1909) classiﬁcation was

based on the idea that Schoenoxiphium and Kobresia

are the base of a reduction series in which the spike-

let rachilla is gradually reduced from an elongated

structure bearing distal male ﬂowers (Schoenox-

iphium, some Kobresia) to a sterile rachilla (some

Kobresia and Schoenoxiphium,Uncinia and a few

unispicate Carex) and then to a vestigial rachilla

(Carex) in an enclosing prophyll (perigynium), the

margins of which change from open in many Kobresia

to sealed, except for a small terminal opening with

only the style, stigmas and rachilla (if present) pro-

truding from that oriﬁce (e.g. Uncinia,Carex,Cymo-

phyllus). Perigynia of Schoenoxiphium vary from

having a rather broad opening to being almost com-

pletely sealed as in most of the other genera.

Many species in the allied genera are unispicate

and androgynous, leading to an early theory that

unispicate Carex spp. are primitive and multispicate

Carex spp. are derived from them (Drejer, 1844). In

line with this idea, Kükenthal (1909) named subge-

nus Primocarex Kük., circumscribing it to include all

unispicate Carex, including C. fraseriana Ker Gawl.,

which was subsequently segregated as the monotypic

genus Cymophyllus by Mackenzie (1913) based on its

unique leaf morphology. Strong opposition to the idea

that unispicate Carex spp. were primitive came from

those who thought that at least some unispicate inﬂo-

rescences were derived from multispicate ones by

reduction. They proposed systems that placed unispi-

cate Carex spp. variously in subgenus Carex or sub-

genus Vignea rather than recognizing a distinct

subgenus Psyllophora (Kreczetovicz, 1936; Ohwi,

1936; Nelmes, 1952; Koyama, 1962; Smith &

Faulkner, 1976). Nelmes (1952) discussed the poly-

phyly of subgenus Psyllophora, speculating that

nearly half of the species were ‘true Carices’ that

could be accommodated in other subgenera of Carex

s.s., whereas at least half of those remaining were

probably derived from Carex s.s. and the remainder

from Uncinia,Kobresia or Schoenoxiphium. Another

view of subgenus Psyllophora was based on associa-

tions with smut fungi (Anthracoidea) and strongly

inﬂuenced by the presence (or not) of the rachilla and

by Heilborn’s chromosome data (Heilborn, 1924;

Savile & Calder, 1953). Savile & Calder (1953)

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considered subgenus Psyllophora to include rachilla-

bearing unispicate species only, referring to this

group as ‘true Primocarex’, and moved the remaining

unispicate species lacking a rachilla either to subge-

nus Vignea or to their new subgenus Kuekenthalia.

They considered this more narrowly circumscribed

subgenus Psyllophora to be derived from Kobresia

and ancestral to subgenera Carex,Vignea and Kuke-

nthalia. Kukkonen supported the basic ideas of Savile

& Calder (1953), but proposed a new phylogenetic

hypothesis in which subgenus Vignea was derived

from Kobresia through subgenus Psyllophora, and

subgenus Carex was independently derived from Kob-

resia through Carex section Acrocystis Dumort. in

subgenus Carex (Kukkonen, 1963).

Hamlin (1959) proposed a putative caricoid ances-

tor with a large branching inﬂorescence having clado-

prophylls and bearing spikelets that each had a basal

female ﬂower and a persistent rachilla bearing male

ﬂowers, the whole spikelet partially enclosed by a

prophyll (perigynium). He postulated that this ances-

tor gave rise to two evolutionary lines, one retaining

the rachilla, but reducing the inﬂorescence branching,

and the other losing the rachilla, but retaining the

compound inﬂorescence. The line retaining the

rachilla then split to give rise to Schoenoxiphium,

Kobresia,Uncinia and some species in Carex subge-

nus Psyllophora. The line retaining the highly

branched inﬂorescence, but losing the rachilla, gave

rise to Carex subgenus Vigneastra from which sub-

genera Vignea and Carex were derived along separate

lines. He hypothesized that Carex subgenus Psyl-

lophora was polyphyletic, comprising species with

unispicate inﬂorescences that arose in parallel by

reduction from each of the six lineages in his evolu-

tionary scenario. Hamlin’s solution to the classiﬁca-

tion problem that this scheme created was the

division of Carex subgenus Psyllophora into several

genera to accommodate those with different ancestry,

rather than uniting the tribe into a single genus

(Hamlin, 1959).

As noted by Kern (1958), discriminating among the

genera in Cariceae was not difficult in the early 19th

century when only a few species of each genus were

described, but, as more species were discovered, the

lines between them became blurred. The broader

ciliate rachillae of Schoenoxiphium have been consid-

ered as a distinctive character contrasting with the

less conspicuous, usually terete, rachillae of Kobresia

(Clarke, 1883; Kükenthal, 1909), but intermediates

(Kükenthal, 1940) and exceptions were discovered as

many more Kobresia spp. were described from Asia.

Nelmes (1952) pointed out that the same reduction

series in inﬂorescence morphology that Kükenthal

postulated for the evolutionary pathway from Schoe-

noxiphium to Kobresia to Uncinia to Carex s.s. could

also be observed within Schoenoxiphium and within

Kobresia, an observation further ampliﬁed by others

(Koyama, 1961; Haines & Lye, 1972, 1983; Smith &

Faulkner, 1976). Both genera vary in the extent of

closure of their perigynia and in the extent of lateral

branching in the inﬂorescence, resulting in overlap of

traits between them. Ivanova (1939) transferred Kob-

resia spp. with paniculiform inﬂorescences to Schoe-

noxiphium and most of the currently recognized

species of Schoenoxiphium into Archaeocarex Börner,

but this genus has never been accepted. Many previ-

ous authors have argued that Kobresia and Schoenox-

iphium could not be reliably distinguished on the

basis of morphology (Nelmes, 1952; Kern, 1958;

Smith & Faulkner, 1976). Koyama (1961) merged the

two genera and made the necessary nomenclatural

transfers from Schoenoxiphium to Kobresia. However,

most 20th century authors maintained the traditional

segregation of the two genera, in part because of their

different geographical ranges and ecological prefer-

ences (Ivanova, 1939; Kukkonen, 1978, 1983; Haines

& Lye, 1983; Rajbhandari & Ohba, 1991; Noltie, 1993;

Zhang, 2001). Segregate genera Elyna Schrad. and

Hemicarex Benth., deﬁned on the basis of their spici-

form inﬂorescences and bisexual and unisexual spike-

lets, respectively, and Blysmocarex N.A.Ivanova,

characterized by distigmatic female ﬂowers, are no

longer recognized, but retained as subgenera of Kob-

resia in most classiﬁcations of that genus (Zhang,

2001). The similarity of Uncinia to Carex s.s. has also

been noted (reviewed by Starr et al., 2008), but only

Koyama (1961) proposed the merging of Uncinia into

Carex s.s., although without making the necessary

nomenclatural transfers. Koyama (1961) recognized

only two genera, Kobresia and Carex s.s., in Cariceae,

the latter with only two subgenera, Carex and Vignea.

Although several authors have commented on the

difficulty of clearly deﬁning genera in Cariceae, only

Mora Osejo (1966) proposed the inclusion of Carex

s.s.,Kobresia,Schoenoxophium and Uncinia in one

genus, recognizing each of them at the subgeneric

level in genus Carex s.l., but without making valid

nomenclatural transfers.

Lack of support from molecular phylogenetics for

previous classiﬁcations

Previously proposed evolutionary scenarios for

Cariceae and classiﬁcations based on them are not

supported by molecular studies (Yen & Olmstead,

2000; Roalson et al., 2001; Starr et al., 2004, 2008;

Waterway & Starr, 2007; Starr & Ford, 2009;

Waterway et al., 2009; Gehrke et al., 2010; Jung &

Choi, 2013) (Fig. 1). Schoenoxiphium and Kobresia

are nested in a clade of species of Carex s.s., rather

than being sister to Carex s.s. as proposed earlier

(Kükenthal, 1909). The Schoenoxiphium lineage is

MAKING CAREX MONOPHYLETIC 11

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distinct from that of Kobresia and includes two small

clades of mostly unispicate Carex s.s. (Gehrke et al.,

2010). Kobresia has been poorly sampled in all pub-

lished papers to date, but it appears to be polyphyletic

in the caricoid clade, a conclusion supported by more

extensive sampling in recent work by Zhang et al.

(2013). These Kobresia lineages are more closely

related to various unispicate Carex s.s. than to Schoe-

noxiphium.Uncinia is ﬁrmly nested in the caricoid

clade and is clearly not the progenitor of other groups

as suggested by Nelmes (1952). The monotypic genus

Cymophyllus, endemic to the south-eastern USA,

is also nested in the caricoid clade and, despite

its unusual leaves, does not warrant the generic

status accorded it in recent North American ﬂoras

(Mackenzie, 1931–1935; Reznicek, 2002). Similarly,

the monotypic genus Vesicarex has been shown to be

related to species of Carex section Abditispicae (core

Carex clade) and treated as Carex collumanthus

(Steyerm.) L.E.Mora (Mora Osejo, 1982; Wheeler,

1989), a conclusion supported by molecular data

(Starr et al., 2004; Waterway & Starr, 2007). Carex

section Siderostictae is not closely related to other

broad-leaved species of Carex subgenus Carex, but

forms a distinct lineage (Waterway et al., 2009).

Recent work on Carex subgenus Vigneastra indicates

that it is polyphyletic; species from sections Hemis-

caposae and Surculosae belong to the same lineage as

the early-diverging Siderostictae clade (Yano et al.,

2014) and two recently sampled species of section

Hypolytroides Nelmes (subgenus Vigneastra) are

sister to this expanded Siderostictae clade (Starr

et al., 2015). The early-diverging position of the

broad-leaved species in the Siderostictae clade had

earlier been suggested by Raymond (1959) for sec-

tions Hemiscaposae and Surculosae and by Egorova

(1999), who considered section Siderostictae, with sec-

tions Decorae and Curvulae, as the least evolutionar-

ily advanced groups in subgenus Carex. The most

recent studies that include subgenus Vigneastra show

that representatives from other sections of subgenus

Vigneastra form one (Gehrke & Linder, 2009) or more

(Waterway et al., 2009; Starr et al., 2015) early-

diverging lineages in the core Carex clade. None of

these groups of subgenus Vigneastra appears to be

closely related to Schoenoxiphium, as Haines & Lye

(1972, 1983) had suggested.

ARGUMENT FOR ONE GENUS

Published molecular phylogenetic hypotheses for

Cariceae are quite consistent, despite differences in

DNA regions used, taxon density and analytical

methods (see Fig. 1 for a summary diagram of rela-

tionships). All recent analyses agree that the expanded

Siderostictae clade or the Siderostictae + Hypol-

ytroides clade, recently termed the ‘minor Carex alli-

ance’ by Starr et al. (2015), is monophyletic and sister

to the rest of the tribe, which is also strongly supported

as monophyletic (Waterway et al., 2009; Jung & Choi,

2013; Yano et al., 2014), that the Vignea clade is

monophyletic (Ford et al., 2006, 2012; Waterway &

Starr, 2007; Starr & Ford, 2009; Waterway et al., 2009;

Jung & Choi, 2013), and that the large core Carex clade

is monophyletic only with the inclusion of at least part

of subgenus Vigneastra with nearly all species of

subgenus Carex (Waterway & Starr, 2007; Gehrke &

Linder, 2009; Starr & Ford, 2009; Waterway et al.,

2009; Starr et al., 2015). Some analyses show a mono-

phyletic caricoid clade, but support for this clade is

weak (Starr et al., 2004, 2008; Gehrke & Linder, 2009)

to moderate (Waterway & Starr, 2007; Starr & Ford,

2009; Waterway et al., 2009). Except for the position of

the Siderostictae clade as sister to the rest of Cariceae,

the molecular data have not yet resolved the relation-

ship among the major clades (for a summary, see Starr

& Ford, 2009). No analysis has contradicted the mono-

phyly of the tribe as a whole.

Given the strong and consistent molecular evidence

for monophyly of Cariceae and the frequent nesting of

the allied genera in one of the major clades of Carex

s.s., it is clear that a new classiﬁcation is needed to

reﬂect the evolutionary relationships in this group

more accurately. With the goal of monophyletic

genera, only three reasonable possibilities exist in the

Linnean system: (1) recognizing each of the four or

ﬁve major clades in the molecular phylogenetic trees

as a distinct genus; (2) recognizing the three strongly

supported clades as distinct genera and naming each

of the dozen or more lineages in the caricoid clade as

a distinct genus; or (3) broadly circumscribing Carex

s.l. to include all species in tribe Cariceae. We discuss

these options in turn below.

Option 1: Major clades as separate genera

As described above, support for three of the clades

(Siderostictae, core Carex and Vignea) is strong in all

recent studies, but there is considerable morphologi-

cal variation in the ﬁrst two of these clades, making

it difficult to ﬁnd consistent morphological traits by

which to diagnose them. Species in the Siderostictae

clade, as expanded by Yano et al. (2014) to include

Carex sections Hemiscaposae and Surculosae, have

similar growth forms and share several other fea-

tures. Vegetative and reproductive culms arise sepa-

rately on short rhizomes and most species have broad

leaves near the base of vegetative culms and blade-

less sheaths at the base of lateral reproductive culms.

Each female ﬂower is enclosed by a perigynium from

which the triﬁd style emerges, and both male ﬂowers

and perigynia are subtended by scale-like glumes. All

species in this clade have low chromosome numbers

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(n= 6 or 12), and some species exhibit euploidy, in

contrast with the aneuploid series that characterizes

Carex (Escudero et al., 2012; Chung, Yang & Lee,

2013; Yano et al., 2014). However, inﬂorescence struc-

ture varies considerably in the expanded Siderostictae

clade. Section Siderostictae is characterized by termi-

nal staminate or androgynous spikes and lateral

androgynous spikes; that is, the main ﬂorescence

terminates in a spike of male ﬂowers, in some cases

with proximal perigynia, and co-ﬂorescences arising

from the culm nodes are usually androgynous with

proximal perigynia and distal male ﬂowers (Chung

et al., 2013). Each paracladium comprises a subtend-

ing sheathing bract varying from spathe-like to folia-

ceous, a cladoprophyll and an androgynous spike.

Nodes are sometimes binate, bearing two pedunculate

spikes rather than the usual single spike. Lateral

spikes have a few spirally arranged perigynia near

the base and a spiral of male ﬂowers produced dis-

tally, sometimes with an elongated internode between

the two types of ﬂower. The terminal spike is simply

the tip of the main culm axis that bears spirally

arranged male ﬂowers, often with a single perigynium

at the base. In contrast, species of sections Hemisca-

posae and Surculosae have paniculiform inﬂores-

cences with higher order branching. Each node of the

reproductive culm in section Hemiscaposae bears one

or two compound inﬂorescences having second- or

third-order branching, subtended by spathe-like

bracts with short blades. Lateral axes form bisexual

androgynous spikes, each subtended by a cladopro-

phyll that resembles a perigynium, but does not bear

a female ﬂower. Species in section Surculosae also

have androgynous lateral branches arranged in com-

pound inﬂorescences, single or binate from nodes,

subtended by smaller, more scale-like bracts.

Raymond (1959) reported that some of the cladopro-

phylls in C. surculosa Raymond (= C. tsiangii

F.T.Wang & Tang) bear fertile female ﬂowers, remi-

niscent of Schoenoxiphium. Despite this high level of

variation, the Siderostictae clade is strongly sup-

ported as monophyletic with molecular data and has

a restricted geographical distribution, being found

only in eastern and south-eastern Asia in temperate

to subtropical forests.

This variation in inﬂorescence structure, together

with variation in leaf morphology from narrow

cauline leaves (e.g. C. tumidula Ohwi, C. grandiligu-

lata Kük.) to basal clusters of broad leaves (one to

several centimetres wide), makes it difficult to con-

sistently deﬁne the Siderostictae clade. In addition,

the features of the Siderostictae clade are not limited

to that clade. Binate inﬂorescences are also found in

the core Carex clade and in Schoenoxiphium, and

compound inﬂorescences composed of androgynous

lateral axes can also be found in all three other major

clades. Broad leaves are also found in shade-tolerant

species of the core Carex clade. In short, characters

that might be considered synapomorphies for the

Siderostictae clade all exhibit extensive homoplasy in

the larger Cariceae clade.

A similar problem exists for the core Carex clade

because of the variability introduced by the inclusion

of species from subgenus Vigneastra in phylogenetic

trees based on molecular data (Gehrke & Linder,

2009; Waterway et al., 2009). Core Carex is by far the

largest clade of Cariceae (c. 1400 species) and

includes most species currently classiﬁed in Carex

subgenus Carex and probably at least half of those in

Carex subgenus Vigneastra, although relatively few

have yet been included in DNA-based phylogenetic

studies. Inﬂorescence structure in subgenus Carex is

similar in the majority of species. Most have terminal

staminate spikes (the main ﬂorescence) and mostly

pedunculate pistillate lateral spikes, each of which is

subtended by a more or less leafy bract and enclosed

at the base of the lateral axis with a tubular clado-

prophyll (the ﬁrst-order paracladia; Fig. 2I). Varia-

tions on this theme occur in a few lineages that are

characterized by having additional distal staminate

spikes or androgynous lateral spikes, especially

towards the apex of the inﬂorescence. In a few cases,

the terminal spikes are gynecandrous rather than

staminate [e.g. some species in sections Racemosae

G.Don, Porocystis Dumort., Hymenochlaenae (Drejer)

L.H.Bailey], and a few species are unispicate and

dioecious [e.g. sections Pictae Kük. and Scirpinae

(Tuck.) Kük.]. Members of subgenus Vigneastra are

characterized by pedunculate, bisexual spikes that

are often much branched and have cladoprophylls

that are relatively large and resemble perigynia in

shape. Sequenced species of Carex subgenus Vigneas-

tra appear to be part of at least two lineages in the

core Carex clade (Starr et al., 1999, 2004, 2008, 2015;

Waterway & Starr, 2007; Gehrke & Linder, 2009;

Waterway et al., 2009). As a result, the core Carex

clade has an even larger range of variation in inﬂo-

rescence structure than the Siderostictae clade,

varying from dioecious unispicate species to species

with unisexual or androgynous lateral spikes, some-

times binate at proximal nodes, with one or more

distal staminate spikes, to species with higher order

branching culminating in androgynous lateral axes

and distal staminate spikes on the main axis. Most

species in this clade have three stigmas as in the

Siderostictae clade, but there are three sections with

distigmatic ﬂowers [Phacocystis s.l.,Bicolores (Tuck.

ex L.H.Bailey) Rouy, Abditispicae] and occasional

reductions to two stigmas in other sections (e.g.

C.saxatilis L. in section Vesicariae). Cladoprophylls

vary from tubular to perigynium-like, a few even

bearing a pistillate ﬂower (Nelmes, 1951), as in some

MAKING CAREX MONOPHYLETIC 13

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Schoenoxiphium spp. In short, although most species

follow a fairly simple inﬂorescence plan with ﬁrst-

order lateral branching only, the variability in inﬂo-

rescence structure and the fact that similar variants

are found in other clades suggest homoplasy in any

characters used to deﬁne the core Carex clade.

