Larridon & al. • Names of subdivisions of Cyperus
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Nomenclature and typification of names of genera and subdivisions of
genera in Cypereae (Cyperaceae): 2. Names of subdivisions of Cyperus
Isabel Larridon,1 Wim Huygh,1 Marc Reynders,1 A. Muthama Muasya, 2 Rafaël Govaerts, 3
David A. Simpson3 & Paul Goetghebeur1
1 Ghent University, Department of Biology, Research Group Spermatophytes, K.L. Ledeganckstraat 35, 9000 Gent, Belgium
2 Botany Department, University of Cape Town, Rondebosch 7701, Cape Town, South Africa
3 Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, U.K.
Author for correspondence: Isabel Larridon, Isabel.Larridon@UGent.be
Abstract This paper is the second in a series of papers on the nomenclature of the names of genera (Huygh & al., 2010) and
subdivisions of genera in Cypereae (Cyperaceae). Here we present a survey of all published names of subdivisions of Cype
rus, designate types where needed, and evaluate priority and legitimacy. Cyperus is the largest genus in Cypereae and is the
second-largest genus in the family. The high diversity in Cyperus and the complex relationships with its segregate genera
resulted in the accumulation of more than 220 names of subdivisions in Cyperus of which 18 are not validly published and
33 are illegitimate.
Keywords Cyperaceae; Cyperus; subdivisions of genera; nomenclature; typification
INTRODUCTION
Cyperus is the second-largest genus in Cyperaceae and
is its most important genus in the tropics. It is the core taxon
of the Cypereae tribe. Recent molecular phylogenetic studies
of Cyperaceae (Muasya & al., 2002, 2009a; Simpson & al.,
2007), based on markers from the chloroplast genome, reveal
the paraphyletic character of Cyperus s.str. In the Cyperus
clade, several taxa are nested which have diverged from the
“typical” Cyperus vegetative, floral and anatomical characters. Spikelets with distichously arranged glumes have traditionally been the diagnostic character for Cyperus and its
closest allies. However, Muasya & al. (2002) conclude that
several taxa with spirally arranged glumes are present in the
Cyperus clade (e.g., Cyperus michelianus, Kyllingiella, Oxy
caryum). Species with distichously arranged glumes within
Ficinia and Isolepis (I. levynsiana) also occur, blurring the
generic delimitations of Cypereae genera. These findings
urge a recircumscription of the genera in this tribe (Muasya
& al., 2009b).
The Cyperus clade has a very complex generic and subdivisional nomenclature with approximately 350 generic and
subdivisional names to accommodate the roughly 950 species
present in the group. The subdivisional classification of Cype
rus alone comprises more than 220 names of which 7% are not
validly published and 12% are found to be illegitimate after
evaluation in accordance with the International Code of Bo
tanical Nomenclature (ICBN , McNeill & al., 2006). The lack
of overview and the limited availability of literature sources
in the past have led to the erroneous use of subdivisional
names with regards to valid publication and legitimacy in
almost all recent and past classifications of the group (e.g.,
Kükenthal, 1935–36; Chermezon, 1937; Kern, 1974; Haines
& Lye, 1983).
868
An overview of all published names of subdivisions of
Cyperus and the nomenclatural priority and typification of
each is a major necessity when creating a modern classification of the Cyperus clade based on the ongoing molecular and
morphological studies. This paper is part of our effort to provide the first complete nomenclatural survey of the names of
genera and subdivisions of genera in Cypereae. In this paper,
the second in a series of four papers, we present a survey of all
published names of subdivisions of Cyperus, designate types
where needed, and evaluate priority and legitimacy. A previous paper treated all names published at generic rank in the
Cyperus clade (Huygh & al., 2010). A third (Reynders & al.,
2011) and fourth paper will respectively present a synopsis of
all infrageneric names published in the segregate genera of
Cyperus, and a synopsis of all names published in taxa of the
Ficinia clade.
History of infrageneric taxonomy of Cyperus. — The morphological diversity and the presence of several convergent
evolutionary lines in Cyperus result in various controversial
classifications. Furthermore, there is no unanimity on the delimitation of the genus. Metcalfe (1971) already realised that to
study relationships within Cyperaceae using only morphological characters is problematic: “The main difficulty about the
classification of the Cyperaceae when the subject is approached
solely along traditional lines is that the flowers are very small,
the exact morphology of their parts is often obscure, and the
morphology of the inflorescences is difficult to interpret.” This
leads to uncertain homologies and conflicting interpretations.
The first infrageneric classification of the genus (and of
related genera which were later included in the genus) is that
of Nees (1834). He was the first to apply a sectional division
in the genus Cyperus, and created eight sections. Often, his
sections have incorrectly been assigned to Kunth (1837) as
stated by Väre & Kukkonen (2006). Bentham (1881) comments
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on Nees’s work that “he created some confusion, as well by
his usual tendency to raise species to the rank of genera, as
by a want of reference to the original papers or work where
Cyperaceae had been described, and by using a terminology
occasionally founded on mistaken views of the homology of
floral organs.” Most genera created by Nees in Cyperaceae are
no longer accepted as such.
Kunth (1837) greatly expanded our knowledge of Cyper
aceae. He classified the species of Cyperus with a trifid style
into 19 groups of unspecified rank, now usually treated at sectional level. He discussed the species of Cyperus with a bifid
style separately. Kunth (1837) accepted Mariscus and Kyllinga
as distinct genera. His main error was, according to Bentham
(1881), “a tendency to give as characters rather what in theory
we ought to see than what we actually do see, and, in his later
works, to describe specimens rather than species”.
Steudel (1854) divided Cyperus into three unnamed sections. The first section included species with a bifid style and
corresponds to the genus Pycreus. His “Sectio II”, split into 17
groups of unspecified rank, comprised the species of Cyperus
with a trifid style. Many of these groups are now treated at
sectional level. The third section held the species with uncertain
affinities. Furthermore, Steudel (1854) recognised Mariscus
and Kyllinga (“Kyllingia”) as segregate genera.
Böckeler (1868–77) studied Cyperaceae in the Berlin Herbarium. Like Steudel (1854), Böckeler (1868) treated Kyllinga
(“Kyllingia”) as a separate genus, and included Pycreus at sectional level in the genus Cyperus alongside a section “Eucype
rus”. However, he did not accept Mariscus at generic rank.
Böckeler (1868) divided his section “Eucyperus” into 20 groups
of unspecified rank, now usually treated at sectional level.
Clarke (1883, 1884, 1893, 1897, 1900, 1902, 1908), the
founder of modern cyperology, originally considered Cype
rus in a wide sense, but in his later publications he proposed
the subdivision of Cyperus s.l. into seven distinct genera, i.e.,
Cyperus, Courtoisina, Juncellus, Kyllinga, Mariscus, Pycreus,
and Torulinium.
The last complete revision of Cyperus was published in
the generic monograph of Kükenthal (1935–36). His infrageneric classification is the one most commonly used at present.
As in Suringar’s (1898) excellent revision of Cyperus in the
Malaysian Archipelago, the genus is taken in its wide sense.
Kükenthal (1935–36) divided Cyperus s.l. into six subgenera
(C. subgg. “Eucyperus”, Juncellus, Pycreus, Mariscus, Kyl
linga, and Torulinium), 61 sections and eight subsections. The
sections were primarily delineated by the nature of branching
of the compound inflorescence, extent of development of the
rhizomes, and the number of stamens and carpels per flower.
Obviously, many of Kükenthal’s sections need revision before
a natural classification of the genus can be proposed. Van der
Veken (1965) studied the embryos of 132 species of Cyperus
belonging in different subdivisions of the genus. The uniformity of the embryos appears to support the wide concept
of Cyperus. Furthermore, the study revealed the presence of
embryos of the Cyperus type in many taxa previously placed
near Scirpus (e.g., Ascolepis, Ficinia, Isolepis, Lipocarpha,
Kyllingiella, and Oxycaryum). Since Van der Veken’s (1965)
Larridon & al. • Names of subdivisions of Cyperus
publication, the inclusion of these genera in Cypereae led to a
more natural circumscription of that tribe. All of these genera
have, until now, been treated separately from Cyperus, except
for the treatment of Lipocarpha under Cyperus by Koyama
(1961). In his treatment of the genus, Kern (1974) only accepted
C. subg. Cyperus, Pycreus and Kyllinga at subgeneric rank,
as he considered Juncellus, Mariscus, and Torulinium too ill
defined to be recognised as subgenera.
Later publications are only regional studies of Cyperus
(e.g., Kern, 1974; Haines & Lye, 1983). Although they present
extensive knowledge of the genus, they are decidedly less valuable in assessing species relationships in a large genus such
as Cyperus, the members of which have successfully utilised
long-distance dispersal as well as more gradual short-distance
migration and in situ diversification.