The Vignea clade is easier to deﬁne, most species

having bisexual terminal spikes, sessile, bisexual

lateral spikes that generally lack cladoprophylls and

female ﬂowers with only two stigmas. Most sections

have androgynous spikes, but a few of the larger

lineages (e.g. sections Ovales Kunth, Glareosae

D.Don, Stellulatae Kunth) have gynecandrous spikes

(Ford et al., 2006; Hipp et al., 2006), and some species

have mesogynous or mesandrous spikes or alternate

staminate and pistillate ﬂowers in the spikes. Excep-

tions to this pattern are relatively few. For example,

a few species are unispicate, some are dioecious (e.g.

section Physoglochin) and a few lineages with con-

densed inﬂorescences have higher order branching

(Ford et al., 2006, 2012; Hipp et al., 2006, 2013;

Waterway & Starr, 2007; Jung & Choi, 2013). Three

stigmas are found in only a few early-diverging

species in the group, including C. gibba Wahlenb.,

which is sister to the rest of the Vignea clade in

molecular analyses (Ford et al., 2006; Waterway et al.,

2009; Jung & Choi, 2013). Although support reported

for the monophyly of subgenus Vignea is strong, the

topology of the tree in the Vignea clade is quite

variable, depending on the genes used, taxon sam-

pling and analytical methods.

Given the inclusion of all four segregate genera of

Cariceae, the caricoid clade is almost impossible to

deﬁne with consistent morphological synapomorphies,

a situation that has been discussed at length by Starr

and co-workers (Starr et al., 2004; Starr & Ford,

2009). Inﬂorescence architecture and vegetative

structure vary widely in the group, perigynia can be

open or closed, and stigma number varies from two to

three. Furthermore, the level of support for the cari-

coid clade is never strong and depends on taxon

density, DNA regions and analytical methods used

(Starr & Ford, 2009; Gehrke et al., 2010). We are thus

hesitant to consider this clade as the basis for recog-

nizing a fourth genus in parallel with the other three

clades. Conferring generic status on this clade would

mean including in it species from all four of Küken-

thal’s subgenera of Carex and species from all four

other genera of Cariceae; this would require at least

as many name changes as uniting the whole tribe into

the genus Carex s.l. With so much variation in form,

it would not be a practical solution to the problem of

paraphyly in Carex s.s., especially because we are not

sure that additional gene and taxon sampling will

continue to support the recognition of the caricoid

clade. Another related possibility might be to recog-

nize each of the two clades that make up the caricoid

clade, especially because they do not together form a

monophyletic group in some analyses (Gehrke et al.,

2010; Jung & Choi, 2013). This would not solve all

problems, however, because the core unispicate clade

also has fairly weak support and is highly variable,

whereas the Schoenoxiphium clade includes several

species of Carex s.s. that do not strongly resemble

Schoenoxiphium (see below for more detail). We thus

reject the ﬁrst option of recognizing each of the major

clades as a distinct genus because that would create

more problems than it solves.

Option 2: Many genera

The second option is to recognize each strongly sup-

ported clade (Siderostictae, core Carex and Vignea)as

a genus, and to divide the caricoid clade into smaller

monophyletic groups that could be recognized as

genera. There is some appeal in this alternative

approach, because a few groups in the caricoid clade

appear to be both monophyletic and distinctive.

Uncinia and Schoenoxiphium have been sampled

quite extensively, although with relatively few genes,

and are monophyletic in the best sampled molecular

studies (Starr et al., 2003, 2004, 2008; Gehrke et al.,

2010; Luceño et al., 2013). An elongate rachilla with a

hooked tip is shared by all Uncinia spp., although the

hook in the controversial U. kingii Boott [= C. kingii

(Boott) Reznicek] has a different derivation (Starr

et al., 2004). However, U. kingii is sister to a clade

including all other sampled Uncinia spp., meaning

that the entire group is also monophyletic, although

its relationships to other species in the caricoid clade

are not clear (Starr et al., 2008). Uncinia spp. also

share a perigynium that is closed, except at the oriﬁce

where the style and rachilla emerge, and all have

three stigmas. Except for the hooked tip of the

rachilla, these features are not unique to Uncinia, but

together they make the genus easy to recognize and

describe. Uncinia also has a coherent and limited

Gondwanan distribution, most species occurring in

New Zealand, Australia and southern South America,

ranging north to Hawaii and the Philippines in the

Eastern Hemisphere and to the Caribbean region in

the Western Hemisphere. The main problem with

recognizing it as a distinct genus is that it is nested

in the caricoid clade with several species of unispicate

Carex s.s. and Kobresia.

Schoenoxiphium also forms a well-supported mono-

phyletic group in the most recent analysis, in which

85% of the named species were sampled (Gehrke

et al., 2010), but Schoenoxiphium is more difficult to

distinguish from other genera of Cariceae, especially

Kobresia. The genus was originally deﬁned by the

ﬂattened ciliate rachilla that often bears male ﬂowers

distally and emerges from the oriﬁce of a closed

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perigynium. These features are in contrast with the

shorter, terete rachillae of many Kobresia spp. in

which the male ﬂowers are fewer in number or sup-

pressed completely and the perigynium is often

open on one side nearly to the base (Fig. 2E). Inﬂo-

rescence nodes in Schoenoxiphium tend to be more

evenly spread along the ﬂowering culm than in Kob-

resia, and inﬂorescence bracts tend to be leaf-like and

often sheathing in some Schoenoxiphium spp., but

much reduced in Kobresia (Reznicek, 1990). Most

Schoenoxiphium spp. also have more highly branched

inﬂorescences than those of most Kobresia spp. Mor-

phological variability in Schoenoxiphium appears to

be almost as great within some species as across the

genus (Levyns, 1945; Haines & Lye, 1983). Some

individuals of S. lehmannii (Nees) Steud. are almost

Carex-like, having perigynia closed except at the

oriﬁce and arranged in lateral androgynous spikes,

as well as a terminal staminate spike, each spike

except the terminal subtended by a foliaceous bract

(Haines & Lye, 1983). Even the rachilla in these

individuals is sterile and short. Other individuals of

the same species have a fully developed rachilla that

emerges from the perigynium and bears a set of male

ﬂowers distally (Haines & Lye, 1983). Branching can

be complex in Schoenoxiphium involving third- or

higher order branching from the main axes and

spikes of spikelets bearing perigynia from within

female or bisexual spikelets on the ultimate ﬂowering

axes (Gehrke et al., 2012). Pistillate ﬂowers borne in

the axil of a branch on which spikelets are borne

have an enclosing, but more open, cladoprophyll

that resembles the unsealed perigynia of Kobresia

(Fig. 2G).

The molecular evidence suggests that two small

clades of species of Carex s.s. with reduced inﬂores-

cence, named the C. andina and C. distachya clades,

and together including representatives from three of

Kükenthal’s subgenera, are more closely related to

Schoenoxiphium than are any Kobresia spp. (Gehrke

et al., 2010; Fig. 1). Schoenoxiphium plus the

C. andina and C. distachya clades form a monophyl-

etic group, but differ strongly in morphology and

distribution. Schoenoxiphium is endemic to eastern

and southern Africa, the C. andina clade is endemic

to southern South America and Australasia, and

species from Europe, the Mediterranean Region,

Macaronesia and central Africa form the C. distachya

clade. Most species in the C. andina clade and some

in the C. distachya clade have at least some unispi-

cate individuals, whereas most Schoenoxiphium spp.

have at least some individuals with higher order

branching in the inﬂorescence. If the option of break-

ing up the caricoid clade into smaller genera, each

with more consistent morphological features, were to

be followed, three genera could be recognized from the

Schoenoxiphium clade, one for Schoenoxiphium itself

and one for each of the sister clades.

The rest of the caricoid clade would be more diffi-

cult to segregate into monophyletic genera (Fig. 1). A

major problem is the genus Kobresia, which is poly-

phyletic in all molecular analyses so far (Yen &

Olmstead, 2000; Starr et al., 2004; Gehrke & Linder,

2009; Gehrke et al., 2010; Jung & Choi, 2013). Some

lineages include both Kobresia spp. and unispicate

Carex spp. However, only a small proportion of Kob-

resia spp. have been included in molecular studies

until now. In a much more comprehensive study of

Kobresia, Zhang et al. (2013) found ﬁve distinct line-

ages variously nested in the caricoid clade, but

without good correspondence to the previous subge-

neric categories of Kobresia. If we continue the logic of

naming monophyletic groups in the caricoid clade as

distinct genera, we would have to recognize at least

these ﬁve lineages that include Kobresia spp. and at

least seven lineages of unispicate Carex. Many of

these small groups already have names at the generic

level as these highly reduced unispicate species were

seen as unusual by many 19th century botanists. One

problem with doing this is that, despite considerable

study of this clade, few genes have been used, and the

tree topology in the caricoid clade is not consistent

across studies using different genes (Starr et al.,

2004; Waterway & Starr, 2007; Gehrke & Linder,

2009; Gehrke et al., 2010). Taking the approach of

recognizing numerous genera in the caricoid clade

would not serve the goal of long-term nomenclatural

stability and, given the superﬁcial similarity of many

unispicate species in this group, would cause consid-

erable confusion.

Option 3: One genus

At this point, it should be clear that there are prob-

lems with recognizing each of the major clades as

distinct genera and even more difficulties with recog-

nizing three of the major clades and at least a dozen

lineages in the caricoid clade as distinct genera.

Neither of these ﬁrst two options is optimal to meet

the goals of nomenclatural stability, generic mono-

phyly and ease of use. Instead, we propose here to

take the third choice listed above, and merge all

genera currently treated as tribe Cariceae (Cymophyl-

lus,Kobresia,Schoenoxiphium,Uncinia) into the

genus Carex. Analysis of molecular data has consist-

ently shown this group of genera to be strongly sup-

ported as monophyletic in comprehensive studies of

Cyperaceae (Muasya et al., 1998, 2009; Simpson

et al., 2007; Jung & Choi, 2010), and in more detailed

analyses of the tribe in relation to outgroups sug-

gested by family-level analyses (Starr et al., 2004;

Waterway & Starr, 2007; Gehrke & Linder, 2009;

Starr & Ford, 2009; Waterway et al., 2009; Gehrke

MAKING CAREX MONOPHYLETIC 15

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et al., 2010; Jung & Choi, 2013; Léveillé-Bourret

et al., 2014). In contrast with the difficulty in ﬁnding

consistent morphological synapomorphies for the

major clades, as detailed above, it is much easier to

list clear synapomorphies for Carex s.l. in the broad-

ened circumscription proposed here. The combination

of unisexual ﬂowers and perigynia surrounding the

female ﬂowers is unique in Cyperaceae. As described

earlier, what appear to be pistillate ﬂowers in

Cariceae are actually reduced spikelets in which the

proximal female ﬂower is surrounded by the enclosing

prophyll (perigynium) and the spikelet axis is either

vestigial or elongated, with or without distal stami-

nate ﬂowers. Processes at play in this change from a

multi-ﬂowered spikelet with a spiral of bisexual

ﬂowers having a bristle perianth, as in the closely

related tribe Scirpeae, involve suppression of peri-

anth, modiﬁcation of sex expression resulting in uni-

sexual ﬂowers and expansion and at least partial

fusion of the prophyll into a perigynium. It is easy to

envisage selection against maintenance of the rachilla

and its distal male ﬂowers within an enclosure like

the perigynium, especially because it is possible for

any axis in the synﬂorescence to produce the needed

staminate ﬂowers directly from the same type of

primordia that produce the reduced female spikelets

(Vrijdaghs et al., 2010).

Classiﬁcations have been based largely on features

of inﬂorescences and perigynia, in part because they

are more obvious than ﬂoral differences within the

perigynia, and in part because vegetative features in

Carex s.l. have more variation within segregate genera

than among them. Other than the unusually thick

leaves of C. fraseriana, which lack a midrib, and the

broad, pseudo-petiolate leaves of some South-East

Asian species, no distinctive vegetative features set

any group apart from the others. As we have described

above, inﬂorescence structure also varies substantially

in clades and even within species. Furthermore,

mutant individuals in the core Carex clade sometimes

exhibit inﬂorescence and ﬂoral forms of the allied

genera. Occasional teratological specimens with

rachillae bearing male ﬂowers in species that normally

have only a tiny vestigial rachilla have been found

naturally and can be induced experimentally by appli-

cation of growth regulators or by damaging the root

tips (Smith & Faulkner, 1976). Section Hangzhouenses

C.Z.Zheng, X.F.Jin & B.Y.Ding was described in Carex

subgenus Vigneastra to accommodate an unusual

specimen of Carex from eastern China that had per-

igynia with protruding, elongated rachillae bearing

staminate ﬂowers (Jin et al., 2005). Comparison of

DNA sequences with species growing on the same cliff,

however, revealed that these specimens were more

likely to be aberrant specimens of Carex simulans

C.B.Clarke of section Rhomboidales Kük., subgenus

Carex (X. F. Jin and M. J. Waterway, unpubl. data)

than a new species in a new section of subgenus

Vigneastra. These results suggest that, although sup-

pression may appear to be genetically ﬁxed in particu-

lar clades, it can be reversed under certain conditions.

Various teratological specimens have been illustrated

showing similar Schoenoxiphium-like spikelets in

specimens of Carex crinita Lam., C. albicans Willd. ex

Spreng. var. emmonsii (Dewey ex Torr.) Rettig,

C. pallescens L., C. sprengelii Steud. and C. sitchensis

Bong. (Penzig, 1894; Holm, 1896; Clarke, 1909;

Svenson, 1972). Perigynia enclosing stamens rather

than a pistil in C. acuta L. and pistils subtended by

glumes rather than enclosed in perigynia have also

been noted (Holm, 1896; Smith & Faulkner, 1976;

Timonen, 1998). Suppression of lateral branching

resulting in unispicate individuals of normally multi-

spicate Carex spp. (e.g. C. ﬂacca Schreb.) has also been

found sporadically in natural Carex populations and

can be easily induced by the application of 2,3,5-

triodobenzoic acid (TIBA) (Smith & Faulkner, 1976).

Reports of these aberrant individuals or populations

frequently mention trampling or other disturbance,

suggesting that damage to meristems or environmen-

tal factors may be involved (Svenson, 1972; Smith &

Faulkner, 1976).

Bisexual ﬂowers are another aberration that have

been reported infrequently in Kobresia and Schoenox-

iphium (Timonen, 1998), most recently as sporadic

occurrences in populations of S. lehmannii and

S. burkei C.B.Clarke (Gehrke et al., 2012). Detailed

observations on several individuals revealed a full

transition series from proximal female spikelets to

bisexual spikelets having distal male ﬂowers on the

rachilla to bisexual ﬂowers directly on the lateral axis

and to distal male ﬂowers at the apex of the lateral

axis. These bisexual ﬂowers were produced directly on

the lateral axis in the transition zone between multi-

ﬂowered bisexual spikelets and distal male ﬂowers.

The bisexual ﬂowers were similar to male ﬂowers in

being subtended by a scale-like glume, but they

lacked a perigynium. In addition to the normal whorl

of three stamens found in male ﬂowers, these

bisexual ﬂowers had a biconvex distigmatic ovary at

the centre, in contrast with the tristigmatic ovary

found in female ﬂowers (Gehrke et al., 2012). These

observations provide further support to the idea that

the position on the axis is important in the control of

sex expression and that primordia on the spikelet can

be regulated, either internally or externally, to

produce either spikelets or single ﬂowers (Vrijdaghs

et al., 2009, 2010). The extreme variability in inﬂo-

rescence structure not only among Schoenoxiphium

spp., but also within species and even within different

parts of the same inﬂorescence, was already noted by

Levyns (1945), whose invaluable observations on

16 GLOBAL CAREX GROUP

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fresh material of four species provide a clear picture

of structural variability. She emphasized the strong

inﬂuence of position in the inﬂorescence and vigour of

the plant in determining branching patterns, sex

expression and whether or not a perigynium contains

a ﬂower. Such variability in ﬂoral and inﬂorescence

structure in response to environment, either internal

or external, calls into question their value as taxo-

nomic characters. An understanding of the intrinsic

and extrinsic controls on inﬂorescence structure

would be useful in evaluating similarities in inﬂores-

cence form across clades in Carex s.l.

With new insights from molecular biology regard-

ing the control of ﬂoral and inﬂorescence structure in

angiosperms, and especially in other commelinids

such as Poaceae, it is becoming clear that even small

changes in the regulation of gene expression can

have signiﬁcant effects on ﬂoral and inﬂorescence

morphology (Doust & Kellogg, 2002; Bommert et al.,

2005; Thompson & Hake, 2009). For example, a

single amino acid substitution in the transcription

factor OsMADS1 can cause paleas, lemmas and lodi-

cules to become leafy and can decrease stamen

number in rice plants with this mutation (Jeon et al.,

2000; Bommert et al., 2005). The behaviour of mer-

istems in the inﬂorescence is regulated by a variety

of transcription factors (e.g. MADS-box genes) that

control whether meristems produce branches or

ﬂowers, the nature of the associated bracts, the

extent of internode elongation and the identity of

different parts of the ﬂowers and spikelets in grasses

(Kellogg, 2000; Bommert et al., 2005; Thompson &

Hake, 2009). A dynamic model of grass inﬂorescence

development describes a series of developmental

switches that determine whether meristems will

branch or will terminate in ﬂowers. These switches

also regulate the number of meristems produced, the

extent of internode or bract growth and the phyllo-

taxy (Kellogg, 2000; Bommert et al., 2005). Given the

conservation of regulatory function among dicots,

there is good reason to expect that similar models

might apply to both sedges and grasses, because they

are both commelinid monocots with spikelets

arranged in complex and variable inﬂorescences. It is

clear from the earlier discussion that control of

branching and the fate of meristems are critical not

only to the architecture of the inﬂorescence of Carex

s.l., but also to patterns of sex expression, pistillate

ﬂowers being produced only at branching events and

staminate ﬂowers most often positioned on axes

where branching has stopped.

Genetic and regulatory controls of inﬂorescence

structure are important, but environmental and hor-

monal signals also play a role (McSteen, 2009). Auxin,

cytokinin and strigolactone are all involved in the fate

of meristems within the inﬂorescence. Basipetal

movement of auxin from the apical meristem inhibits

bud meristems from growing (apical dominance) and

acropetal movement of cytokinin and strigolactone

promotes and inhibits bud growth, respectively.

Shading is also known to inhibit branching, and soil

nutrients can promote it. Genetic, regulatory, hormo-

nal and environmental controls thus determine the

ﬁnal structure of an inﬂorescence (McSteen, 2009). It

is important to remember that the induction of ﬂow-

ering is a dynamic process, mediated by growth regu-

lators, which are inﬂuenced by intrinsic and extrinsic

factors, including the carbohydrate status of the

plant, temperature and day length. Position in rela-

tion to root and shoot meristems that produce growth

regulators can be a strong inﬂuence on whether male

or female ﬂowers are produced in monoecious plants

such as Carex s.l. We do not yet have a complete

understanding of the way in which these various

regulatory pathways interact, but it appears likely

that simple changes in the regulation of ﬂoral and

inﬂorescence structure can explain the variability we

see in Carex s.l. Understanding how these regulatory

systems work should help us evaluate which aspects

of ﬂoral and inﬂorescence structure are stable within

clades and which are not, making it easier to select

appropriate characters to deﬁne subgroupings in

Carex s.l.