Goetghebeur (1989) discussed the problems in the lectotypification of names relating to the infrageneric classification
of Cyperus. He specifically excluded all taxa with laterally
compressed nutlets, i.e. Kyllinga, Pycreus and Queenslandiella,
and several highly specialised taxa, i.e., Alinula, Courtoisina,
Kyllingiella, Oxycaryum, Remirea and Sphaerocyperus, from
Cyperus s.str. However, he included such segregates as Ano
sporum, Galilea, Juncellus, Mariscus, Sorostachys and Tor
ulinium. Goetghebeur (1998) retained this perspective in his
treatment of the family Cyperaceae.
Current views on the infrageneric classification of
Cyperus. — Cyperus s.str. is commonly divided into two units,
determined by the alternative character states of an anatomical
and of an inflorescence character set. The presence of Kranz
anatomy, correlated with C4 photosynthesis, has been used in
the classification of Cyperus since Rikli (1895). The vegetative anatomy is eucyperoid or chlorocyperoid, characterising
“C. subg. Eucyperus” (Grisebach, 1846) and “C. subg. Chloro
cyperus” (Schischkin, 1935). The inflorescence is composed
of digitately clustered spikelets and is often condensed, or the
inflorescence is an anthela composed of spikes of spikelets,
characterising C. subg. Pycnostachys and C. subg. Choris
tachys (Clarke, 1893). As demonstrated by Goetghebeur (1989),
the two classification systems are not completely congruent.
Either a few sections of subg. Chlorocyperus (with predominantly open inflorescence) exhibit a (partly) condensed and
depauperate inflorescence (Kükenthal, 1935–36), or a few sections of the predominantly eucyperoid subgenus Pycnostachys
reveal a chlorocyperoid type of anatomy (Druyts-Voets, 1970)
linked to C4 photosynthesis. Inevitably, this means that at least
one, or perhaps both of these presumed apomorphic character
states (chlorocyperoid anatomy, condensed inflorescence) has
or have evolved several times. This raises questions about the
switch from C3 to C4 anatomy either having evolved repeatedly
with no significant morphological divergence or only once
with subsequent convergence in the morphology of unrelated
species. Clearly these are problems that cannot be resolved
using morphological data alone. As already suggested by
Raynal (1973) and Goetghebeur (1989), Soros & Bruhl (2000)
confirmed that the chlorocyperoid anatomy type arose only
once within Cyperus. Soros & Bruhl (2000) stated that the C4
photosynthetic pathway arose at least four separate times in
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Larridon & al. • Names of subdivisions of Cyperus
Cyperaceae. However, in the tribe Cypereae only the chlorocyperoid anatomical variant occurs. Ongoing molecular research
suggests that the Cyperus C3 species form a paraphyletic group
with a monophyletic C4 subclade nested within the C3 group
(Muasya & al., 2002, 2009a; Simpson & al., 2007; Larridon &
al., in press). On the other hand, Goetghebeur (1989) remarked
that a multiple origin of the condensed inflorescence within Cy
perus is much more probable. There are many instances of species exhibiting both inflorescence types (open vs. condensed),
in Cyperaceae in general, and in Cyperus in particular.
Consequently, within Cyperus at least two main infrageneric groups have been recognised. An eucyperoid subgenus,
uniting plants without the Kranz syndrome, and with an inflorescence composed of digitately clustered spikelets; and a
chlorocyperoid subgenus, uniting plants with a chlorocyperoid
type of Kranz syndrome, and an inflorescence composed of
spikes of spikelets or condensed spikes. Within Cyperus, two
subgenera thus circumscribed were recognised by Chermezon (1937), Raynal (1973), Tucker (1983) and Hooper (1985).
However, Hooper (1985) also upheld C. subg. Juncellus, accommodating the chlorocyperoid species with a condensed
inflorescence and dorsiventrally flattened nutlets. Even more
subgenera were proposed by Haines & Lye (1983), who considered Cyperus in a broad sense, although in Lye’s earlier work
generic segregates such as Anosporum, Sorostachys (Lye,
1981b), Kyllinga (Lye, 1982), and Pycreus (Lye, 1981c) were
recognised. Goetghebeur (1989) remarked that he failed to see
the advantage of merging such easily recognisable taxa with
obvious synapomorphies, like Alinula, Ascolepis, Courtoisina,
Kyllinga, Kyllingiella, Lipocarpha, Oxycaryum, Pycreus and
Queenslandiella. On the other hand Lye (1992) convincingly
demonstrated the polyphyly of Mariscus within Cyperus. Most
authors now accept Mariscus as an element of Cyperus (e.g.,
Goetghebeur, 1998; Simpson & Koyama, 1998). Goetghebeur
(1989) proposed the recognition within Cyperus of only two
subgenera, respectively composed of the eucyperoid and of
the chlorocyperoid species, further arranged in sections. The
recent molecular studies (Muasya & al., 2002, 2009a; Simpson
& al., 2007; Larridon & al., in press) reveal the paraphyletic
character of Cyperus s.str. including several highly derived
lineages, urging a redefinition of generic delimitations in
Cypereae.
Materials and methods
The typifications were made after careful review of the
original descriptions of the taxa, examination of herbarium
specimens, and of live specimens in the Ghent University Botanical Garden. They are founded on expertise in Cyperaceae
obtained through long-standing research into the systematics
of this family by the Research Group Spermatophytes of the
Ghent University. References to Articles refer to the Interna
tional Code of Botanical Nomenclature (ICBN , McNeill &
al., 2006). References to accepted names refer to the World
Checklist of Cyperaceae (Govaerts & al., 2007, 2010). The classification used is that of Goetghebeur (1998).
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The type of each name of a subdivision of Cyperus is indicated. If in the protologue only one validly published species
name is cited or referred to (Art. 10.3 of the ICBN), the entry
appears simply as “Type: …” Where a type has been selected
later, the entry appears as “Lectotype” with a parenthetical
reference to the publication in which the selection was made.
Names that are homotypic, e.g., under Art. 7.5, are indicated
appropriately. If, when first published, the name of a subdivision of Cyperus had been assigned to a generic subdivision
of higher rank, that assignment is given in parentheses even
although it has no nomenclatural significance.
The large number of nomina nuda published by Chermezon are a consequence of the fact that Chermezon (1922, 1931)
did not provide descriptions for his intended new names, and
later Chermezon (1937) only provided a French diagnosis;
therefore his names are not validly published (Art. 32.1(d)
and 36.1).
Of the names of subdivisions of Cyperus with epithets that
are derived from the epithet of a constituent species and hence
typified under Art. 22.6 by the type of that species name, a
significant number, when published, included the already established type of a name of another generic subdivision of the
same rank, thereby making them superfluous and under a strict
reading of Art. 52, also illegitimate. As Art. 22.6 establishes
that such a name has a different type from that of the name that
ought to have been adopted, automatic typification under Art.
7.5 does not apply. The resultant situation is very unsatisfactory
in that perfectly appropriate epithets of subdivisions of Cyperus
based on the names of familiar species, are blocked from use
even when the taxon is circumscribed to exclude the type of
any earlier name at that rank. One might suppose by analogy
with superfluous names formed from legitimate basionyms
(Art. 52.3) that such names would be available for use when
the cause of the superfluity was removed, but a strict reading
of Art. 52.3 does not permit this. As it would seem logical that
the Code should extend the provisions of Art. 52.3 to such
cases, which are particularly important in large genera such as
Cyperus, we have noted the superfluous situation in the listings
below, but have refrained from adding the word “illegitimate”.
Nomenclature
Cyperus [unranked] Alati C.B. Clarke (see Cyperus L. sect.
Cyperus).
Cyperus sect. Alati Kük., nom. illeg. (see Kyllinga sect. Alatae
(Kük.) Tang & F.T. Wang; Reynders & al., 2011).
Cyperus (subg. Pycreus) sect. Albomarginati Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 359. 1936 – Type: Cyperus
albomarginatus (Mart. & Schrad. ex Nees) Steud. [=
Pycreus macrostachyos (Lam.) J. Raynal] (Art. 22.6).
This sectional epithet has not been combined in Pycreus.
Cyperus subg. Alinula (J. Raynal) Lye in Nordic J. Bot. 3: 230.
1983 ≡ Alinula J. Raynal in Adansonia, ser. 2, 17: 43. 1977
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Larridon & al. • Names of subdivisions of Cyperus
– Type: Cyperus lipocarphioides (Kük.) Lye [≡ Alinula
lipocarphioides (Kük.) J. Raynal (Art. 7.4)].
≡ Alinula J. Raynal (see Huygh & al., 2010).
Repentes Kunth are validly published under Art. 34.2. Of the
two names Jaubert & Spach (1844) selected the epithet of the
former for this section.
Cyperus sect. Allagostachyi Nees in Linnaea 9: 284. 1834 –
Lectotype (designated here): Cyperus hexastachyos Rottb.
[= C. rotundus L.].
Nees (1834) noted “ex affinitate Cyperi hexastachyi ” in his
description of C. sect. Allagostachyi. According to Kükenthal
(1936), however, Nees interpreted this species incorrectly, his
plant being C. zollingeri Steud.