Although the similarity of form in genera of

Cariceae has long been recognized (e.g. Clarke, 1883;

Nelmes, 1951; Kern, 1958), they have been main-

tained as distinct genera for so long in part because of

their allopatry. Schoenoxiphium is restricted to

eastern and southern Africa, with a single species

reaching the south-western parts of the Arabian Pen-

insula, whereas Uncinia grows around Antarctica,

ranging from Australia, New Zealand, Chile and

Argentina north to the Philippines, Hawaii and the

Caribbean. Kobresia has its centre of distribution and

greatest species richness in the Himalayas, with a

few wide-ranging circumboreal species. Given the

ﬂexibility of inﬂorescence structure and the possibili-

ties for differential suppression of parts that might

become genetically ﬁxed, it is reasonable to expect

that the same basic constraints on the inﬂorescence

might give parallel results in lineages that colonized

different regions of the world. Although not identical

by descent, the rachillae bearing male ﬂowers apically

in Schoenoxiphium and Kobresia have positional

homology and could reasonably represent parallel

evolution of similar structures (Vrijdaghs et al., 2010).

Similarly, selection pressures in the harsh, windy and

often nutrient-poor conditions in which many unispi-

cate species of Carex s.l. (including Kobresia) are

found may have favoured reduced branching and

smaller stature, again resulting in parallel develop-

ment of the same traits in different lineages.

MAKING CAREX MONOPHYLETIC 17

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CONCLUSIONS

Much has been accomplished towards an understand-

ing of Carex s.l. since the publication of Kükenthal’s

(1909) monograph, from species discovery to detailed

analysis of ﬂoral and inﬂorescence morphology to

DNA-based phylogenetic trees. Species of Carex s.l.

continue to be discovered at a surprising rate even in

well-explored areas, such as Europe, North America,

Japan, Australia and New Zealand, and at even

higher rates in China, South-East Asia, Africa and

South America. Detailed studies of ﬂoral and inﬂo-

rescence morphology have been conducted for most

major groups of Carex s.l., although more work is

needed on the Vigneastra group and Kobresia. At least

one gene has been sequenced for more than half of the

species in the tribe, and nuclear and plastid DNA

regions, coding and non-coding, have been sequenced

for more than half of these in ongoing phylogenetic

studies. Despite this progress, there has been under-

standable hesitancy for anyone to reclassify a large

and widespread group such as Cariceae unilaterally,

despite frequent comments on the difficulty of distin-

guishing the genera in it. As a global group of Cyper-

aceae and Cariceae specialists, we think it is time to

overcome the inertia of the traditional classiﬁcation

and make the required nomenclatural changes to

recognize all species in the tribe as the genus Carex

s.l. Much remains to be accomplished on a global scale

to continue exploration and species discovery, to

understand the ways in which genetic and environ-

mental factors inﬂuence inﬂorescence variability

within and between species, and to take advantage of

the constantly expanding set of molecular tools and

analytical methods to formulate phylogenetic hypoth-

eses for the monophyletic genus Carex s.l.

The new broader circumscription of Carex proposed

here is just the ﬁrst step in the reclassiﬁcation of the

genus. The Global Carex Group is working towards a

complete global sampling of species of Carex s.l.,

sequencing multiple DNA regions per species, to aid

in placing species into natural sectional groups in the

genus. Molecular phylogenetic work already com-

pleted on Carex section Ovales (Hipp et al., 2006),

section Spirostachyae (Drejer) L.H.Bailey (Escudero

et al., 2007; Escudero & Luceño, 2009, 2011), section

Phyllostachyae Tuck. ex Kük. (Starr et al., 1999) and

section Ceratocystis Dumort. (Jiménez-Mejías,

Martín-Bravo & Luceño, 2012; Derieg et al., 2013)

illustrates the potential for clarifying the relation-

ships at the sectional level with DNA analyses. Pro-

jects currently in progress by our group include: an

expanded phylogenetic analysis and new monograph

of Schoenoxiphium as a section of Carex, a new

molecular phylogenetic analysis of Carex subgenus

Vignea, and additional work on the rest of the caricoid

clade to elucidate natural groups in Kobresia and

unispicate species. We are also working on reclassiﬁ-

cations of sections Glareosae,Phleoideae (Meinsh.)

T.V. Egorova and Remotae (Asch.) C.B.Clarke in the

Vignea clade and sections Aulocystis Dumort., Chlo-

rostachyae Tuckerm. ex Meinsh., Hymenochlaenae

s.l.,Porocystis,Phacocystis s.l.,Racemosae,Mitratae

Kük., Rhomboidales,Laxiﬂorae (Kunth) Mack., Pan-

iceae G.Don, Bicolores,Careyanae Tuck. ex Kük.,

Griseae (L.H.Bailey) Kük., Granulares (O.Lang)

Mack., Rostrales Meinsh., Vesicariae,Paludosae,

Carex,Lupulinae,Squarrosae J.Carey, Sylvaticae

Rouy, Rhynchocystis Dumort. and Indicae Tuck. in the

core Carex clade based on global sampling and

informed by molecular data.

TAXONOMIC TREATMENT

NEW NAME IN CAREX

Carex zikae E.H.Roalson & M.J.Waterway, nom.

nov.

Replaced synonym: Carex brevicaulis Mack., Bull.

Torrey Bot. Club 40: 547. 1913, nom. illeg. Carex

deﬂexa var. brevicaulis B.Boivin, Le Naturaliste

Canadien 94(4): 522. 1967; non Carex brevicaulis

Thouars (1808).

Distribution: North-western North America.

Etymology: The new name for this Paciﬁc Northwest

species recognizes the important contributions of

Peter Zika (WTU, University of Washington, Seattle,

WA, USA) to Carex systematics in western North

America.

Note: Despite the long usage of Carex brevicaulis

Mack. in North America, it is clear that this is a later

homonym of Carex brevicaulis Thouars (Esquisse Fl.

Tristan d’Acugna: 35. 1808), a species most recently

treated in Uncinia.Carex brevicaulis Thouars has

priority in Carex over C. brevicaulis Mack.

INCLUSION OF CYMOPHYLLUS MACK.EX BRITTON &

A.BR.IN CAREX L.

Carex fraseriana Ker Gawl., Bot. Mag. 33: t.

1391. 1811.

Cymophyllus fraserianus (Ker Gawl.) Kartesz &

Gandhi, Rhodora 93: 138. 1991.

Carex fraseri Andrews, Bot. Repos. 10: t. 639. 1811.

Olamblis fraseri (Andrews) Raf., Good Book: 26. 1840.

Cymophyllus fraseri (Andrews) Mack. in N.L.Britton

& A.Brown, Ill. Fl. N. U.S., ed. 2, 1: 441. 1913.

Mapania sylvatica Pursh, Fl. Amer. Sept. 1: 47.

1813, nom. illeg.

Carex lagopus Muhl., Descr. Gram.: 265. 1817.

18 GLOBAL CAREX GROUP

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Distribution: eastern USA (southern Appalachian

Mts.).

TRANSFERS FROM KOBRESIA WILLD.TO CAREX L.

Carex alatauensis S.R.Zhang, nom. nov.

Replaced synonym: Elyna humilis C.A.Mey. ex

Trautv., Trudy Imp. S.-Peterburgsk. Bot. Sada 1: 21.

187. 1871. Kobresia humilis (C.A.Mey. ex Trautv.)

Serg. in Schischkin, Fl. URSS 3: 111. 1935. Kobresia

royleana (Nees) Boeckeler var. humilis (C.A.Mey. ex

Trautv.) Kük. in Engler (ed.), Pﬂanzenr. 38(IV, 20): 46.

1909; non Carex humilis Leyss. (1761).

Kobresia persica Kük. & Bornm., Oesterr. Bot. Z.

47: 133. 1897; non Carex persica Nelmes (1939).

Kobresia royleana (Nees) Boeckeler var. parvinux

T.Koyama, Acta Phytotax. Geobot. 16: 168. 1956.

Distribution: Central Asia, north-western China.

Etymology: The type of the replaced synonym, Elyna

humilis C.A.Mey. ex Trautv., was collected from the

Alatau Mountains in Central Asia.

Carex bhutanensis S.R.Zhang, nom. nov.

Replaced synonym: Kobresia prainii Kük., Bull. Herb.

Boissier, sér. 2, 4: 50. 1904; non Carex prainii Kük.

(1903).

Kobresia utriculata C.B.Clarke, Bull. Misc. Inform.

Kew, Addit. Ser. 8: 67. 1908; non Carex utriculata

Boott (1939).

Kobresia prainii Kük. var. elliptica Y.C.Yang in

C.Y.Wu, Fl. Xizang. 5: 387, f. 217. 1987.

Distribution: Eastern Himalaya (Nepal, Sikkim,

Bhutan) to south-western China.

Etymology: The type of the replaced name, Kobresia

prainii Kük., was collected in Bhutan.

Carex bistaminata (W.Z.Di & M.J.Zhong)

S.R.Zhang, comb. nov.

Basionym: Kobresia bistaminata W.Z.Di & M.J.Zhong,

Acta Bot. Boreal.-Occid. Sin. 6(4): 275. 1986. Kobresia

myosuroides (Vill.) Fiori ssp. bistaminata (W.Z.Di &

M.J.Zhong) S.R.Zhang, Novon, 9: 453. 1999.

Kobresia kashgarica Dickoré, Stapﬁa, 39: 79. 1995.

Distribution: From Karakorum to western China.

Carex bonatiana (Kük.) Ivanova, Bot. Zhurn.

SSSR 24: 501. 1939

Basionym: Kobresia bonatiana Kük., Bull. Géogr. Bot.

22: 250. 1912.

Kobresia fragilis C.B.Clarke, J. Linn. Soc., Bot. 36:

267. 1903. Schoenoxiphium fragile (C.B.Clarke)

C.B.Clarke, Bull. Misc. Inform. Kew, Addit. Ser. 8: 67.

1908. non Carex fragilis Boott (1858).

Carex curvata Boott, Ill. Gen. Carex 1: 2, pl. 5.

1858, non Knaf (1847). Kobresia curvata C.B.Clarke,

Bull. Misc. Inform. Kew, Addit. Ser. 8: 68. 1908.

Schoenoxiphium clarkeanum Kük., Bull. Herb.

Boissier, sér. 2, 4: 49. 1904. Kobresia clarkeana (Kük.)

Kük. in Engler (ed.), Pﬂanzenr. 38(IV, 20): 48. 1909;

non Carex clarkeana Kük. (1904).

Kobresia clarkeana (Kük.) Kük. var. megalantha

Kük., Bull. Géogr. Bot. 22: 249. 1912.

Kobresia hispida Kük., Acta Horti Gothob. 5: 39.

1930.

Kobresia yuennanensis Hand.-Mazz., Symb. Sin. 7:

1256. 1936.

Kobresia curticeps (C.B.Clarke) Kük. var. gyiron-

gensis Y.C.Yang in C.Y.Wu, Fl. Xizang. 5: 391, f. 222.

1987.

Distribution: Nepal, Sikkim, Bhutan, south-western

China.

Note: The correct name for this species if the segre-

gate genus is recognized is Kobresia fragilis.

Carex borealipolaris S.R.Zhang, nom. nov.

Replaced synonym: Kobresia sibirica (Turcz. ex

Ledeb.) Boeckeler, Linnaea 39: 7. 1875. Elyna sibirica

Turcz. ex Ledeb., Fl. Ross. 4(2): 262. 1852; non Carex

sibirica Willd. ex Kunth (1837).

Kobresia arctica A.E.Porsild, Sargentia 4: 15. 1943,

non Meinsh. (1901).

Kobresia macrocarpa Clokey ex Mack., N. Amer. Fl.

18: 5. 1931. Kobresia bellardii (All.) Degl. ex Loisel.

var. macrocarpa (Clokey ex Mack.) H.D.Harr., Man.

Pl. Colorado 641. 1954; non Carex macrocarpa Phil.

(1858).

Kobresia smirnovii N.A.Ivanova, Bot. Zhurn.

S.S.S.R. 24: 480. 1939; non Carex smirnovii V.I.Krecz.

(1935).

Kobresia hyperborea A.E.Porsild, Bull. Natl. Mus.

Canada 121: 103. 1951; non Carex hyperborea Drejer

(1841).

Kobresia hyperborea A.E.Porsild var. alaskana

Duman, Bull. Torrey Bot. Club 83: 194. 1956.

Kobresia hyperborea A.E.Porsild var. lepagei

Duman, Bull. Torrey Bot. Club 83: 194. 1956. Kobre-

sia schoenoides (C.A.Mey.) Steud. var. lepagei

(Duman) B.Boivin, Naturaliste Canad. 94: 525. 1967.

Distribution: Subarctic to north-western USA.

MAKING CAREX MONOPHYLETIC 19

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Etymology: The species is circumboreal, and common

in the North Polar area. The epithet refers to the

distributional pattern of the species.

Carex brandisii (C.B.Clarke ex Jana &

R.C.Srivast.) O.Yano, comb. nov.

Basionym: Kobresia brandisii C.B. Clarke ex Jana &

R.C.Srivast., J. Jap. Bot. 89: 205, f. 1&2. 2014.

Distribution: Western Himalaya (India).

Carex breviprophylla O.Yano, nom. nov.

Replaced synonym: Kobresia gandakiensis Rajbh. &

H.Ohba in H. Ohba & S. B. Malla (eds.), Himal. Pl. 2:

132, f. 4h-o. 1991; non Carex gandakiensis Katsuy.

(2008).

Distribution: Nepal to Sikkim.

Etymology: This species is characterized by a prophyll

that is shorter than the nutlet.

Carex burangensis (Y.C.Yang) S.R.Zhang, comb.

nov.

Basionym: Kobresia burangensis Y.C.Yang in C.Y.Wu,

Fl. Xizang. 5: 374, f. 208. 1986.

Distribution: South-western China (western Tibet).

Carex capillifolia (Decne.) S.R.Zhang, comb. nov.

Basionym: Elyna capillifolia Decne. in Jacquemont,

Voy. Inde. 4(Bot.): 173. 1844. Kobresia capillifolia

(Decne.) C.B.Clarke, J. Linn. Soc., Bot. 20: 378. 1883.

Elyna spicata Boiss., Fl. Orient. 5: 394. 1882, non

Schrad. (1806).

Kobresia brunnescens Boeckeler, Beitr. Cyper. 1: 40.

1888.

Kobresia elata Boeckeler, Beitr. Cyper. 2: 32. 1890.

Kobresia macrolepis Meinsh., Trudy Imp.

S.-Peterburgsk. Bot. Sada 18: 276. 1901. Elyna mac-

rolepis (Meinsh.) Fomin & Woronow, Opred. Rast.

Kavk. 1: 173. 1909.

Kobresia capilliformis Ivanova, Bot. Zhurn. SSSR

24: 484. 1939.

Kobresia thomsonii Maxim. ex Ivanova, Bot. Zhurn.

SSSR 24: 486. 1939, pro syn.

Kobresia oviczinnikovii T.V.Egorova in Grubov, Pl.

As. Centr. 3: 33. 1967, syn. nov.

Kobresia yushuensis Y.C.Yang, Acta Biol. Plateau

Sin. 2: 6. 1984.

Distribution: From Caucasus to western China.

Carex cercostachys Franch., Bull. Soc. Philom.

Paris, 8, 7: 27. 1895

Kobresia cercostachys (Franch.) C.B.Clarke, J. Linn.

Soc., Bot. 37: 267. 1903.

Kobresia stiebritziana Hand.-Mazz., Akad. Wiss.

Wien, Math.-Naturwiss. Kl., Anz. 57: 54. 1920.

Kobresia nepalensis (Nees) Kük. var. stiebritziana

(Hand.-Mazz.) R.C.Srivast., Novon 8(2): 203. 1998.

Distribution: Bhutan, Sikkim, south-western China.

Carex clavispica S.R.Zhang, nom. nov.

Replaced synonym: Kobresia duthiei C.B.Clarke in

J.D.Hooker, Fl. Brit. India 6: 697. 1894; non Carex

duthiei C.B.Clarke (1894).

Kobresia rostrata C.B.Clarke ex Ivanova, Bot.

Zhurn. SSSR 24: 500. 1939, nom. nud., pro syn.; non

Carex rostrata Stokes (1787).

Distribution: Himalaya, south-western China.

Etymology: The inﬂorescence of the species is a

clavate spike.

Carex coninux (F.T.Wang & Tang) S.R.Zhang,

comb. nov.

Basionym: Kobresia coninux F.T.Wang & Tang, Acta

Phytotax. Sin. 1: 182. 1951.

Kobresia pusilla Ivanova, Bot. Zhurn. SSSR 24:

496. 1939; non Carex pusilla Arv.-Touv. (1872).

Kobresia helanshanica W.Z.Di & M.J.Zhong, Acta

Bot. Boreal.-Occid. Sin. 5 (4): 311, f. 1. 1985.

Kobresia daqingshanica X.Y.Mao, Acta Sci. Nat.

Univ. Intramongol. 19(2): 341, f. 1. 1988.

Kobresia karakorumensis Dickoré, Stapﬁa 39: 77.

1995, syn. nov.

Distribution: Karakorum, western Himalaya, north-

ern and western China (Gansu, Hebei, Nei Mongol,

Qinghai, Shanxi, Sichuan, Tibet and Yunnan).

Note: The correct name for this species if the segre-

gate genus is recognized is Kobresia pusilla.

Carex curticeps C.B.Clarke in J.D.Hooker, Fl.

Brit. India 6: 749. 1894

Kobresia curticeps (C.B.Clarke) Kük. in Engler (ed.),

Pﬂanzenr. 38(IV, 20): 47. 1909.

Distribution: Central & eastern Himalaya to southern

Tibet.

Carex deasyi (C.B.Clarke) O.Yano & S.R.Zhang,

comb. nov.

Basionym: Kobresia deasyi C.B.Clarke, Bull. Misc.

Inform. Kew, Addit. Ser. 8: 68. 1908.

20 GLOBAL CAREX GROUP

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Kobresia schoenoides (C.A.Mey.) Steud., Syn. Pl.

Glumac. 2: 246. 1855. Elyna schoenoides C.A.Mey.,

Verzeichn. Pﬂ. Cauc. Casp.: 29. 1831; non Carex

schoenoides Schrank (1789).

Kobresia pamiroalaica Ivanova, Bot. Zhurn. SSSR

24: 481. 1939.

Kobresia pamiralaica Ivanova in R.R. Schreder, Fl.

Uzbekist. 1: 347, 540. 1941.

Kobresia septatonodosa Koyama, Acta Phytotax

Geobot. 16: 168. 1956.

Kobresia maquensis Y.C.Yang, Acta Biol. Plateau

Sin. 2: 4, f. 3. 1984.

Kobresia lacustris P.C.Li, Acta Bot Yunnan. 12(1):

14. 1990.

Kobresia glaucifolia F.T.Wang & Tang ex P.C.Li,

Acta Phytotax. Sin. 37(2): 153. 1999.

Distribution: From Caucasus to western China.

Note: The correct name for this species if the segre-

gate genus is recognized is Kobresia schoenoides.

Carex esbirajbhandarii (Rajbh. & H.Ohba)

O.Yano, comb. nov.

Basionym: Kobresia esbirajbhandarii Rajbh. &

H.Ohba, J. Jap. Bot. 62: 272. 1987.

Distribution: Nepal.

Carex esenbeckii Kunth, Enum. Pl. 2: 522. 1837

Kobresia esenbeckii (Kunth) Noltie, Edinburgh J. Bot.

50(1): 43. 1993. Kobresia esenbeckii (Kunth) F.T.Wang

& Tang ex P.C.Li in W.T.Wang, Vasc. Pl. Hengduan

Mount. 2: 2352. 1994, later isonym.