Cyperus sect. Aristati Nees in Linnaea 9: 284. 1834 – Type:
Cyperus aristatus Rottb., nom. illeg. (Art. 52.1) (≡ Scirpus
intricatus L., non C. intricatus Schrad.) [= C. squarro
sus L.] (Art. 22.6).
Often, this section has been incorrectly assigned to Kunth
(1837) as stated by Väre & Kukkonen (2006).
Cyperus sect. Alopecuroidei Nees in Linnaea 9: 285. 1834 –
Type: Cyperus alopecuroides Rottb. (Art. 22.6).
Väre & Kukkonen (2006) remarked that this sectional
name has often been incorrectly assigned to Kunth (1837).
Cyperus sect. Alternifolii (Kunth) C.B. Clarke in J. Linn. Soc.,
Bot. 20: 288. 1883 ≡ C. [unranked] Alternifolii Kunth,
Enum. Pl. 2: 31. 1837 – Type: Cyperus alternifolius L.
(Art. 22.6).
Cyperus sect. Amabiles C.B. Clarke in Hooker, Fl. Brit. India
6: 598. 1893 – Type: Cyperus amabilis Vahl (Art. 22.6).
Clarke (1893) originally includes three species in this section, i.e., Cyperus amabilis, C. castaneus Willd., and C. cus
pidatus Kunth.
Cyperus (sect. Diffusi) [unranked] Angustifolii C.B. Clarke in
Bull. Misc. Inform. Kew, Addit. Ser. 8: 98. 1908 (‘Angusti
foliae’) – Lectotype (designated here): Cyperus derreilema
Steud.
We designate C. derreilema as lectotype since its characters (narrow leaves) fit very well the description of C. [unranked] Angustifolii. Kükenthal (1936) later organised the 11
species of this group of unspecified rank into the sections
Diffusi (4 spp.), Fusci (2 spp.), Haspani (1 sp.), and Incurvi
(4 spp.).
Cyperus subg. Anosporum (Nees) C.B. Clarke in J. Linn. Soc.,
Bot. 21: 34. 1884 ≡ Anosporum Nees in Linnaea 9: 287.
1834 ≡ C. sect. Anosporum (Nees) Pax in Engler, Nat.
Pflanzenfam. 2, 2: 107. 1887 – Type: Cyperus monocepha
lus Roxb. [= C. cephalotes Vahl].
Cyperus subsect. Apteri Nakai (see Cyperus sect. Compressi
Nees).
Cyperus sect. Arenarii (Kunth) Jaub. & Spach, Ill. Pl. Orient.
2, 11: t. 101. 1844 ≡ C. [unranked] Arenarii sive Repentes
Kunth, Enum. Pl. 2: 46. 1837 – Type: Cyperus arenarius
Retz. (Art. 22.6).
Väre & Kukkonen (2006) remark that as to the name of the
section, Kunth (1837) caused some confusion, because he simultaneously proposed two different names based on the same
type for the same taxon, viz. “Arenarii sive Repentes” both
the names C. [unranked] Arenarii Kunth and C. [unranked]
Cyperus subg. Aristomariscus Lye in Nordic J. Bot. 3: 213.
1983 ≡ C. (“sect. Eucyperus”) [unranked] Squarrosi
Benth., Fl. Austral. 7: 255. 1878 – Type: Cyperus squar
rosus L.
Bentham (1878) published C. [unranked] Squarrosi without
mention of rank. Lye (1983) published C. subg. Aristomariscus
based on the same type.
“Cyperus sect. Bernieriani Cherm.” in Humbert, Fl. Madagasc.
29: 104. 1937, nom. nud. (Art. 32.1(d) and 36.1).
This intended name was based on C. bernieri Cherm.
Cyperus sect. Bobartia (L.) C.B. Clarke in J. Linn. Soc., Bot.
21: 110. 1884 – Type: Bobartia indica L. (Art. 7.4).
≡ Bobartia L.
Clarke (1884) based C. sect. Bobartia on the monotypic genus Bobartia L. The only species described by Linnaeus (1753),
Bobartia indica L., has since been typified with a specimen
in the Hermann Herbarium (BM), which applies the name to
an iridaceous plant from southern Africa, traditionally placed
in the genus Bobartia (Strid, 1974; Väre & Kukkonen, 2006).
Although Väre & Kukkonen (2006) stated that this means that
the name of C. sect. Bobartia should be attributed to Clarke
(1884) alone, according to Art. 7.4 a new name formed from a
previously published legitimate name is, in all circumstances,
typified by the type of the basionym, even though it may have
been applied erroneously to a taxon now considered not to
include that type.
Clarke (1884) considered Bobartia indica L. to represent
C. pachyrrhizus, but later Clarke (1893) identified it with
C. arenarius Retz. (a species he explicitly excluded when he
established C. sect. Bobartia) when he established C. sect. Con
glomerati, nom. illeg., a section uniting the species C. arenar
ius, C. conglomeratus, and C. pachyrrhizus. Kükenthal (1936)
and Koyama (1985) cite Bobartia indica L. as a synonym of
C. arenarius.
Cyperus [unranked] Bracteati Boeck., nom. illeg. (see Cyperus
sect. Luzuloidei (Kunth) C.B. Clarke).
Cyperus sect. Brevifoliati C.B. Clarke, nom. illeg. (see Cyperus
sect. Corymbosus (Kunth) C.B. Clarke).
Cyperus subsect. Brevifolii Kük. (see Kyllinga sect. Brevifoliae
(Kük.) Kukkonen; Reynders & al., 2011).
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Larridon & al. • Names of subdivisions of Cyperus
Cyperus (subg. Mariscus) subsect. Brevistyli Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 522. 1936 – Lectotype (designated here): Cyperus cylindrostachyus Boeck.
Cyperus cylindrostachyus is the type of the name C. sect.
Umbellati, nom. illeg. in which C. subsect. Brevistyli is included.
Cyperus subg. Bulbocaules (C.B. Clarke) Lye in Nordic. J. Bot.
3: 230. 1983 (‘Bulbocaulis’) ≡ Mariscus subg. Bulbocaules
C.B. Clarke in Hooker, Fl. Brit. India 6: 619. 1893 ≡ M. sect.
Bulbocaules (C.B. Clarke) C.B. Clarke in Dyer, Fl. Cap. 7:
185. 1897 ≡ C. (subg. Mariscus) sect. Bulbocaules (C.B.
Clarke) Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 538.
1936 – Lectotype (designated here): Mariscus dregeanus
Kunth [= Cyperus dubius Rottb.].
Clarke (1893) originally included two species in this
subgenus, i.e., M. dregeanus and M. bulbosus C.B. Clarke,
nom. illeg., non Steud. [≡ C. clarkei T. Cooke]. Lye (1983)
was incorrect to indicate C. dubius as lectotype of his new
combination C. subg. Bulbocaules (C.B. Clarke) Lye because
Clarke (1893: 620) definitely excluded the type of C. dubius
from Mariscus dregeanus Kunth (1837: 120). Based on Lye’s
choice for C. dubius in 1983, we here correctly designate
Mariscus dregeanus as lectotype of the basionym M. subg.
Bulbocaules C.B. Clarke (1893) and by extension as type of
all names formed from this previously published legitimate
name (Art. 7.4). However, the currently accepted name in
Cyperus for Mariscus dregeanus is still Cyperus dubius
Rottb. (Govaerts & al., 2007, 2010). It is important to note
that Cyperus dregeanus Kunth (1837: 31) is different from
Mariscus dregeanus Kunth (1837: 120).
Clarke (1908) added 29 species (including M. bulbocaulis
Hochst. ex A. Rich. [= C. plateilema (Steud.) Kük.]) and published this taxon at sectional level. He divided C. sect. Bulbo
caules in two unnamed series differing in having either one or
more nutlets ripening per spikelet.
Cyperus subg. Bulbomariscus Lye in Nordic J. Bot. 3: 214.
1983 – Type: Cyperus amauropus Steud.
Cyperus sect. Bulbosi C.B. Clarke in Hooker, Fl. Brit. India
6: 611. 1893 – Type: Cyperus bulbosus Vahl (Art. 22.6).
Chermezon also lists a C. sect. Bulbosi in 1922 and 1937
without any author citation nor description and with C. escu
lentus L. as only species. It is unclear whether Chermezon
intended this as a new section, but in that case the name would
be a nomen nudum and would also not be validly published
(Art. 22.2), as C. esculentus was selected as type of Cyperus
(Britton, 1907).
Cyperus subg. Caespitosi Nakai (see Cyperus sect. Irioidei
Nees).
Cyperus (subg. Mariscus) subsect. Capenses Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 538. 1936 – Type: Cyperus
capensis (Steud.) Endl. (Art. 22.6).
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Cyperus [unranked] Capitati Boeck. (see Cyperus sect. Leuco
cephali Cherm. ex Kük.).
“Cyperus [unranked] Capitati multinervii Steud.”, Syn. Pl.