Carex trinervis Nees in Wight, Contr. Bot. India

120. 1834, non Degl. (1807). Kobresia trinervis (Nees)

Boeckeler, Linnaea 39: 4. 1875. Hemicarex trinervis

(Nees) C.B.Clarke, J. Linn. Soc., Bot. 20: 382. 1883.

Kobresia seticulmis Boeckeler, Linnaea 39: 3. 1875.

Holmia seticulmis (Boeckeler) Fedde & J.Schust.,

Just’s Bot. Jahresber. 41(2): 10. 1913 (publ. 1918).

Kobresia hookeri Boeckeler, Linnaea 39: 4. 1875.

Hemicarex hookeri (Boeckeler) Benth., J. Linn. Soc.,

Bot. 18: 367. 1881.

Carex mutans Boott ex C.B.Clarke, J. Linn. Soc.,

Bot. 20: 383. 1883.

Carex polygyna Boeckeler, Beitr. Cyper. 1: 40. 1888.

Kobresia hookeri Boeckeler var. dioica C.B.Clarke

in J.D.Hooker, Fl. Brit. India 6: 695. 1894.

Kobresia angusta C.B.Clarke in J.D.Hooker, Fl.

Brit. India 6: 695. 1894.

Kobresia foliosa C.B.Clarke in J.D.Hooker, Fl. Brit.

India 6: 696. 1894.

Kobresia trinervis (Nees) Boeckeler var. foliosa

(C.B.Clarke) Kük. in Engler (ed.), Pﬂanzenr. 38(IV,

20): 43. 1909.

Distribution: Himalaya to south-western China.

Carex ﬁlispica S.R.Zhang, nom. nov.

Replaced synonym: Hemicarex ﬁlicina C.B.Clarke, J.

Linn. Soc., Bot. 20: 384. 1883. Kobresia ﬁlicina

(C.B.Clarke) C.B.Clarke in J.D.Hooker, Fl. Brit. India

6: 696. 1894; non Carex ﬁlicina Nees (1834).

Kobresia ﬁlicina (C.B.Clarke) C.B.Clarke var. sub-

ﬁlicinoides P.C.Li, Acta Phytotax. Sin. 37(2): 155.

1999; non Carex subﬁlicinoides Kük. (1930).

Distribution: Eastern Himalaya, south-western

China.

Etymology: The ﬁrst part of the name, ﬁli-, thread-

like, from ﬁlum, a thread, here refers to the narrowly

linear spike of this species.

Carex ﬁssiglumis (C.B.Clarke) S.R.Zhang &

O.Yano, comb. nov.

Basionym: Kobresia ﬁssiglumis C.B.Clarke in

J.D.Hooker, Fl. Brit. India 6: 696. 1894. Kobresia

esenbeckii (Kunth) Noltie var. ﬁssiglumis (C.B.Clarke)

Noltie, Edinburgh J. Bot. 50(1): 43. 1993.

Distribution: Central Himalaya (western Nepal) to

south-western China.

Carex gammiei (C.B.Clarke) S.R.Zhang & O.Yano,

comb. nov.

Basionym: Kobresia gammiei C.B.Clarke, Bull. Misc.

Inform. Kew, Addit. Ser. 8: 68. 1908.

Kobresia williamsii Koyama, Bot. Mag. (Tokyo) 86

(1004): 279, pl. 3. 1973.

Distribution: Nepal, Bhutan, Sikkim, south-western

China.

Carex handel-mazzettii (Ivanova) S.R.Zhang,

comb. nov.

Basionym: Kobresia handel-mazzettii Ivanova, Bot.

Zhurn. SSSR 24: 494. 1939.

Kobresia capillifolia (Decne.) C.B.Clarke var. con-

densata Kük., Notes Roy. Bot. Gard. Edinburgh 7:

134. 1912.

Kobresia condensata (Kük.) S.R.Zhang & Noltie, Fl.

China 23: 274. 2010.

Kobresia royleana (Nees) Boeckeler var. himalaica

Rajbh. & H.Ohba in H. Ohba & S. B. Malla (eds.),

Himal. Pl. 2: 150. 1991.

MAKING CAREX MONOPHYLETIC 21

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Distribution: South-western China (Sichuan, Tibet,

Yunnan), Nepal.

Carex harae (Rajbh. & H.Ohba) O.Yano, comb.

nov.

Basionym: Kobresia harae Rajbh. & H.Ohba, J. Jap.

Bot. 62: 193 (1987), as ‘harai’.

Distribution: India, Nepal.

Carex hohxilensis (R.F.Huang) S.R.Zhang, comb.

nov.

Basionym: Kobresia hohxilensis R.F.Huang, Biol. &

Human Physiol. Hoh Xil Region, 101. 1996.

Kobresia stolonifera Y.C.Tang ex P.C.Li, Acta Phy-

totax. Sin. 37(2): 154. 1999.

Distribution: Western China (Gansu, Qinghai, Tibet).

Carex hughii S.R.Zhang, nom. nov.

Replaced synonym: Kobresia graminifolia C.B.Clarke,

J. Linn. Soc., Bot. 36: 268. 1903; non Carex gramini-

folia Cherm. (1923).

Distribution: Western China (Gansu, Qinghai,

Shaanxi, Sichuan, Tibet, Yunnan).

Etymology: The epithet of the species is adopted to

commemorate Rev. Fr. Hugh, the collector of the type

of Kobresia graminifolia C.B.Clarke.

Carex kanaii (Rajbh. & H.Ohba) S.R.Zhang &

O.Yano, comb. nov.

Basionym: Kobresia kanaii Rajbh. & H.Ohba in H.

Ohba & S. B. Malla (eds.), Himal. Pl. 2: 135, f. 6.

1991.

Distribution: Nepal, Sikkim.

Carex kangdingensis S.R.Zhang, nom. nov.

Replaced synonym: Kobresia falcata F.T.Wang & Tang

ex P.C.Li, Acta Bot. Yunnan 12 (1): 18, f. 9. 1990; non

Carex falcata Turcz. (1838).

Distribution: China (Sichuan, Gansu).

Etymology: Kangding (Sichuan, China) is the locality

in which the type of Kobresia falcata F.T.Wang &

Tang ex P.C.Li was collected.

Carex kobresioidea (Kük.) S.R.Zhang, comb. nov.

Basionym: Schoenoxiphium kobresioideum Kük.,

Bull. Jard. Bot. Buitenzorg III 16: 312. 1940. Kobresia

kobresioidea (Kük.) J. Kern, Acta Bot. Neerl. 7: 795.

1958.

Distribution: Northern Sumatra.

Carex kokanica (Regel) S.R.Zhang, comb. nov.

Basionym: Elyna kokanica Regel, Trudy Imp.

S.-Peterburgsk. Bot. Sada 7: 563. 1880.

Trilepis royleana Nees, Linnaea 9: 305. 1834; non

Carex royleana Nees (1834). Kobresia royleana (Nees)

Boeckeler, Linnaea 39: 8. 1875. Kobresia stenocarpa

(Kar. & Kir.) Steud. var royleana (Nees) C.B.Clarke,

J. Linn. Soc., Bot. 20: 381. 1883.

Elyna stenocarpa Kar. & Kir., Bull. Soc. Imp. Natu-

ralistes Moscou 15(3): 526. 1842; non Carex steno-

carpa Turcz. ex Krecz. (1935). Kobresia stenocarpa

(Kar. & Kir.) Steud., Synop. Pl. Glum. 2: 246. 1854.

Kobresia stenocarpa (Kar. & Kir.) Steud. var.

simplex Y.C.Yang in C.Y.Wu, Fl. Xizang. 5: 395, f. 224.

1987.

Kobresia paniculata Meinsh., Trudy Imp.

S.-Peterburgsk. Bot. Sada 18(3): 279. 1901. Kobresia

royleana (Nees) Boeckeler var. paniculata (Meinsh.)

Kük. in Engler (ed.), Pﬂanzenr. 38(IV, 20): 46. 1909.

K. minshanica F.T.Wang & Tang ex Y.C.Yang, Acta

Biol. Plateau Sin. 2: 1. 1984. Kobresia royleana (Nees)

Boeckeler ssp. minshanica (F.T.Wang & Tang ex

Y.C.Yang) S.R.Zhang, Novon 9: 453. 1999.

K. menyuanica Y.C.Yang, Acta Biol. Plateau Sin. 2:

3. 1984.

Distribution: Western Asia (Afghanistan), Central

Asia, Himalaya, western China.

Note: The correct name for this species if the segre-

gate genus is recognized is Kobresia royleana.

Carex lepidochlamys (F.T.Wang & Tang ex

P.C.Li) S.R.Zhang, comb. nov.

Basionym: Kobresia lepidochlamys F.T.Wang & Tang

ex P.C.Li, Acta Bot. Yunnan. 12(1): 15, f. 6. 1990.

Kobresia cuneata Kük., Acta Horti Gothob. 5: 39.

1930; non Carex cuneata Ohwi (1931).

Distribution: South-western China (Gansu, Sichuan,

Tibet, Yunnan).

Note: The correct name for this species if the segre-

gate genus is recognized is Kobresia cuneata.

Carex liangshanensis S.R.Zhang, nom. nov.

Replaced synonym: Kobresia kuekenthaliana Hand.-

Mazz., Symb. Sin. 7: 1258. 1936. Schoenoxiphium

kuekenthalianum (Hand.-Mazz.) Ivanova, Bot. Zhurn.

SSSR 24: 501. 1939; non Carex kuekenthaliana Appel

& A.Brückn. (1891).

Distribution: South-western China (Sichuan).

22 GLOBAL CAREX GROUP

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Etymology: Liangshan (Sichuan, China) is the locality

in which the type of the replaced synonym was

collected.

Carex littledalei (C.B.Clarke) S.R.Zhang, comb.

nov.

Basionym: Kobresia littledalei C.B.Clarke, Bull. Misc.

Inform. Kew, Addit. Ser. 8: 67. 1908. Kobresia tibetica

Maxim. var. littledalei (C.B.Clarke) P.C.Li in

W.T.Wang, Vasc. Pl. Hengduan Mount. 2: 2349. 1994.

Distribution: South-western China (Sichuan and

Tibet).

Carex macroprophylla (Y.C.Yang) S.R.Zhang,

comb. nov.

Basionym: Kobresia ﬁlifolia (Turcz.) C.B.Clarke var.

macroprophylla Y.C.Yang, Acta Biol. Plateau Sin. 2: 8,

f. 5. 1984. Kobresia macroprophylla (Y.C.Yang) P.C.Li

in L.K.Dai & S.Y.Liang, Fl. Reipubl. Popul. Sin. 12:

17. 2000.

Kobresia ﬁlifolia (Turcz.) C.B.Clarke, J. Linn. Soc.,

Bot. 20: 381. 1883. Elyna ﬁlifolia Turcz., Bull. Soc.

Imp. Naturalistes Moscou 28(1): 353. 1855; non Carex

ﬁlifolia Nutt. (1818). Kobresia capillifolia (Decne.)

C.B.Clarke var. ﬁlifolia (Turcz.) Kük., Finska Vet.-

Soc. Föhr. 65(8): 1. 1902–1903.

Kobresia gracilis Meinsh., Acta Horti Petrop. 18:

276. 1901; non Carex gracilis Curtis (1782).

Kobresia pratensis Freyn, Oesterr. Bot. Z. 40: 266.

1890; non Carex pratensis Hosé (1797).

Distribution: Northern China, Mongolia, Siberia.

Note: The correct name for this species if the segre-

gate genus is recognized is Kobresia ﬁlifolia.

Carex mallae (Rajbh. & H.Ohba) O.Yano, comb.

nov.

Basionym: Kobresia mallae Rajbh. & H.Ohba, J. Jap.

Bot. 62: 270. 1987.

Distribution: Nepal.

Carex myosuroides Vill., Prosp. Hist. Pl.

Dauphine 17–18. 1779

Kobresia myosuroides (Vill.) Fiori in Fiori et al., Fl.

Anal. Ital. 1: 125. 1896. Elyna myosuroides (Vill.)

Fritsch ex Janch., Mitt. Naturwiss. Vereins Univ.

Wien 5: 110. 1907.

Carex bellardii All., Fl. Pedem. 2: 264. 1785. Kob-

resia bellardii (All.) Degl. ex Loisel., Fl. Gall. 2: 626.

1807.

Carex hermaphrodita J. F. Gmel, Syst. Nat. 2: 138.

1791.

Kobresia scirpina Willd., Sp. Pl. ed. 4, 4(1): 205.

1805.

Elyna spicata Schrad., Fl. Germ. 1: 155. 1806.

Carex affinis R.Br., Bot. App. 750. 1823.

Kobresia nardina Hornem. in G. C. Oeder & al., Fl.

Dan. 74. 1827.

Kobresia ﬁliformis Dewey, Amer. J. Sci. Arts 29:

253. 1836.

Elyna ﬁliformis Steud., Syn. Pl. Glumac. 2: 245.

1855.

Carex vulcanicola Nakai, Bot. Mag. (Tokyo) 28: 327.

1914.

Distribution: Europe, northern Asia, northern North

America, Greenland.

Carex neesii S.R.Zhang, nom. nov.

Replaced synonym: Kobresia nepalensis (Nees) Kük.

in Engler (ed.), Pﬂanzenr. 38(IV, 20): 46: 40, f. 9. 1909.

Uncinia nepalensis Nees in Wight, Contr. Bot. India

129. 1834; non Carex nepalensis Spreng. (1826).

Carex linearis Boott, Ill. Gen. Carex 1: 51, pl. 136.

1858, non Clairv. (1811). Hemicarex linearis (Boott)

Benth., J. Linn. Soc., Bot. 18: 367. 1881.

Carex linearis Boott var. elachista C.B.Clarke in

J.D.Hooker, Fl. Brit. India 6: 713. 1894. Kobresia

nepalensis (Nees) Kük. var elachista (C.B.Clarke)

Kük. in Engler (ed.), Pﬂanzenr. 38(IV, 20): 40. 1909.

Distribution: Western Himalaya to south-western

China.

Etymology: The epithet of the species is adopted to

commemorate Christian Gottfried Daniel Nees von

Esenbeck (1776–1858) who published the ﬁrst name

for this species.

Carex noltiei S.R.Zhang, nom. nov.

Replaced synonym: Kobresia woodii Noltie, Edin-

burgh J. Bot. 50(1): 48, f. 1h-l. 1993; non Carex woodii

Dewey (1846).

Distribution: Bhutan, south-western China (southern

Tibet).

Etymology: The epithet of the species is adopted to

commemorate Henry John Noltie (Royal Botanic

Garden, Edinburgh) who published the replaced

synonym Kobresia woodii.

Carex nudicarpa (Y.C.Yang) S.R.Zhang, comb.

nov.

Basionym: Blysmocarex nudicarpa Y.C.Yang, Acta

Bot. Yunnan. 4(4): 325. 1982. Kobresia nudicarpa

(Y.C.Yang) S.R.Zhang, Acta Phytotax. Sin. 33(2): 160.

1995. Kobresia macrantha Boeckeler var. nudicarpa

MAKING CAREX MONOPHYLETIC 23

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(Y.C.Yang) P.C.Li in L.K.Dai & S.Y.Liang, Fl. Reipubl.

Popul. Sin. 12: 26. 2000. Blysmocarex macrantha

(Boeckeler) Ivanova ssp. nudicarpa (Y.C.Yang)

D.S.Deng, Guihaia 22: 120. 2002.

Kobresia macrantha Boeckeler, Beitr. Cyper. 1: 39.

1888. Blysmocarex macrantha (Boeckeler) Ivanova,

Bot. Zhurn. SSSR 24: 502. 1939; non Carex macran-

tha Boeckeler (1888).

Distribution: South-western China, Himalaya,

eastern Karakorum.

Note: The correct name for this species if the segre-

gate genus is recognized is Kobresia macrantha.

Carex ovoidispica O.Yano, nom. nov.

Replaced synonym: Kobresia nitens C.B.Clarke, J.

Linn. Soc., Bot. 20: 379, pl. 30, f. 7. 1883; non Carex

nitens Phil. (1873).

Distribution: Northern Pakistan to Nepal.

Etymology: The speciﬁc epithet refers to the conspicu-

ous ovoid spikes.

Carex paramjitii (Jana, Noltie, R.C.Srivast &

A.Mukh.) O.Yano, comb. nov.

Basionym: Kobresia paramjitii Jana, Noltie, R.C.Sriv-

ast & A.Mukh., Indian J. Plant Sci. 3 (online): 106.

2014.

Distribution: Sikkim.

Carex parvula O.Yano, nom. nov.

Replaced synonym: Kobresia pygmaea (C.B.Clarke)

C.B.Clarke in J.D.Hooker, Fl. Brit. India 6: 696. 1894.

Hemicarex pygmaea C.B.Clarke, J. Linn. Soc., Bot.

20: 383. 1883; non Carex pygmaea Boeckeler (1876).

Kobresia pygmaea C.B.Clarke var. ﬁliculmis Kük.,

Acta Horti Gothob. 5: 37. 1930; non Carex ﬁliculmis

Franch. & Sav. (1878).

Kobresia microstachya Ivanova, Bot. Zhurn. SSSR

24: 488. 1939; non Carex microstachya Ehrh. (1784).

Kobresia koelzii Kük. ex Ivanova, Bot. Zhurn. SSSR

24: 498. 1939, pro syn.

Distribution: From Himalaya to northern and

western China.

Etymology: The speciﬁc epithet refers to the dwarf

habit.

Carex peichuniana S.R.Zhang, nom. nov.

Replaced synonym: Kobresia inﬂata P.C.Li, Acta Bot.

Yunnan. 12(1): 16. 1990; non Carex inﬂata Huds.

(1762).

Distribution: South-western China (north-western

Yunnan, south-eastern Tibet).

Etymology: The epithet of the species is adopted to

commemorate Pei-Chun Li (Shenzen Fairy Lake

Botanical Garden, Guangdong, China), who published

the replaced synonym Kobresia inﬂata.

Carex prainii Kük., Bull. Herb. Boissier, sér. 2, 4:

51. 1903

Replaced synonym: Kobresia sikkimensis Kük. in

Engler (ed.), Pﬂanzenr. 38 (IV, 20): 47. 1909; non

Carex sikkimensis C.B.Clarke (1894).

Distribution: Nepal to Assam.

Carex pseudogammiei S.R.Zhang, nom. nov.

Replaced synonym: Kobresia loliacea F.T.Wang &

Tang ex P.C.Li, Acta Bot. Yunnan. 12(1): 13. 1990; non

Carex loliacea L. (1753).

Distribution: South-western China (north-western

Yunnan, western Sichuan).

Etymology: The species is morphologically similar to

Carex gammiei (C.B.Clarke) S.R.Zhang.

Carex pseudolaxa (C.B.Clarke) O.Yano &

S.R.Zhang, comb. nov.

Basionym: Kobresia pseudolaxa C.B.Clarke, J. Linn

Soc., Bot. 20: 381. 1883.

Kobresia laxa Nees in Wight, Contr. Bot. India 119.

1834; non Carex laxa Wahlenb. (1803). Elyna laxa

(Nees) Kunth, Enum. Pl. 2: 534. 1837. Hemicarex laxa

(Nees) Benth., J. Linn. Soc., Bot. 18: 367. 1881. Schoe-

noxiphium laxum (Nees) Ivanova, Bot. Zhurn. SSSR

24: 501. 1939.

Schoenoxiphium hissaricum Pissjauk., Bot. Mater.

Gerb. Bot. Inst. Komarova Akad. Nauk SSSR. 12: 72.

1950. Kobresia hissarica (Pissjauk.) Soják, Nár. Mus.

Odd. Prír. 148: 194. 1979 (publ. 1980). Kobresia laxa

Nees ssp. hissarica (Pissjauk.) Kukkonen, Ann.

Naturhist. Mus. Wien, B 98B(Suppl.): 91. 1996.

Kobresia afghanica Raymond, Dansk Bot. Ark. 14:

17. 1965.

Distribution: Kashmir to central Himalaya.

Note: The correct name for this species if the segre-

gate genus is recognized is Kobresia laxa.

24 GLOBAL CAREX GROUP

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Carex pseuduncinoides (Noltie) O.Yano &

S.R.Zhang, comb. nov.

Basionym: Kobresia pseuduncinoides Noltie, Edin-

burgh J. Bot. 50(1): 47, f. 1a–g. 1993.

Kobresia kansuensis Kük., Acta Horti Gothob. 5: 38.

1930; non Carex kansuensis Nelmes (1939).

Distribution: Bhutan, India, Nepal, western China.

Note: The correct name for this species if the segre-

gate genus is recognized is Kobresia kansuensis.

Carex rcsrivastavae (Jana) E.H.Roalson,

comb. nov.

Basionym: Kobresia rcsrivastavae Jana, Indian J.

Fundam. Appl. Life Sci. 2: 256. 2012.

Distribution: India (Uttaranchal).

Carex sanguinea Boott, Proc. Linn. Soc. London

1: 285. 1846

Kobresia sanguinea (Boott) Raymond, Biol. Skr. 14(4):

19. 1965.

Distribution: Eastern Afghanistan to western Hima-

laya.

Carex sargentiana (Hemsl.) S.R.Zhang, comb.

nov.

Basionym: Kobresia sargentiana Hemsl., J. Linn.

Soc., Bot. 30: 139. 1894. Kobresia robusta Maxim. var.

sargentiana (Hemsl.) Kük. in Engler (ed.), Pﬂanzenr.

38(IV, 20): 36. 1909.

Kobresia robusta Maxim., Bull. Acad. Imp. Sci. St.-

Petersbourg, n.s., 29: 218. 1883; non Carex robusta F.

Nyl. (1844).

Distribution: Western China, Sikkim.

Note: The correct name for this species if the segre-

gate genus is recognized is Kobresia robusta.

Carex setschwanensis (Hand.-Mazz.) S.R.Zhang,

comb. nov.

Basionym: Kobresia setschwanensis Hand.-Mazz.,

Symb. Sin. 7: 1254. 1936.

Kobresia longearistita P.C.Li, Acta Bot. Yunnan.

12(1): 16, f. 8. 1990.

Kobresia pinetorum F.T.Wang & Tang ex P.C.Li,

Acta Bot. Yunnan: 12(1): 14, f. 5. 1990.

Distribution: Western China (southern Gansu, south-

ern Qinghai, Sichuan, Tibet, Yunnan).

Carex siamensis (Ohwi) S.R.Zhang, comb. nov.

Basionym: Kobresia siamensis Ohwi, Acta Phytotax.

Geobot. 23: 109. 1968.

Kobresia curvirostris (C.B.Clarke) C.B.Clarke in

J.D.Hooker, Fl. Brit. India 6: 699. 1894. Hemicarex

curvirostris C.B.Clarke, J. Linn. Soc., Bot. 20: 384.

1883; non Carex curvirostra Hartm. (1832).

Distribution: Eastern Himalaya to northern Thailand.

Note: The correct name for this species if the segre-

gate genus is recognized is Kobresia curvirostris.

Carex simpliciuscula Wahlenb., Kongl. Vetensk.

Acad. Nya Handl. 24(2): 141. 1803

Kobresia simpliciuscula (Wahlenb.) Mack., Bull.

Torrey Bot. Club 50: 349. 1923.

Carex bipartita All., Fl. Pedem. 2: 265. 1785, nom.

rej. Kobresia bipartita (All.) Dalla Torre, Atlas

Alpenﬂ. 2: 216. 1882.

Schoenus monoicus Sm., Engl. Bot. t. 1410. 1805.

Kobresia caricina Willd., Sp. Pl. ed. 4, 4: 206. 1805.

Carex lacustris Balbis ex Willd., Sp. Pl. ed. 4, 4:

206. 1805, pro syn.

Carex hybrida Schuhr ex Willd., Sp. Pl. 4: 206.

1805, pro syn.

Carex mirabilis Host, Icon. Descr. Gram. Austriac.

4: 44, pl. 78. 1809.

Elyna caricina Mert. et Koch in Röhling, Deutschl.

Fl. ed. 3, 1: 458. 1823.

Carex lobata Willd. ex Kunth, Enum. Pl. 2: 533.

1837, pro syn.

Kobresia simpliciuscula (Wahlenb.) Mack. var.

americana Duman, Bull. Torrey Bot. Club 83: 194.

1956.

Kobresia ﬁlifolia (Turcz.) C.B.Clarke ssp. subﬁlifo-

lia T.V.Egorova, Jurtzev & V.V.Petrovsky, Bot. Zhurn.

(Moscow & Leningrad) 66(7): 1042. 1981. Kobresia

simpliciuscula (Wahlenb.) Mack. ssp. subﬁlifolia

(T.V.Egorova, Jurtzev & V.V.Petrovsky) T.V.Egorova,

Novosti Sist. Vyssh. Rast., 20: 84. 1983. Kobresia

simpliciuscula (Wahlenb.) Mack. var. subﬁlifolia

(T.V.Egorova, Jurtzev & V.V.Petrovsky) A.E.Kozhevn.,

Sosud. Rast. Sovet. Dal’nego Vostoka, 3: 229. 1988.

Kobresia simpliciuscula (Wahlenb.) Mack. ssp. sub-

holarctica T.V.Egorova, Novosti. Sist. Vyssh. Rast., 20:

83. 1983. Kobresia simpliciuscula (Wahlenb.) Mack.

var. subholarctica (T.V.Egorova) A.E.Kozhevn., Sosud.

Rast. Sovet. Dal’nego Vostoka, 3: 229. 1988. Kobresia

subholarctica (T.V.Egorova) T.V.Egorova, Bot. Zhurn.

(Moscow & Leningrad) 76(12): 1736 1991.

Distribution: Europe to Caucasus, subarctic America

to western USA.

MAKING CAREX MONOPHYLETIC 25

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Carex squamiformis (Y.C.Yang) S.R.Zhang,

comb. nov.

Basionym: Kobresia squamiformis Y.C.Yang, Acta

Biol. Plateau Sin. 2: 9, f. 6. 1984, published as:

‘squmaeformis’.

Kobresia setschwanensis Hand.-Mazz. ssp. squami-

formis (Y.C.Yang) S.R.Zhang, Novon 9: 453. 1999.

Distribution: North-western China (southern Gansu,

eastern Qinghai).

Carex tibetikobresia S.R.Zhang, nom. nov.

Replaced synonym: Kobresia tibetica Maxim., Bull.

Acad. Imp. Sci. St.-Petersbourg, n.s., 29: 219. 1884.

Kobresia capillifolia (Decne.) C.B.Clarke var.

tibetica (Maxim.) Kük. in Engler (ed.), Pﬂanzenr.

38(IV, 20): 36. 1909; non Carex thibetica Franch.,

orth. var. (1887).

Distribution: Bhutan, western China.

Etymology: The epithet is based on the replaced

synonym, Kobresia tibetica. The species is mainly

distributed in the eastern Tibetan Plateau.

Carex tunicata (Hand.-Mazz.) S.R.Zhang, comb.

nov.

Basionym: Kobresia tunicata Hand.-Mazz., Symb.

Sin. 7: 1254. 1936.

Distribution: South-western China (north-western

Yunnan).

Carex uncinoides Boott, Ill. Gen. Carex 1: 8, pl.

23. 1858

Kobresia uncinoides (Boott) C.B.Clarke in

J.D.Hooker, Fl. Brit. India 6: 698. 1894.

Distribution: Eastern Himalaya (Nepal to Bhutan),

northern Myanmar, south-western China.

Carex vaginosa (C.B.Clarke) S.R.Zhang, comb.

nov.

Basionym: Kobresia vaginosa C.B.Clarke in

J.D.Hooker, Fl. Brit. India 6: 695. 1894. Kobresia

nepalensis (Nees) Kük. var. vaginosa (C.B.Clarke)

Kük. in Engler, Pﬂanzenr. 38(IV, 20): 40. 1909. Kob-

resia nepalensis (Nees) Kük. ssp. vaginosa

(C.B.Clarke) Koyama in Hara et al., Enum. Fl. Pl.

Nepal 1: 113. 1978. Kobresia nepalensis (Nees) Kük.

var. vaginosa (C.B.Clarke) R.C.Srivast., Fl. Sikkim 1:

225. 1996.

Kobresia cercostachys (Franch.) C.B.Clarke var.

capillacea P.C.Li, Acta Bot. Yunnan. 12(1): 17. 1990.

Distribution: Nepal, Sikkim, south-western China.

Carex vibhae (Jana, R.C.Srivast & Bhaumik)

O.Yano, comb. nov.

Basionym: Kobresia vibhae Jana, R.C.Srivast &

Bhaumik, Indian J. Plant Sci. 3 (online): 110, pl. 5.

2014.

Distribution: South-eastern Himalaya.

Carex vidua Boott ex C.B.Clarke in J.D.Hooker, Fl.

Brit. India 6: 713. 1894

Kobresia vidua (Boott ex C.B.Clarke) Kük. in Engler

(ed.), Pﬂanzenr. 38(IV, 20): 40. 1909.

Kobresia prattii C.B.Clarke, J. Linn. Soc., Bot. 36:

268. 1903.

Kobresia harrysmithii Kük., Acta Horti Gothob. 5:

37. 1930.

Distribution: Himalaya (Nepal, Sikkim, Bhutan) to

western China.

Carex yadongensis (Y.C.Yang) S.R.Zhang, comb.

nov.

Basionym: Kobresia yadongensis Y.C.Yang in C.Y.Wu,

Fl. Tibet. 5: 388, f. 219. 1987.

Distribution: South-western China (southern Tibet).

Carex yangii (S.R.Zhang) S.R.Zhang, comb. nov.

Basionym: Kobresia yangii S.R.Zhang, Acta Phytotax.

Sin. 33(2): 160. 1995.

Kobresia gracilis Y.C.Yang, Acta Biol. Plateau Sin.

2: 11, f. 7. 1984, non Meinsh. (1901).

Distribution: South-western China (south-western

Sichuan).

TRANSFERS FROM SCHOENOXIPHIUM NEES TO

CAREX L.

Carex basutorum (Turrill) Luceño &

Martín-Bravo, comb. nov.

Basionym: Schoenoxiphium basutorum Turrill, Bull.

Misc. Inform. Kew 1914: 19. 1914.

Distribution: South Africa (Free State), Lesotho.

Carex burkei (C.B.Clarke) Luceño &

Martín-Bravo, comb. nov.

Basionym: Schoenoxiphium burkei C.B.Clarke, J.

Linn. Soc. Bot. 20: 386. 1883.

26 GLOBAL CAREX GROUP

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Distribution: South Africa (Cape Province, Natal),

Lesotho.

Carex capensis Thunb., Prodr. Pl. Cap.: 14. 1794

Schoenoxiphium ecklonii Nees, Linnaea 10: 200.

1836. Archaeocarex ecklonii (Nees) Pissjauk., Bot.

Mater. Gerb. Bot. Inst. Komarova Akad. Nauk

S.S.S.R. 12: 83. 1950. Kobresia ecklonii (Nees)

T.Koyama, J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 8: 80.

1961.

Schoenoxiphium thunbergii Nees, Linnaea 9: 305.

1834. Archaeocarex thunbergii (Nees) Pissjauk., Bot.

Mater. Gerb. Bot. Inst. Komarova Akad. Nauk

S.S.S.R. 12: 83. 1950.

Schoenoxiphium altum Kukkonen, Notes Roy. Bot.

Gard. Edinburgh 43: 365. 1986.

Carex bisexualis C.B.Clarke in Harvey & auct. suc.

(eds.), Fl. Cap. 7: 302. 1898.

Carex capensis Schkuhr, Beschr. Riedgräs. 2: 39.

1806, nom. illeg.

Carex zeyheri C.B.Clarke in Harvey & auct. suc.

(eds.), Fl. Cap. 7: 303. 1898.

Schoenoxiphium ecklonii var. unisexuale Kük. in

Engler, Pﬂanzenr. 38(IV, 20): 33. 1909.

Distribution: South Africa (Cape Province).

Note: The correct name for this species if the segre-

gate genus is recognized is Schoenoxiphium ecklonii.

Carex chermezonii Luceño & Martín-Bravo, nom.

nov.

Replaced synonym: Schoenoxiphium gracile Cherm.,

Bull. Soc. Bot. France 70: 300. 1923; non Carex gra-

cilis Curtis (1782).

Distribution: Northern Madagascar (Mt. Tsaratan-

ana).

Etymology: The epithet honours Henri Chermezon

(1885–1939), a French botanist and explorer, who ﬁrst

described this species in 1923.

Carex distincta (Kukkonen) Luceño &

Martín-Bravo, comb. nov.

Basionym: Schoenoxiphium distinctum Kukkonen,

Bot. Not. 131: 263. 1978.

Distribution: South Africa (Free State?, Natal),

Lesotho.

Carex killickii Nelmes, Kew Bull. 10: 89. 1955

Replaced synonym: Schoenoxiphium ﬁliforme Kük.,

Bull. Misc. Inform. Kew 1910: 129. 1910; non Carex

ﬁliformis L. (1753).

Schoenoxiphium strictum Kukkonen, Notes Roy.

Bot. Gard. Edinburgh 43: 366. 1986.

Schoenoxiphium molle Kukkonen, Notes Roy. Bot.

Gard. Edinburgh 43: 366. 1986.

Distribution: South Africa (Cape Province, Free State,

Natal), Lesotho.

Carex kukkoneniana Luceño & Martín-Bravo,

nom. nov.

Replaced synonym: Schoenoxiphium buchananii

C.B.Clarke ex C.B.Clarke in Harvey & auct. suc.

(eds.), Fl. Cap. 7: 305. 1898. Carex buchananii

(C.B.Clarke ex C.B.Clarke) C.B.Clarke in Harvey &

auct. suc. (eds.), Fl. Cap. 7: 305. 1898, nom. illeg.; non

Carex buchananii Bergr. (1880). Kobresia buchananii

(C.B.Clarke) T.Koyama, J. Fac. Sci. Univ. Tokyo, Sect.

3, Bot. 8: 80. 1961.

Distribution: South Africa (Natal), Lesotho.

Etymology: The epithet honours Ilkka Kukkonen, a

Finnish botanist who studied Schoenoxiphium and

described several species.

Carex lancea (Thunb.) Baill., Hist. Pl. 12: 341.

1894

Basionym: Schoenus lanceus Thunb., Prodr. Pl. Cap.:

17. 1794. Schoenoxiphium lanceum (Thunb.) Kük. in

Engler (ed.), Pﬂanzenr. 38(IV, 20): 28. 1909. Kobresia

lancea (Thunb.) Koyama, J. Fac. Sci. Univ. Tokyo,

Sect. 3. Bot. 8: 80. 1961.

Carex ramosa Eckl. ex Kunth, Enum. Pl. 2: 531.

1837, nom. illeg.

Schoenoxiphium capense Nees, Linnaea 7: 533.

1832.

Schoenoxiphium meyerianum Kunth, Enum. Pl. 2:

530. 1837.

Schoenoxiphium sickmannianum Kunth, Enum. Pl.

2: 530. 1837.

Distribution: South Africa (Cape Province).

Carex ludwigii (Hochst.) Luceño & Martín-Bravo,

comb. nov.

Basionym: Schoenoxiphium ludwigii Hochst., Flora,

28: 764. 1845.

Schoenoxiphium rufum Nees in Linnaea 10: 201.

1836. Carex rufa (Nees) Baill., Hist. Pl. 12: 340. 1894,

nom. illeg., non Lam. (1779), non Schrank (1789).

Archaeocarex rufus (Nees) Fedde & J.Schust., Just’s

Bot. Jahresber. 41(2): 7. 1913 (publ. 1918). Kobresia

rufa (Nees) T.Koyama, J. Fac. Sci. Univ. Tokyo, Sect.

3, Bot. 8: 80. 1961.

Schoenoxiphium dregeanum Kunth, Enum. Pl. 2:

529. 1837; non Carex dregeana Kunth (1837).

MAKING CAREX MONOPHYLETIC 27

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Schoenoxiphium rufum var. pondoense Kük. in

Engler (ed.), Pﬂanzenr. 38(IV, 20): 31. 1909.

Schoenoxiphium burkei sensu Govaerts et al., World

Checklist of Cyperaceae (2007): 673, non C.B.Clarke.

Schoenoxiphium buchananii sensu Govaerts et al.,

World Checklist of Cyperaceae (2007): 673, non

C.B.Clarke.

Distribution: South Africa (Cape Province, Natal,

Northern provinces, Swazilandia), Lesotho.

Carex multispiculata Luceño & Martín-Bravo,

nom. nov.

Replaced synonym: Schoenoxiphium madagascariense

Cherm., Bull. Soc. Bot. France 70: 299. 1923; non

Carex madagascariensis Boeckeler (1884).

Distribution: South Africa (Natal, Northern prov-

inces), Madagascar.

Etymology: From the Latin multus, many, and

spicula, spikelet.

Carex perdensa (Kukkonen) Luceño &

Martín-Bravo, comb. nov.

Basionym: Schoenoxiphium perdensum Kukkonen,

Bot. Not. 131: 265. 1978.

Distribution: South Africa (Cape Province, Natal).

Carex pseudorufa Luceño & Martín-Bravo,

nom. nov.

Replaced synonym: Schoenoxiphium burttii Kuk-

konen, Notes Roy. Bot. Gard. Edinburgh 43: 365.

1986; non Carex burttii Noltie (1993).

Distribution: South Africa (Natal).

Etymology: From the Greek ψευδη

´ς(pseudo, resem-

bling but not equalling) and the Latin rufus,-a,-um

(red), alluding to the resemblance of this species to

Carex ludwigii, which was formerly known as Schoe-

noxiphium rufum.

Carex schimperiana Boeckeler, Linnaea 40: 373.

1876

Schoenoxiphium schimperianum (Boeckeler)

C.B.Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8:

67. 1908.

Carex densenervosa Chiov. in Ann. Bot. (Rome) 9:

149. 1911.

Schoenoxiphium bracteosum Kukkonen in Notes

Royal Botanic Garden Edinburgh 43: 365. 1986.

Distribution: Ethiopia to South Africa, Arabian

Peninsula.

Carex schweickerdtii (Merxm. & Podlech) Luceño

& Martín-Bravo, comb. nov.

Basionym: Schoenoxiphium schweickerdtii Merxm. &

Podlech, Mitt. Bot. Staatssamml. München 3: 529.

1960.

Distribution: South Africa (Natal).

Carex spartea Wahlenb., Kongl. Vetensk. Acad.

Nya Handl. 1803: 149. 1803

Schoenoxiphium sparteum (Wahlenb.) C.B. Clarke,

Bull. Misc. Inform. Kew, Addit. Ser. 8: 67. 1908.