Glumac. 2: 27. 1854, nom. inval.
Steudel (1854) used two words to name this proposed subdivision. The Code does not directly address the use of such
names of subdivisions of genera. However, we conclude that
this name is not in accordance with Art. 21.2, which states
that the epithet of a subdivision of a genus is either of the same
form as a generic name, or a noun in the genitive plural, or a
plural adjective agreeing in gender with the generic name, but
not a noun in the genitive singular. In addition, Art. 20.3 states
that the name of a genus may not consist of two words, unless
these words are joined by a hyphen, and Steudel did not link
the words in this way.
Cyperus sect. Capituligeri Nees in Linnaea 9: 284. 1834 ≡
C. sect. Tenelli C.B. Clarke in Bull. Misc. Inform. Kew,
Addit. Ser. 8: 96. 1908 (‘Tenellae’) – Type: Cyperus tenel
lus L. f. [≡ Isolepis levynsiana Muasya & D.A. Simpson]
(lectotype C. sect. Capituligeri designated here – type
C. sect. Tenelli Art. 22.6).
We designate C. tenellus applicable to a species now included in Isolepis (Muasya & al., 2006, 2007), as lectotype of
this name. Nees (1834) included 29 species in C. sect. Capitu
ligeri based on the presence of a head-like inflorescence or
an umbel with head-like inflorescences. Although Nees mentioned C. tenellus in both his C. sect. Capituligeri and C. sect.
Allagostachyi, since it has head-like spikelet clusters it clearly
belongs in C. sect. Capituligeri.
Since Nees (1834), C. sect. Capituligeri has not been used
by any later author; his work from 1834 has been overlooked
by most authors as mentioned by Väre & Kukkonen (2006).
The combination in Isolepis has not been made.
Cyperus sect. Cephalotes J.V. Suringar, Cyperus: 76. 1898,
nom. superfl. – Type: Cyperus cephalotes Vahl (Art. 22.6).
= Cyperus sect. Anosporum (Nees) Pax.
When published, Cyperus sect. Cephalotes included the
type of C. sect. Anosporum (Nees) Pax (1887) and is therefore
superfluous.
“Cyperus subg. Chlorocyperus (Rikli) Schischk.” in Komarov,
Fl. URSS 3: 17. 1935, nom. inval. (Art. 22.2).
“Cyperus subg. Chlorocyperus” is not validly published
(Art. 22.2), as it included C. esculentus which was selected as
type of Cyperus (Britton, 1907). This proposed name was based
on: Chlorocyperus Rikli, Jahrb. Wiss. Bot. 27: 563. 1895, nom.
illeg. (see Huygh & al., 2010).
Cyperus subg. Choristachys C.B. Clarke, Cyperus sect. Chori
stachys (C.B. Clarke) C.B. Clarke (see Cyperus L. sect.
Cyperus).
Cyperus sect. Chrysanthi (C.B. Clarke) J. Kern (see Pycreus
sect. Chrysanthi C.B. Clarke; Reynders & al., 2011).
TAXON 60 (3) • June 2011: 868–884
“Cyperus sect. Compacti Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 31. 1922, nom. nud. (Art. 32.1(d) and
36.1).
This intended name was based on C. compactus Retz.
Cyperus sect. Compressi Nees in Linnaea 9: 284. 1834 ≡
C. (“sect. Eucyperus”) [unranked] Compressi (Nees)
Benth., Fl. Austral. 7: 255. 1878 ≡ C. [unranked] Sub
exalati C.B. Clarke in Oliver, Fl. Trop. Afr. 8: 313. 1901
(‘Subexalatae’) ≡ C. (sect. Pseudohaspani) subsect. Apteri
Nakai in Bot. Mag. (Tokyo) 26: 189. 1912 – Type: Cyperus
compressus L. (type C. sect. Compressi Art. 22.6 – type
C. subsect. Apteri – lectotype Subexalati designated here).
Väre & Kukkonen (2006) remarked that C. sect. Com
pressi has often been incorrectly assigned to Kunth (1837).
We designate C. compressus as lectotype of C. [unranked]
Subexalati as this species clearly has an unwinged rachilla.
Clarke (1901) created this group of unspecified rank, based
on his former C. subg. Choristachys series A from 1893, and
having a wingless or scarcely winged rachilla to differentiate
it from C. [unranked] Alati in which the rachilla is obviously
winged. Clarke (1901) divided C. [unranked] Subexalati into
three sections but named only the first section (i.e., C. sect.