Archaeocarex spartea (Wahlenb.) Pissjauk. Bot.

Mater. Gerb. Bot. Inst. Komarova Akad. Nauk

S.S.S.R. 12: 83. 1950. Kobresia spartea (Wahlenb.) T.

Koyama, J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 8: 80.

1961. Uncinia spartea (Wahlenb.) Spreng., Syst. Veg.

3: 830. 1826.

Schoenoxiphium kunthianum Kük. in Engler (ed.),

Pﬂanzenr. 38(IV, 20): 31. 1909. Archaeocarex kunthi-

ana (Kük.) Pissjauk., Bot. Mater. Gerb. Bot. Inst.

Komarova Akad. Nauk S.S.S.R. 12: 83. 1950. Kobresia

kunthiana (Kük.) T. Koyama, J. Fac. Sci. Univ. Tokyo,

Sect. 3, Bot. 8: 80. 1961.

Carex bolusii C.B.Clarke in Harvey & auct. suc.

(eds.), Fl. Cap. 7: 304. 1898.

Carex dregeana Kunth, Enum. Pl. 2: 511. 1837.

Carex dregeana var. major C.B.Clarke in Harvey &

auct. suc. (eds.), Fl. Cap. 7: 304. 1898.

Carex esenbeckiana Boeckeler, Linnaea 40: 372.

1876.

Carex indica Schkuhr, Beschr. Riedgräs. 1: 37.

1801, nom. illeg.

Schoenoxiphium caricoides C.B.Clarke, Bull. Misc.

Inform. Kew, Addit. Ser. 8: 67. 1908.

Schoenoxiphium caricoides var. major (C.B.Clarke)

C.B.Clarke, Bull. Misc. Inform. Kew, Addit. Ser. 8: 67.

1908.

Uncinia sprengelii Nees, Linnaea 10: 205. 1836,

nom illeg.; Carex sprengelii (Nees) Boeckeler, Linnaea

40: 371. 1876, nom. illeg.

Distribution: Uganda, Kenya, South Africa,

Madagascar.

Carex uhligii K.Schum. ex C.B.Clarke, Bot. Jahrb.

Syst. 38(2): 136. 1906

Replaced synonym: Schoenoxiphium lehmannii (Nees)

Kunth ex Steud., Syn. Pl. Glumac. 2: 245. 1855.

Uncinia lehmannii Nees, Linnaea 10: 206. 1836.

Schoenoxiphium sparteum var. lehmannii (Nees) Kük.

in Engler, Pﬂanzenr. 38(IV, 20): 32. 1909. Kobresia

28 GLOBAL CAREX GROUP

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lehmannii (Nees) T.Koyama, J. Fac. Sci. Univ. Tokyo,

Sect. 3, Bot. 8: 80. 1961; non Carex lehmannii Drejer

(1844).

Distribution: Ethiopia to South Africa.

Note: The correct name for this species if the

segregate genus is recognized is Schoenoxiphium

lehmannii.

TRANSFERS FROM UNCINIA PERS.TO CAREX L.

Carex aspericaulis (G.A.Wheeler) J.R.Starr, comb.

nov.

Basionym: Uncinia aspericaulis G.A.Wheeler, Dar-

winiana 45: 131. 2007.

Distribution: Juan Fernández Islands (Alejandro

Selkirk).

Carex astricta K.A.Ford, nom. nov.

Replaced synonym: Uncinia caespitosa Colenso ex

Boott in J.D.Hooker, Fl. Nov.-Zel. 1: 287. 1853; non

Carex caespitosa L. (1753).

Uncinia caespitosa var. collina Petrie, Trans. &

Proc. New Zealand Institute 52: 19. 1920.

Distribution: New Zealand (North & South Islands,

Stewart Island).

Etymology: From the Latin astrictus, drawn together

tight, referring to the densely caespitose habit of this

species.

Carex auceps (de Lange & Heenan) K.A.Ford,

comb. nov.

Basionym: Uncinia auceps de Lange & Heenan, Phy-

totaxa 104 (1): 12–20. 2013.

Distribution: New Zealand (Chatham Islands).

Carex aucklandica (Hamlin) K.A.Ford, comb.

nov.

Basionym: Uncinia aucklandica Hamlin, Domin.

Mus. Bull. 19: 63. 1959.

Distribution: New Zealand (South Island, Stewart

Island, Auckland Islands, Campbell Island).

Carex austrocompacta K.L.Wilson, nom. nov.

Replaced synonym: Uncinia compacta R.Br., Prodr. Fl.

Nov. Holl.: 241. 1810. Carex compacta (R.Br.) Poir. in

Lamarck, Encycl., Suppl. 3: 282. 1813, nom. illeg.;

non Carex compacta Lam. (1779).

Distribution: South-eastern Australia.

Etymology: The ﬁrst component of the name is from

the Latin australis, southern, referring to the South-

ern Hemisphere occurrence of this species, added to

Brown’s original epithet, which presumably refers to

the dense inﬂorescences of this species, to provide a

link between the new name and the original name.

Carex austroﬂaccida K.L.Wilson, nom. nov.

Replaced synonym: Uncinia ﬂaccida S.T.Blake, Proc.

Roy. Soc. Queensland 51: 49. 1939 (publ. 1940); non

Carex ﬂaccida Sw. ex Kunth (1837).

Uncinia tenella var. robustior Kük., Bot. Centralbl.

76: 211 (1898).

Distribution: South-eastern Australia.

Etymology: The ﬁrst component of the name is from

the Latin australis, southern, referring to the South-

ern Hemisphere occurrence of this species, added to

Blake’s original epithet, which refers to the soft-

textured leaves of this species, to provide a link

between the new name and the original name.

Carex austrosulcata K.L.Wilson, nom. nov.

Replaced synonym: Uncinia sulcata K.L.Wilson,

Telopea 5: 620. 1994; non Carex sulcata Schur (1858).

Distribution: South-eastern Australia.

Etymology: The ﬁrst component of the name is from

the Latin australis, southern, referring to the South-

ern Hemisphere occurrence of this species, added to

the original epithet, which refers to the rather chan-

nelled leaves of this species, to provide a link between

the new name and the original name.

Carex austrotenella K.L.Wilson, nom. nov.

Replaced synonym: Uncinia tenella R.Br., Prodr. Fl.

Nov. Holl.: 241. 1810.

Carex tenella Poir. in Lamarck, Encycl., Suppl. 3:

282. 1813, nom. illeg.; non Carex tenella Thuill.

(1790).

Distribution: South-eastern Australia.

Etymology: The ﬁrst component of the name is from

the Latin australis, southern, referring to the South-

ern Hemisphere occurrence of this species, added to

the original epithet, which presumably refers to the

small stature of this species, to provide a link

between the new name and the original name.

Carex banksiana K.A.Ford, nom. nov.

Replaced synonym: Uncinia banksii Boott in

J.D.Hooker, Fl. Nov.-Zel. 1: 287. 1853. Uncinia riparia

MAKING CAREX MONOPHYLETIC 29

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var. banksii (Boott) C.B.Clarke, J. Linn. Soc., Bot. 20:

392. 1883; non Carex banksii Boott (1846).

Uncinia capillaris Colenso, Trans. & Proc. New

Zealand Institute 20: 210. 1888; non Carex capillaris

L. (1753).

Distribution: New Zealand (North & South Islands).

Etymology: This name honours Sir Joseph Banks

(1743–1820), as did the previous epithet.

Carex brevicaulis Thouars, Esquisse Fl. Tristan

d’Acugna: 35. 1808

Uncinia brevicaulis (Thouars) Kunth, Enum. Pl. 2:

528. 1837.

Uncinia breviculmis Carmich., Trans. Linn. Soc.

London 12: 508. 1819.

Uncinia rigida Boeckeler, Flora 65: 64. 1882.

Uncinia brevicaulis var. rigida (Boeckeler) Kük. in

Engler (ed.), Pﬂanzenr. 38(IV, 20): 52. 1909.

Uncinia cylindrica Franch., Miss. Sci. Cap Horn 5:

379. 1889. Uncinia macloviana var. cylindrica

(Franch.) Kük., Bot. Centralbl. 76: 212. 1898.

Uncinia phleoides var. laticarpa Kük., Bot. Cen-

tralbl. 82: 130. 1900. Uncinia brevicaulis var. lati-

carpa (Kük.) Kük. in Engler (ed.), Pﬂanzenr. 38 (IV,

20): 52. 1909.

Uncinia brevicaulis f. montana Kük. in Engler (ed.),

Pﬂanzenr. 38 (IV, 20): 52. 1909.

Distribution: Hawaii (eastern Maui), Peru, southern

Chile to subantarctic islands.

Carex cheesemanniana (Boeckeler) K.A.Ford,

comb. nov.

Basionym: Uncinia cheesemanniana Boeckeler, Bot.

Jahrb. Syst. 5: 521. 1884.

Uncinia nervosa Boott in J.D.Hooker, Fl. Tasman.

2: 102. 1858. Uncinia compacta var. nervosa (Boott)

C.B.Clarke, J. Linn. Soc., Bot. 20: 395. 1883.

Distribution: South-eastern Australia, New Zealand

(South Island).

Note: The correct name for this species if the segre-

gate genus is recognized is Uncinia nervosa.

Carex corynoidea K.A.Ford, nom. nov.

Replaced synonym: Uncinia clavata (Kük.) Hamlin,

Domin. Mus. Bull. 19: 68. 1959. Uncinia australis var.

clavata Kük. in Cheeseman, Man. New Zealand Fl.:

802. 1906. Uncinia uncinata var. clavata (Kük.) Kük.

in Engler (ed.), Pﬂanzenr. 38 (IV, 20): 62. 1909; non

Carex clavata Thunb. (1794).

Distribution: New Zealand (North & South Islands).

Etymology: From the Greek koryne, club or mace,

Latinized as coryne, plus the adjectival suffix -oideus,

-a,-um, indicating resemblance of this species’ spikes

to small clubs.

Carex crispa K.A.Ford, nom. nov.

Replaced synonym: Uncinia involuta Hamlin, Domin.

Mus. Bull. 19: 49. 1959; non Carex × involuta (Bab.)

Syme (1870).

Distribution: New Zealand (North & South Islands,

Stewart Island).

Etymology: The name refers to the curled leaf tips in

this species.

Carex cyanea K.A.Ford, nom. nov.

Replaced synonym: Uncinia leptostachya Raoul, Ann.

Sci. Nat., Bot., III, 2: 116. 1844; non Carex leptos-

tachya Boiss. (1882).

Distribution: New Zealand (North & South Islands).

Etymology: The name refers to the bluish colour of the

leaves; from the Latin cyaneus, dark blue.

Carex dawsonii (Hamlin) K.L.Wilson, comb. nov.

Basionym: Uncinia dawsonii Hamlin, Trans. Roy. Soc.

New Zealand, Bot. 2: 128. 1963.

Distribution: New Caledonia.

Carex debilior (F.Muell.) K.L.Wilson, comb. nov.

Basionym: Uncinia debilior F.Muell., Fragm. 8: 151.

1874. Uncinia ﬁliformis var. debilior (F.Muell.)

W.R.B.Oliv., Trans. & Proc. New Zealand Institute 49:

128. 1917.

Distribution: Australia (Lord Howe Island).

Carex delacosta Kuntze, Revis. Gen. Pl. 3(2): 332.

1898

Replaced synonym: Uncinia macloviana Gaudich.,

Voy. Uranie: 412. 1829. Uncinia gracilis var. maclo-

viana (Gaudich.) C.B.Clarke, J. Linn. Soc., Bot. 20:

400. 1883. Uncinia brevicaulis var. macloviana

(Gaudich.) Kük. in Engler (ed.), Pﬂanzenr. 38(IV, 20):

52. 1909; non Carex macloviana d’Urv. (1826).

Uncinia montana Phil., Anales Univ. Chile 1865(2):

322. 1865. Uncinia macloviana var. montana (Phil.)

Kük., Bot. Centralbl. 82: 132. 1900; non Carex

montana L. (1753).

Uncinia delacosta Steud. in W.Lechler, Berberid.

Amer. Austral.: 52. 1857, nom. nud.

Distribution: Chile to southern Argentina.

30 GLOBAL CAREX GROUP

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Note: Uncinia delacosta Steudel is a nomen nudum

and therefore of no nomenclatural consequence.

Kuntze can therefore be interpreted under the provi-

sions of the Code (Art. 6.11 and Art. 58.1) as having

published a replacement name in Carex for Uncinia

macloviana Gaudich. There is no description or type

mentioned by Kuntze associated with his new name,

but he clearly indicated that he is publishing a

replacement name for U. macloviana, which is a valid

and legitimate name.

Carex dikei (Nelmes) K.L.Wilson, comb. nov.

Basionym: Uncinia dikei Nelmes, Kew Bull. 4: 377.

1949.

Distribution: South Africa (Marion Island, Prince

Edward Island).

Note: Nelmes published the epithet as ‘dykei’ in the

mistaken belief that Dyke was the surname of the

person concerned. Nelmes corrected the spelling to

dikei when he found out that the correct spelling of

the surname was Dike (Nelmes, 1949).

Carex dolichophylla J.R.Starr, nom. nov.

Replaced synonym: Uncinia macrophylla Steud., Syn.

Pl. Glumac. 2: 244. 1855; non Carex macrophylla

Hochst. ex Steud. (1855).

Uncinia bella Phil., Linnaea 30: 204. 1859; non

Carex bella L.H.Bailey (1892).

Uncinia phalaroides Boott ex C.B.Clarke, J. Linn.

Soc., Bot. 20: 396. 1883; non Carex phalaroides Kunth

(1837).

Uncinia bracteosa Phil., Anales Univ. Chile 93: 503.

1896; non Carex bracteosa Schwein. (1824).

Distribution: Southern Chile.

Etymology: The epithet combines the Greek word for

long (dolichos) with the Greek word for leaves (phylla)

to highlight the long leaves that typically surpass the

inﬂorescence of this rather large species.

Carex drucei (Hamlin) K.A.Ford, comb. nov.

Basionym: Uncinia drucei Hamlin, Domin. Mus. Bull.

19: 58. 1959.

Uncinia drucei var. payciﬂora Hamlin, Domin. Mus.

Bull. 19: 59. 1959.

Distribution: New Zealand (North & South Islands,

Stewart Island).

Carex ecuadorensis (G.A.Wheeler & Goetgh.)

J.R.Starr, comb. nov.

Basionym: Uncinia ecuadorensis G.A.Wheeler &

Goetgh., Aliso 15: 10. 1996 (publ. 1997).

Distribution: Northern and central Ecuador.

Carex edura K.A.Ford, nom. nov.

Replaced synonym: Uncinia divaricata Boott in

J.D.Hooker, Fl. Nov.-Zel. 1: 286. 1853. Uncinia com-

pacta var. divaricata (Boott) Hook.f., Handb. N. Zeal.

Fl. 1: 309. 1864; non Carex divaricata Kük. (1903).

Uncinia clarkei Petrie, Trans. & Proc. New Zealand

Institute 20: 185. 1888. Uncinia compacta var. clarkei

(Petrie) Cheeseman, Man. New Zealand Fl.: 800.

1906. non Carex clarkii E.W.Berry, Am. Nat. 39: 347

(1905).

Uncinia compacta var. Petriei C.B.Clarke in

Cheeseman, Man. New Zealand Fl.: 800. 1906.

Uncinia divaricata var. Petriei (C.B.Clarke) Hamlin,

Domin. Mus. Bull. 19: 57. 1959.

Distribution: Australia (Macquarie Island), New

Zealand (North & South Islands, Campbell Island).

Etymology: From the Latin edurus, or tough, referring

to the harsh environmental conditions this species

withstands.

Note: We consider that Art. 53.3. of the Code, which

states that confusingly similar names should be

treated as homonyms, applies here, making the

epithet clarkei unavailable in Carex because of the

publication of the fossil species Carex clarkii

E.W.Berry in 1905.

Carex egmontiana (Hamlin) K.A.Ford,

comb. nov.

Basionym: Uncinia egmontiana Hamlin, Domin. Mus.

Bull. 19: 33. 1959.

Uncinia silvestris var. squamata Hamlin, Domin.

Mus. Bull. 19: 28. 1959.

Distribution: New Zealand (North & South Islands,

Stewart Island).

Carex erebus K.A.Ford, nom. nov.

Replaced synonym: Uncinia hookeri Boott in

J.D.Hooker, Fl. Antarct.: 91. 1844. Uncinia riparia

var. hookeri (Boott) Kük. in Engler (ed.), Pﬂanzenr. 38

(IV, 20): 63. 1909; non Carex hookeri Kunth (1837).

Distribution: Australia (Macquarie Island), New

Zealand (Stewart Island, Antipodes Island, Auckland

Islands, Campbell Island).

Etymology: Named for the ship HMS Erebus on which

Joseph Dalton Hooker sailed on the Voyage to the

MAKING CAREX MONOPHYLETIC 31

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Antarctic 1839–1843, during which this species was

ﬁrst collected from the Auckland Islands.

Carex erinacea Cav., Icon. 5: 40. 1799

Uncinia erinacea (Cav.) Pers., Syn. Pl. 2: 534. 1807.

Agistron erinacea (Cav.) Raf., Good Book: 28. 1840.

Uncinia longifolia Kunth, Enum. Pl. 2: 527. 1837.

Uncinia longiaristata Steud., Syn. Pl. Glumac. 2:

243. 1855.

Uncinia philippii Hohen. ex Steud., Syn. Pl.

Glumac. 2: 243. 1855.

Uncinia macrotricha Franch., Miss. Sci. Cap Horn

5: 379. 1889.

Uncinia erinacea var. angustata Kük., Bot. Cen-

tralbl. 82: 101. 1900.

Distribution: Magellan Region of South America.

Carex erythrovaginata K.A.Ford, nom. nov.

Replaced synonym: Uncinia laxiﬂora Petrie, Trans. &

Proc. New Zealand Institute 17: 271. 1885; non Carex

laxiﬂora Lam. (1792).

Distribution: New Zealand (North & South Islands,

Stewart Island).

Etymology: The name refers to the reddish sheaths of

the leaves of this species.

Carex fernandesiana (Nees ex Boeckeler)

J.R.Starr, comb. nov.

Basionym: Uncinia fernandesiana Nees ex Boeckeler,

Linnaea 41: 347. 1877.

Uncinia douglasii Boott in J.D.Hooker, Fl. Antarct.

2: 369. 1846. Uncinia macloviana var. douglasii

(Boott) Kük., Bot. Centralbl. 82: 133. 1900.

Uncinia angusta Nees, Linnaea 9: 305. 1834, nom.

inval.

Uncinia angustata Boeckeler, Linnaea 41: 347.

1877.

Distribution: Juan Fernández Islands.

Note: The correct name for this species if the segre-

gate genus is recognized is Uncinia douglasii.

Carex ﬁrmula (Kük.) J.R. Starr, comb. &

stat. nov.

Basionym: Uncinia tenuis f. ﬁrmula Kük., Repert.

Spec. Nov. Regni Veg. 16: 433. 1920.

Uncinia tenuis Poepp. ex Kunth, Enum. Pl. 2: 525.

1837; non Carex tenuis Rudge (1804). Uncinia gracilis

Decne. in Dumont d’Urville, Voy. Pôle Sud, Atlas: t. 6,

f. B. 1843; non Carex gracilis Curtis (1782).

Distribution: Central America to Falkland Islands.

Note: The correct name for this species if the segre-

gate genus is recognized is Uncinia tenuis.

Carex goetghebeuri J.R.Starr, nom. nov.

Replaced synonym: Uncinia tenuifolia G.A.Wheeler &

Goetgh., Aliso 14: 144. 1995; non Carex tenuifolia

Poir. (1789).

Distribution: South-eastern Ecuador.

Etymology: The epithet honours the prominent cyper-

ologist Paul Goetghebeur of Ghent University (GENT,

Belgium) who described this species with Gerald A.

Wheeler (MIN, USA).

Carex hamata Sw., Prodr. Veg. Ind. Occ.: 18. 1788

Uncinia hamata (Sw.) Urb., Symb. Antill. 2: 169.

1900.

Carex jamaicensis Poir. in Lamarck, Encycl., Suppl.

3: 246. 1813.

Carex uncinata Schkuhr ex Steud., Nomencl. Bot.,

ed. 2, 1: 297. 1840.

Uncinia phleoides C.A.Mey., Bull. Acad. Roy. Sci.

Bruxelles 9(2): 249. 1842, nom. illeg.

Uncinia jamaicensis Liebm., Mexic. Neldeagt. Pl.,

V, 2: 272. 1851, nom. illeg.

Uncinia mexicana Steud., Syn. Pl. Glumac. 2: 243.

1855. Uncinia hamata var. mexicana (Steud.) Kük. in

Engler (ed.), Pﬂanzenr. 38 (IV, 20): 54. 1909.

Uncinia galeottii Boott ex C.B.Clarke, J. Linn. Soc.,

Bot. 20: 400. 1883.

Uncinia multifolia Boeckeler, Bot. Jahrb. Syst. 8:

207. 1887.

Uncinia hamata f. angustifolia Kük. in Engler (ed.),

Pﬂanzenr. 38 (IV, 20): 54. 1909.

Distribution: Mexico to Tropical America.

Carex hamlinii K.A.Ford, nom. nov.

Replaced synonym: Uncinia astonii Hamlin, Domin.

Mus. Bull. 19: 64. 1959; non Carex astonii Hamlin

(1968).

Distribution: New Zealand (North & South Islands).

Etymology: The epithet of the species is adopted to

recognize Bruce G. Hamlin (1929–1976) and his

important contributions to the ﬂora of New Zealand,

where this species is found.

32 GLOBAL CAREX GROUP

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Carex healyi K.A.Ford, nom. nov.

Replaced synonym: Uncinia scabra Colenso ex Boott

in J.D.Hooker, Fl. Nov.-Zel. 1: 285. 1853; non Carex

scabra Hoppe (1800).

Uncinia disticha Colenso, Trans. & Proc. New

Zealand Institute 20: 210. 1888; non Carex disticha

Huds. (1762).

Distribution: New Zealand (North & South Islands).

Etymology: The epithet of the species is adopted to

recognize Arthur J. Healy (1917–2011) and his impor-

tant contributions to the ﬂora of New Zealand.

Carex horizontalis (Colenso) K.A.Ford,

comb. nov.

Basionym: Uncinia horizontalis Colenso, Trans. &

Proc. New Zealand Institute 15: 334. 1883.

Uncinia rupestris Raoul, Ann. Sci. Nat., Bot., II, 2:

117. 1844; non Carex rupestris All. (1785).

Uncinia compacta var. viridis C.B.Clarke, J. Linn.

Soc., Bot. 20: 395. 1883. Uncinia caespitosa var.

viridis (C.B.Clarke) Hamlin, Domin. Mus. Bull. 19:

52. 1959. Uncinia viridis (C.B.Clarke) Edgar in Moore

& Edgar Fl. N. Zeal. 2: 229. 1970.

Uncinia compacta var. caespitiformis Kük. in

L.Cockayne, Rep. Bot. Surv. Stewart I.: 42. 1909.

Distribution: New Zealand (North & South Islands,

Chatham Islands, Stewart Island).

Note: The correct name for this species if the segre-

gate genus is recognized is Uncinia rupestris.

Carex imbecilla K.A.Ford, nom. nov.

Replaced synonym: Uncinia gracilenta Hamlin,

Domin. Mus. Bull. 19: 47. 1959; non Carex gracilenta

Boott ex Boeckeler (1877).

Distribution: New Zealand (North & South Islands,

Stewart Island).

Etymology: The name refers to the fragile habit of this

species.

Carex koyamae (Gómez-Laur.) J.R.Starr,

comb. nov.

Basionym: Uncinia koyamae Gómez-Laur., Brenesia

18: 92. 1980.

Distribution: Mexico (Chiapas), Costa Rica.

Carex laegaardii J.R.Starr, nom. nov.

Replaced synonym: Uncinia paludosa G.A.Wheeler &

Goetgh., Aliso 14: 142. 1995; non Carex paludosa

Gooden. (1794).

Distribution: North-eastern Colombia to Peru.

Etymology: The new name honours Simon Laegaard

(AAU, Denmark) who collected the holotype for this

and three other Carex spp. from northern South

America that were formerly treated in Uncinia

(Wheeler & Goetghebeur, 1995, 1997).

Carex lechleriana (Steud.) J.R.Starr, comb. nov.

Basionym: Uncinia lechleriana Steud., Syn. Pl.

Glumac. 2: 244. 1855.

Distribution: Chile to southern Argentina.

Carex lectissima K.A.Ford, nom. nov.

Replaced synonym: Uncinia ﬁliformis Colenso ex

Boott in J.D.Hooker, Fl. Nov.-Zel. 1: 286. 1853; non

Carex ﬁliformis L. (1753).

Uncinia rupestris var. capillacea Kük. in Engler

(ed.), Pﬂanzenr. 38 (IV, 20): 64. 1909.

Distribution: New Zealand (North & South Islands,

Stewart Island).

Etymology: From the superlative of the Latin adjec-

tive lectus, selected for the delicate ﬁne-leaved habit

of this species.

Carex longifructus (Kük.) K.A.Ford, comb. nov.

Basionym: Uncinia tenella var. longifructus Kük. in

Engler (ed.), Pﬂanzenr. 38 (IV, 20): 66. 1909. Uncinia

longifructus (Kük.) Petrie, Trans. & Proc. New

Zealand Institute 52: 17. 1920.

Distribution: New Zealand (North & South Islands).

Carex macloviformis (G.A.Wheeler) J.R.Starr,

comb. nov.

Basionym: Uncinia macloviformis G.A.Wheeler, Dar-

winiana 45: 136. 2007.

Distribution: Juan Fernández Islands (Alejandro

Selkirk).

Carex macrotrichoides J.R.Starr, nom. nov.

Replaced synonym: Uncinia chilensis G.A.Wheeler,

Aliso 15: 1. 1996 (publ. 1997); non Carex chilensis

Brongn. (1833).

Distribution: South-central Chile to Argentina (Rio

Negro).

Etymology: When described by Wheeler (1997b), this

species was only known from Chile, but it is now

documented from at least two localities in Argentina

(Starr, 2001; Wheeler, 2005). The new epithet com-

MAKING CAREX MONOPHYLETIC 33

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bines the Greek macros, long, with the Greek tri-

choides, hair-like, to highlight the extremely long

rachillae of this species, which are probably the

longest known in Carex (Wheeler, 1997b).

Carex madida J.R.Starr, nom. nov.

Replaced synonym: Uncinia lacustris G.A.Wheeler,

Aliso 14: 141. 1995; non Carex lacustris Willd. (1805).

Distribution: North-central Ecuador.

Etymology: The new epithet comes from the Latin

madidus for moist or wet, and it refers to the occur-

rence of this páramo species in humid habitats, such

as those at the margins of lakes (Wheeler &

Goetghebeur, 1995).

Carex megalepis K.A.Ford, nom. nov.

Replaced synonym: Uncinia ferruginea Boott in

J.D.Hooker, Fl. Nov.-Zel. 1: 288. 1853. Uncinia aus-

tralis var. ferruginea (Boott) C.B.Clarke in Cheese-

man, Man. New Zealand Fl.: 802. 1906. Uncinia

unciniata var. ferruginea (Boott) Kük. in Pﬂanzenr.

38: 62. 1909; non Carex ferruginea Scop. (1772).

Uncinia nigra Colenso, Trans. & Proc. New Zealand

Institute 17: 253. 1885; non Carex nigra (L.) Reichard

(1778).

Uncinia variegata Colenso, Trans. & Proc. New

Zealand Institute 20: 211. 1888; non Carex variegata

(All.) Lam. (1792).

Distribution: New Zealand (North & South Islands,

Stewart Island).

Etymology: This species has large glumes that are

much longer than the perigynia; from the Greek

mega-, big, and lepis,lepidos, a scale.

Carex meridensis (Steyerm.) J.R.Starr,

comb. nov.

Basionym: Uncinia meridensis Steyerm., Fieldiana,

Bot. 28(1): 61. 1951.

Uncinia macrolepis Decne. in Dumont d’Urville,

Voy. Pôle Sud 2: 13. 1853; non Carex macrolepis DC.

(1813).

Uncinia smithii Philcox, Kew Bull. 15: 229. 1961.

Distribution: North-western Venezuela to subantarc-

tic islands.

Note: The correct name for this species if the segre-

gate genus is recognized is Uncinia macrolepis.

Carex minor (Kük.) K.A.Ford, comb. & stat. nov.

Basionym: Uncinia caespitosa var. minor Kük. in

T.F.Cheeseman, Man. New Zealand Fl.: 802. 1906.

Uncinia angustifolia Hamlin, Domin. Mus. Bull. 19:

42. 1959.

Uncinia rupestris var. planifolia Kük. in Engler

(ed.), Pﬂanzenr. 38 (IV, 20): 64. 1909.

Distribution: New Zealand (North & South Islands,

Stewart Island).

Note: The correct name for this species if the segre-

gate genus is recognized is Uncinia angustifolia.

Carex multifaria (Nees ex Boott) J.R.Starr,

comb. nov.

Basionym: Uncinia multifaria Nees ex Boott in

J.D.Hooker, Fl. Antarct. 2: 369. 1846.

Uncinia macrostachya É.Desv. in C.Gay, Fl. Chil. 6:

229. 1854. Uncinia multifaria var. macrostachya

(É.Desv.) Kük., Bot. Centralbl. 82: 102. 1900.

Distribution: South-central & southern Chile.

Carex negeri (Kük.) J.R.Starr, comb. nov.

Basionym: Uncinia negeri Kük., Bot. Centralbl. 76:

210. 1898.

Uncinia negeri var. araucana Gunckel, Revista

Univ. (Santiago) 30: 58. 1945.

Distribution: Chile to south-western Argentina.

Carex nemoralis (K.L.Wilson) K.L.Wilson, comb.

nov.

Basionym: Uncinia nemoralis K.L.Wilson, Telopea 5:

620. 1994.

Distribution: South-eastern Australia.

Carex obtusifolia (Heenan) K.A.Ford, comb. nov.

Basionym: Uncinia obtusifolia Heenan, New Zealand

J. Bot. 34 (1): 11. 1996.

Distribution: New Zealand (North & South Islands,

Stewart Island).

Carex papualpina K.L.Wilson, nom. & stat. nov.

Replaced synonym: Uncinia compacta var. alpina

Noot., Blumea 24: 519. 1978; non Carex alpina

Schrank (1789).

Distribution: New Guinea (Mt Wilhelm, Mt Giluwe).

Etymology: The ﬁrst component of the epithet is

taken from an earlier name for this broad region,

34 GLOBAL CAREX GROUP

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Papua, added to the original epithet, referring to the

habitat of this taxon on the two highest mountains in

Papua New Guinea.

Carex parvispica K.A.Ford, nom. nov.

Replaced synonym: Uncinia sinclairii Boott in

J.D.Hooker, Handb. N. Zeal. Fl. 1: 309. 1864; non

Carex sinclairii Boott ex Cheeseman (1906).

Distribution: Eastern New Zealand (South Island);

also in south-eastern Australia (probably naturalized

there).

Etymology: The name refers to the small spikes found

in this species.

Carex penalpina K.A.Ford, nom. nov.

Replaced synonym: Uncinia fuscovaginata Kük., Bull.

Herb. Boissier, II, 4: 50 (1904). Uncinia purpurata

var. fuscovaginata (Kük.) Cheeseman, Man. New

Zealand Fl.: 801. 1906; non Carex fuscovaginata Kük.

(1904).

Uncinia fuscovaginata var. caespitans Hamlin,

Domin. Mus. Bull. 19: 21. 1959.

Distribution: New Zealand (North & South Islands,

Stewart Island).

Etymology: The name refers to this species being

often found in almost alpine tussock-grassland; from

the Latin paene or pene, nearly, and alpinus, alpine.

Carex perplexa (Heenan & de Lange) K.A.Ford,

comb. nov.

Basionym: Uncinia perplexa Heenan & de Lange,

New Zealand J. Bot. 39 (3): 376. 2001.

Distribution: New Zealand (North Island).

Carex phleoides Cav., Icon. 5: 40. 1799

Uncinia phleoides (Cav.) Pers., Syn. Pl. 2: 534. 1807.

Agistron phleoides (Cav.) Raf., Good Book: 28. 1840.

Uncinia trichocarpa C.A.Mey., Cyperac. Nov.: 11.

1831. Uncinia phleoides var. trichocarpa (C.A.Mey.)

C.B.Clarke, J. Linn. Soc., Bot. 20: 399. 1883.

Uncinia cumingii Nees, Linnaea 9: 305. 1834, nom.

inval.

Uncinia longifolia É.Desv. in C.Gay, Fl. Chil. 6: 226.

1854, nom. illeg.

Uncinia trichocarpa É.Desv. in C.Gay, Fl. Chil. 6:

227. 1854.

Uncinia durvillei Steud., Syn. Pl. Glumac. 2: 243.

1855.

Uncinia urvillei Steud., Syn. Pl. Glumac. 2: 243.

1855.

Uncinia longispica Boeckeler, Flora 41: 650. 1858.

Uncinia trichocarpa var. longispica (Boeckeler) Kük.,

Bot. Centralbl. 82: 131. 1900.

Uncinia montteana Phil., Linnaea 30: 205. 1859.

Uncinia chlorostachya Phil., Linnaea 33: 275. 1865.

Uncinia leptostachya Phil., Linnaea 33: 274. 1865.

Uncinia lasiocarpa Steud. ex Boeckeler, Linnaea

41: 349. 1877.

Uncinia longifolia Phil. ex C.B.Clarke, J. Linn.

Soc., Bot. 20: 399. 1883.

Uncinia phleoides var. nux-nigra C.B.Clarke, J.

Linn. Soc., Bot. 20: 399. 1883.

Uncinia phleoides f. longispica Franch., Miss. Sci.

Cap Horn 5: 378. 1889.

Uncinia loliacea Phil., Anales Univ. Chile 93: 503.

1896.

Uncinia phleoides var. brachytricha Speg., Revista

Fac. Agron. Univ. Nac. La Plata 3: 626. 1897.

Uncinia phleoides var. krausei Kük., Bot. Centralbl.

76: 211. 1898.

Distribution: Central Mexico, north-western Ven-

ezuela to southern South America.

Carex plurinervata J.R.Starr, nom. nov.

Replaced synonym: Uncinia costata Kük., Repert.

Spec. Nov. Regni Veg. 16: 433. 1920; non Carex costata

Schwein. (1824).

Distribution: Juan Fernández Islands (Alejandro

Selkirk).

Etymology: The epithet costata, ribbed, refers to the

many prominent veins on the perigynium of this

species known only from its type locality (Wheeler,

2007). The new epithet plurinervata combines the

Latin preﬁx pluri-, many, with nervata, nerved, to

convey the same meaning.

Carex potens K.A.Ford, nom. nov.

Replaced synonym: Uncinia affinis (Colenso ex

C.B.Clarke) Hamlin, Domin. Mus. Bull. 19: 30. 1959.

Uncinia riparia var. affinis Colenso ex C.B.Clarke, J.

Linn. Soc., Bot. 20: 392. 1883. non Carex affinis R.Br.

in J.Richardson, Bot. App: 750 (1823).

Uncinia purpurata var. subcaespitosa Kük. in

Engler (ed.), Pﬂanzenr. 38 (IV, 20): 61. 1909.

Distribution: New Zealand (North & South Islands).

Etymology: The name refers to the strong and harsh

habit of this species.

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Carex punicea K.A.Ford, nom. nov.

Replaced synonym: Uncinia rubra Colenso ex Boott in

J.D.Hooker, Fl. Nov.-Zel. 1: 287. 1853; non Carex

rubra H.Lév. & Vaniot (1909).

Uncinia rubra var. fallax Kük. in Engler (ed.),

Pﬂanzenr. 38 (IV, 20): 64. 1909.

Distribution: New Zealand (North & South Islands,

Stewart Island).

Etymology: The name refers to the red colour of the

whole plant.

Carex purpurata (Petrie) K.A.Ford, comb. nov.

Basionym: Uncinia purpurata Petrie, Trans. & Proc.

New Zealand Institute 17: 272. 1885.

Distribution: New Zealand (South Island).

Carex rapaensis (H.St.John) K.L.Wilson,

comb. nov.

Basionym: Uncinia rapaensis H.St.John, Nordic J.

Bot. 4: 60. 1984.

Distribution: Austral Islands (Rapa-Iti).

Carex ×rubrovaginata (Hamlin) K.A.Ford, comb.

nov.

Basionym: Uncinia ×rubrovaginata Hamlin, Domin.

Mus. Bull. 19: 24. 1959. U. fuscovaginata ×U. rubra.

Distribution: New Zealand (North Island).

Carex salticola J.R.Starr, nom. nov.

Replaced synonym: Uncinia andina G.A.Wheeler,

Hickenia 2: 218. 1997; non Carex andina Phil. (1896).

Distribution: South-central Chile to south-western

Argentina.

Etymology: The epithet refers to the fact that the

species grows in forests (saltus = forest;

-cola = dweller).

Carex scabrida J.R.Starr, nom. nov.

Replaced synonym: Uncinia scabriuscula

G.A.Wheeler, Hickenia 2: 215. 1997; non Carex scabri-

uscula Mack. (1908).

Distribution: Southern Chile to south-western

Argentina.

Etymology: The epithet for the new name means

somewhat scabrous in Latin, and is used to highlight

the characteristically scabrid culms of this species

(Wheeler, 1997a)

Carex sclerophylla (Nelmes) K.L.Wilson, comb.

nov.

Basionym: Uncinia sclerophylla Nelmes, Kew Bull. 4:

143 (1949).

Uncinia ohwiana T.Koyama, Bot. Mag. (Tokyo) 69:

214 (1956).

Distribution: New Guinea highlands.

Carex silvestris (Hamlin) K.A.Ford, comb. nov.

Basionym: Uncinia silvestris Hamlin, Domin. Mus.

Bull. 19: 26. 1959.

Distribution: New Zealand (North Island).

Carex strictissima (Kük.) K.A.Ford, comb. nov.

Basionym: Uncinia rubra var. strictissima Kük. in

Engler (ed.), Pﬂanzenr. 38 (IV, 20): 64. 1909. Uncinia

strictissima (Kük.) Petrie, Trans. & Proc. New

Zealand Institute 47: 55. 1915.

Uncinia rigida Petrie, Trans. & Proc. New Zealand

Institute 17: 271. 1885, nom. illeg.

Distribution: New Zealand (North & South Islands,

Antipodes Islands).

Carex subsacculata (G.A.Wheeler & Goetgh.)

J.R.Starr, comb. nov.

Basionym: Uncinia subsacculata G.A.Wheeler &

Goetgh., Aliso 14: 145. 1995.

Distribution: Ecuador (Pichincha).

Carex subtilis K.A.Ford, nom. nov.

Replaced synonym: Uncinia elegans (Kük.) Hamlin,

Domin. Mus. Bull. 19: 11. 1959. Uncinia sinclairii var.

elegans Kük. in Cheeseman, Man. New Zealand Fl.:

799. 1906. Uncinia macrolepis var. elegans (Kük.)

Kük. in Engler (ed.), Pﬂanzenr. 38 (IV, 20): 60. 1909;

non Carex elegans Willd. (1787).

Distribution: Australia (Tasmania); New Zealand

(South Island).

Etymology: The name refers to the delicate habit of

this species.

36 GLOBAL CAREX GROUP

The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, ••, ••–••

Carex subtrigona (Nelmes) K.L.Wilson,

comb. nov.

Basionym: Uncinia subtrigona Nelmes, Kew Bull. 4:

144 (1949).

Uncinia riparia var. stolonifera Kük. & Steen., Bull.

Jard. Bot. Buitenzorg, III, 13: 213 (1934).

Distribution: Borneo (Mt Kinabalu), Philippines (Mt

Apo), New Guinea highlands.

Carex subviridis K.A.Ford, nom. nov.

Replaced synonym: Uncinia distans Colenso ex Boott

in J.D.Hooker, Fl. Nov.-Zel. 1: 286. 1853; non Carex

distans L. (1759).

Uncinia nelmesii Hamlin, Trans. Roy. Soc. New

Zealand, Bot. 2: 127. 1963; non Carex nelmesii

H.E.Hess (1953).

Distribution: New Zealand (North Island).

Etymology: The name refers to the light green leaves

of this species.

Carex triangula J.R.Starr, nom. nov.

Replaced synonym: Uncinia triquetra Kük., Bot. Cen-

tralbl. 82: 97. 1900. Uncinia lechleriana var. triquetra

(Kük.) Kük. in Engler (ed.), Pﬂanzenr. 38 (IV, 20): 58.

1909; non Carex triquetra Boott (1846).

Distribution: Magellan Region of South America.

Etymology: The new epithet triangula has the same

meaning in Latin as triquetra (three-cornered).

Carex turbaria J.R.Starr, nom. nov.

Replaced synonym: Uncinia austroamericana

G.A.Wheeler, Darwiniana 43: 271. 2005; non Carex

austroamericana G.A.Wheeler (1986).

Distribution: Southern Chile to Tierra del Fuego.

Etymology: The epithet is derived from the Latin

turbarium for peat-bog and refers to the occurrence of

this species in persistently wet, base-poor sites, such

as Sphagnum bogs.

Carex umbricola K.L.Wilson, nom. nov.

Replaced synonym: Uncinia riparia R.Br., Prodr. Fl.

Nov. Holl.: 241 (1810). Carex riparia (R.Br.) Poir. in

Lamarck, Encycl., Suppl. 3: 282 (1813), nom. illeg.,

non Carex riparia Curtis (1783).

Distribution: South-eastern Australia.

Etymology: From the Latin umbra, shade, and -cola,

the Latin for a dweller, referring to the shady habitat

preferred by this species.

Carex uncinata L.f., Suppl. Pl.: 413. 1782

Uncinia uncinata (L.f.) Kük. in Engler (ed.),

Pﬂanzenr. 38 (IV, 20): 62. 1909.

Uncinia australis Pers., Syn. Pl. 2: 534. 1807, nom.

illeg.

Carex hamosa Thouars, Esquisse Fl. Tristan

d’Acugna: 35. 1808, nom. illeg.

Uncinia scaberrima Nees, Linnaea 9: 305. 1834,

nom. inval.

Uncinia lindleyana Kunth, Enum. Pl. 2: 526. 1837.

Uncinia rigidula Steud., Syn. Pl. Glumac. 2: 245.

1855.

Uncinia alopecuroides Colenso, Trans. & Proc. New

Zealand Institute 15: 335. 1883.

Uncinia bractata Colenso, Trans. & Proc. New

Zealand Institute 16: 341. 1884.

Uncinia pedicellata Kük. in Engler (ed.), Pﬂanzenr.

38 (IV, 20): 61. 1909. Uncinia uncinata var. pedicel-

lata (Kük.) Petrie, Trans. & Proc. New Zealand Insti-

tute 47: 54. 1915.

Uncinia uncinata var. laxior Carse, Trans. & Proc.

New Zealand Institute 48: 240. 1916.

Uncinia uncinata var. uliginosa Skottsb., Acta

Horti Gothob. 15: 328. 1944.

Distribution: New Zealand (North & South Islands,

Chatham Islands, Stewart Island, Auckland Islands),

Paciﬁc islands, New Caledonia, Hawaii.

Carex wheeleri J.R.Starr, nom. nov.

Replaced synonym: Uncinia araucana G.A.Wheeler,

Aliso 15: 3. 1996 (publ. 1997); non Carex araucana

Phil. (1896).

Distribution: South-central Chile (La Araucaria).

Etymology: The name honours Gerald A. Wheeler

(MIN, USA), who described this species and dozens of

others in a continuing series of signiﬁcant revisions of

the genus Carex (including Uncinia) in South

America.

Carex zotovii (Hamlin) K.A.Ford, comb. nov.

Basionym: Uncinia zotovii Hamlin, Domin. Mus. Bull.

19: 37. 1959.

Distribution: New Zealand (North & South Islands,

Stewart Island, Chatham Islands).

TRANSFER FROM VESICAREX STEYERM.TO CAREX L.

Carex collumanthus (Steyerm.) L.E.Mora, Acta

Biol. Colomb. 1: 40. 1982

Basionym: Vesicarex collumanthus Steyerm., Fieldi-

ana, Bot. 28(1): 63. 1951.

MAKING CAREX MONOPHYLETIC 37

The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, ••, ••–••

Distribution: Western South America to north-

western Venezuela.

INCERTAE SEDIS

Uncinia obtusata Colenso, Trans. & Proc. New

Zealand Institute 16: 341. 1884.

ACKNOWLEDGEMENTS

We are grateful to the John D. and Catherine T.

MacArthur Foundation for funding of the Biodiversity

Synthesis Group of the Encyclopedia of Life (EOL)

project, which funded our BioSynC Synthesis meeting

at the Field Museum in Chicago in September 2011,

when the Global Carex Group was formed. We also

thank the US National Science Foundation (NSF) for

funding our continuing international collaborative

work on the phylogeny and classiﬁcation of Carex

under grants DEB 1255901 to ALH and MJW, and

DEB 1256033 to EHR. We also acknowledge with

thanks funding for nomenclatural research and for

attendance at our second meeting during the Mono-

cots V conference in New York in July, 2013, from the

Natural Sciences and Engineering Research Council,

Canada (NSERC) to MJW and JRS; University of

Mainz to BG; JSPS KAKENHI Grant no. 25840136 to

OY; Korea National Arboretum to SK; CGL2012-

38744 project from the Spanish Ministry of Economy

and Competitiveness to ML; project 30870178 from

the National Natural Science Foundation of China to

SRZ, and a University of Wisconsin-Madison Raper

Travel Grant to DS. The ﬁgures were prepared with

invaluable technical advice from H. C. Rimmer.

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Genetic diversity and variations among Oklahoma and Texas sedge germplasm

Article

Full-text available

Mar 2025
CROP SCI

Sedges (Carex spp.) are herbaceous perennials with more than 2100 species recognized, offering significant potential for use in mowed turf, landscape gardens, and ecosystem restoration. Breeding and development of improved cultivars within various species could have a broad impact across these industries. The lack of information on genetic diversity and genetic variation within most relevant species has limited cultivar development. The objective of this study was to estimate the genetic diversity, population structure, and genetic variability of morphological and reproductive traits of 35 Carex plants collected from Oklahoma and Texas. A total of 20,689 single nucleotide polymorphic markers were used for genetic diversity characterization. Three subpopulations were identified from the germplasm panel that were largely determined by the species and partially by geographic locations. Substantial genetic diversity was observed within and among species. In addition, substantial genetic variation was found for morphological and reproductive traits and the reliability for those traits was moderate to high (𝑖2 = 0.38–0.83). The findings of this study provide insights into the genetic diversity of Carex germplasm from Oklahoma and Texas, offering essential knowledge for future Carex breeding efforts. Additionally, the study highlights the need for future research to understand the genetic mechanisms underlying reproductive traits and to develop reliable molecular tools for species differentiation.

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

Article

Full-text available

Jan 2025

Carex yangchunensis (Cyperaceae), a new species of Carex sect. Cryptostachyae in limestone regions of Guangdong, China, is described and illustrated. Both morphological 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.

Chitosan induced cold tolerance in Kobresia pygmaea by regulating photosynthesis, antioxidant performance, and chloroplast ultrastructure

Article

Full-text available

Nov 2024

Introduction Cold stress is the primary factor that limits the growth and development of Kobresia pygmaea in the Tibetan Plateau, China. Chitosan (CTS) has been recognized for its ability to enhance agricultural production and tolerance to stress. Methods This study examined the effect of treating seedlings under cold stress with chitosan. Results and Discussion The results demonstrated that cold stress inhibited the growth of seedlings and adversely affected the photosynthetic capacity [net photosynthetic rate (Pn), stomatal conductance (Gs), transpiration rate (Tr), maximum efficiency of photosystem II (Fv/Fm), quantum yield of photosystem II (φ PSII ), electron transport rate (ETR), and non-light-induced non-photochemical fluorescence quenching Y(NPQ)] and destroyed PSII and the chloroplast structure. Under regular temperatures, low concentrations of CTS (0.005% and 0.01%) inhibited the soluble protein content, ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco, EC 4.1.1.39) activity, and photosynthetic capacity. However, the application of 0.015% CTS increased the levels of soluble sugar, fructose, and protein, as well as those of the levels of ions, such as iron and magnesium, chlorophyll, photosynthetic capacity, and the activities of Rubisco, superoxide dismutase, and phenylalanine amino-lyase (PAL). Under cold stress, treatment with CTS decreased the contents of starch and sucrose; improved the contents of fructose, soluble protein, and antioxidants, such as ascorbic acid and glutathione; and enhanced the photosynthesis capacity and the activities of Rubisco, chitinase, and PAL. Exogenous CTS accelerated the development of the vascular bundle, mitigated the damage to chloroplast structure induced by cold, and promoted the formation of well-organized thylakoids and grana lamellae. Additionally, CTS upregulated the expression of genes related to cold tolerance in K. pygmaea, such as KpBSK2/KpERF/KpDRE326. These findings indicate that CTS enhances the cold tolerance in K. pygmaea by improving development of the vascular bundle, increasing the accumulation of solutes and antioxidants, regulating the transformation of carbohydrates, repairing the chloroplast structure, and maintaining the photosynthetic capacity and Rubisco activity.

Standing on the shoulders of giants: molecular data confirm Kükenthal’s systematic placement of the Australian endemic Carex archeri (Cyperaceae)

Article

Full-text available

Oct 2024

Archer’s sedge (Carex archeri Boott) is a small, rare (or possibly overlooked by collectors due to the diminutive size) species restricted to alpine and subalpine habitats in south-eastern mainland Australia and Tasmania. The systematic placement has been obscure with the species having been historically associated with sections in four of the six recognised Carex subgenera. We investigated the placement of C. archeri by addition to the available Carex molecular phylogenetic framework. Our results support C. archeri belonging to sect. Junciformes (in subg. Psyllophorae), making this the only representative of the subgenus in Australia. This placement was first proposed by Kükenthal (1909) who regarded C. archeri as a synonym of the New Zealand endemic C. acicularis Boott but our phylogenetic analyses support C. archeri as a separate taxon. Our approach highlights the utility of molecular barcoding for elucidating systematic relationships of poorly known taxa. Biogeographic reconstruction suggests Late Miocene dispersal from South America to the south-western Pacific but does not clarify whether New Zealand or Australia was colonised first. We evaluate the conservation status of Carex archeri using IUCN criteria as Endangered at the global level. At the state level, we propose Critically Endangered status in New South Wales, Endangered in Victoria and Data Deficient in Tasmania.

An upgraded key for identifying all native species, subspecies and varieties of the genus Carex (Cyperaceae) in Europe and the Caucasus

Article

Full-text available

Apr 2025
NORD J BOT

The last pan‐European key to Carex taxa was published in 1980 by Chater. Since that time several new species have been described, and numerous nomenclatural changes, including the recognition that the former genus Kobresia should be incorporated into Carex as C. subg. Euthyceras, have been made. This article provides a comprehensive key to identify all native European and Caucasian Carex taxa, plus two North American taxa, C. crawfordii and C. vulpinoidea , which are commonly regarded as being widely established in Europe. The key presented includes a general access key that directs the reader to one of five detailed keys (keys A–E), thus enabling the identification of all 316 European and Caucasian Carex taxa: 239 species, 63 subspecies and 14 varieties. Hybrids and introduced taxa, other than the two species mentioned above, are not included.

Southern Islands Vascular Flora (SIVFLORA) dataset: A global plant database from Southern Ocean islands

Article

Full-text available

Mar 2025

The Southern Islands Vascular Flora (SIVFLORA) dataset is a globally significant, open-access resource that compiles essential biodiversity data on vascular plants from islands across the Southern Ocean. The SIVFLORA dataset was generated through five steps: study area delimitation, compiling the dataset, validating and harmonizing taxonomy, structuring dataset attributes, and establishing file format and open access. Covering major taxonomic divisions, SIVFLORA offers a comprehensive overview of plant occurrences, comprising 14,589 records representing 886 species, 95 families, and 42 orders. This dataset documents that 58.62% of the taxa are native, 9.61% are endemic, and 31.77% are alien species. The Falkland/Malvinas Archipelago, the most species-rich, contrast sharply with less diverse islands like the South Orkney Archipelago. SIVFLORA serves as a taxonomically harmonized, interoperable resource for investigating plant diversity patterns, ecosystem responses to climate change in extreme environments, island biogeography, endemism, and the effects of anthropogenic pressures on Southern Ocean flora.

A taxonomic revision of New Zealand species of Carex section Inversae Kük. (Carex subgenus Vignea, Cyperaceae)

Article

Full-text available

Feb 2025
NEW ZEAL J BOT

Kerry A. Ford

This study revises an expanded Carex section Inversae for New Zealand. Eleven indigenous species are recognised, 10 of which are endemic. Notable features are the presence of unisexual spikes associated with gynodioecious and dioecious sexual systems, androgyny, and diandry. Based on results from morphometric analyses of Carex kirkii s.l., C. kirkii var. membranacea is raised to specific rank as C. ambita, C. pilifolia is described as new to science, and C. kirkii var. elatior is identified as a hybrid. Carex kirkii s.s. is found in alpine tussock grasslands and herbfield, C. ambita is a montane species of inland tarns, wetlands and tussock grasslands, and C. pilifolia is part of a rich turf kettlehole flora on moraine in the inter-montane basins of the eastern South Island. Both gynecandry and androgyny are present, with androgyny associated with diandry and found in all seven endemic androgynous species: C. ambita, C. kaloides, C. kirkii, C. muelleri, C. pilifolia, C. pterocarpa, and C. trachycarpa. Carex kaloides is the only species exhibiting an orange or reddish leaf coloration, a trait also observed in species of three other lineages of Carex in New Zealand. Typifications for three taxa are proposed: C. kirkii var. membranacea, C. pterocarpa, and C. inversa var. radicata. Updated descriptions, distributions, illustrations, habitat notes, and a diagnostic key are provided. Threat statuses are assessed for all species. Two species are reported as threatened, C. pilifolia is assessed as Nationally Critical, and C. ambita is assessed as Nationally Vulnerable, following the New Zealand Threat Classification System.

Comparative genomics reveals the molecular mechanisms of two drought-tolerant forages adapted to extreme environments in the northwest of China

Article

Dec 2024
IND CROP PROD

Chromosome number report of six Carex sect. Mitratae taxa from the Korean Peninsula (Cyperaceae)

Article

Full-text available

Sep 2024

Carex is the most species-rich group in the flowering plants in the temperate zones with more than 2,000 species worldwide. Coupled with the high species diversity, the genus possesses holocentric chromosomes, which miss constricted centromeres during cell divisions. Holocentric chromosomes are supposed to contribute to great variation in chromosome numbers by chromosome fragmentation (fission) and/or merging (fusion). Carex section Mitratae Kük. occurs mainly in East Asia and comprises about 80 taxa. Meiotic chromosome numbers of six taxa in the section were observed from 15 populations in South Korea: C. breviculmis R. Br. (n=32II, 33II, 34II), C. brevispicula G. H. Nam & G. Y. Chung (n=34II), C. candolleana H. Lév. & Vaniot (n=35II), C. conica Boott (n=18II, 19II), C. fibrillosa Franch. & Sav. (n=34II), and C. polyschoena H. Lév. & Vaniot (n=34II, 36II). All the chromosomes observed were very small (about 1–2 µm long) with diffuse centromeres (without constricted centromeres). Chromosome numbers vary in C. breviculmis, C. conica, and C. polyschoena. C. breviculmis and C. polyschoena show variation within a species (among populations), and C. conica exhibits two different numbers within a population. C. brevispicula, a Korean endemic, has a consistent chromosome number with previous counts, and C. fibrillosa shows the same chromosome number as the count from a Japanese population. For the first time, the chromosome number for C. candolleana is counted. Further investigations of chromosomes in Carex will help to understand the species diversity of the most species-rich group in flowering plants.

The genome assembly of Carex breviculmis provides evidence for its phylogenetic localization and environmental adaptation

Article

Jun 2024
ANN BOT-LONDON

Background and Aims Carex breviculmis is a perennial herb with good resistance and is widely used for forage production and turf management. It is important in ecology, environmental protection and biodiversity conservation, but faces several challenges due to human activities. However, the absence of genome sequences has limited basic research and the improvement of wild plants. Methods We annotated the genome of C. breviculmis and conducted a systematic analysis to explore its resistance to harsh environments. We also conducted a comparative analysis of Achnatherum splendens, which is similarly tolerant to harsh environments. Key Results The assembled the genome comprises 469.01 Mb, revealing 37 372 genes with a BUSCO completeness score of 99.0 %. The genome has 52.03 % repetitive sequences, primarily influenced by recent LTR insertions that have contributed to its expansion. Phylogenetic analysis suggested that C. breviculmis diverged from C. littledalei ~6.61 million years ago. Investigation of repetitive sequences and expanded gene families highlighted a rapid expansion of tandem duplicate genes, particularly in areas related to sugar metabolism, synthesis of various amino acids, and phenylpropanoid biosynthesis. Additionally, our analysis identified crucial genes involved in secondary metabolic pathways, such as glycolysis, phenylpropanoid biosynthesis and amino acid metabolism, which have undergone positive selection. We reconstructed the sucrose metabolic pathway and identified significant gene expansions, including 16 invertase, 9 sucrose phosphate synthase and 12 sucrose synthase genes associated with sucrose metabolism, which showed varying levels of expansion. Conclusions The expansion of these genes, coupled with subsequent positive selection, contributed to the ability of C. breviculmis to adapt to environmental stressors. This study lays the foundation for future research on the evolution of Carex plants, their environmental adaptations, and potential genetic breeding.