Compressi). Cyperus subsect. Apteri Nakai has the same type
as C. sect. Compressi Nees (1834) because C. compressus is the
only species included by Nakai (1912), but C. subsect. Apteri is
legitimate with priority at the rank of subsection.
Cyperus sect. Congesti C.B. Clarke in J. Linn. Soc., Bot. 20:
285. 1883 – Lectotype (designated here): Cyperus exilis
Willd ex Kunth.
Originally, Clarke (1883) included five species in this section based on the presence of a simple to congested anthela:
C. dubius Rottb., C. obtusiflorus Vahl [= C. niveus var. leuco
cephalus (Kunth) Fosberg], C. exilis, C. stoloniferus Retz., and
C. pervillei Boeck [= Pycreus pervillei (Boeck.) C.B. Clarke].
Cyperus sect. Conglomerati C.B. Clarke in Hooker, Fl. Brit.
India 6: 602. 1893, nom. illeg. – Type: Cyperus conglo
meratus Rottb. (Art. 22.6).
= Cyperus sect. Arenarii (Kunth) Jaub. & Spach.
Cyperus sect. Conglomerati C.B. Clarke (1893) is illegitimate (Art. 52.1) since it includes the type of the older name
C. sect. Arenarii (Kunth) Jaub. & Spach (1844) (Art. 11.4).
Cyperus sect. Corymbosi (Kunth) C.B. Clarke in J. Linn.
Soc., Bot. 21: 153. 1884 ≡ C. [unranked] Corymbosi
Kunth, Enum. Pl. 2: 53. 1837 ≡ C. ([unranked] Corym
bosi) [unranked] Subaphylli Boeck. in Linnaea 36: 271.
1870 ≡ C. sect. Brevifoliati C.B. Clarke in Hooker, Fl. Brit.
India 6: 611. 1893 (‘Brevefoliati’), nom. illeg. (Art. 52.1) ≡
C. sect. Tunicati C.B. Clarke in Oliver, Fl. Trop. Afr. 8: 314.
1901 (‘Tunicatae’), nom. illeg. (Art. 52.1) – Type: Cyperus
corymbosus Rottb. (type of C. sect. Corymbosi Art. 22.6
– lectotype of C. [unranked] Subaphylli designated here –
type of C. sect. Brevifoliati and C. sect. Tunicati Art. 7.5).
= Cyperus L. sect. Cyperus (by lectotypification).
Larridon & al. • Names of subdivisions of Cyperus
Bentham (1878: 256) mentioned Kunth’s name Corymbosi
as a group of unspecified rank under C. sect. Papyrus.
Böckeler (1870) described C. [unranked] Subaphylli as a
subgroup under C. [unranked] Corymbosi characterised by
reduced leaves. Several authors published similar names of
unspecified rank for the same group of species. These names
were not validly published and might not have been intended as
names of subdivisions of Cyperus, i.e., “Aphylli ” (Kunth, 1837:
53), “Culmo aphyllo” (Steudel, 1854: 30), “Folia abbreviata
aut subnulla” (Clarke, 1883: 291) and “Sub-efoliati ” (Suringar,
1898: 121). All these authors also published names for an analogous group of species at unspecified rank characterised by
well-developed leaves, i.e., “Foliati ” (Kunth, 1837: 57), “Culmo
foliato” (Steudel, 1854: 32), “Foliati ” (Böckeler, 1870: 277),
“Foliati ” (Clarke, 1883: 292) and “Foliati ” (Suringar, 1898:
107). However, all these names were published without a diagnosis or description (Art. 32.1(d) and 36.1).
Cyperus sect. Brevifoliati C.B. Clarke (1893) and C. sect.
Tunicati C.B. Clarke (1901) are illegitimate (Art. 52.1) since
they include the type of the older name C. sect. Corymbosi
(Kunth) C.B. Clarke (1884).
Cyperus sect. Corymbosi Nakai, nom. illeg. (see Cyperus sect.
Irioidei Nees).
Cyperus subg. Courtoisia (Nees) Lye in Nordic J. Bot. 3: 230.
1983 ≡ Courtoisia Nees in Linnaea 9: 286. 1834, nom.
illeg. (Art. 53.1), non Marchand (1830) ≡ Cyperus subg.
Courtoisina (Soják) Lye in Lidia 3, 2: 52. 1992, nom. illeg.
(Art. 52.1) – Type: Cyperus pseudokyllingioides Kük. as
nomen novum of Courtoisia cyperoides (Roxb.) Nees (Kyl
linga cyperoides Roxb.) [≡ Courtoisina cyperoides (Roxb.)
Soják] (Art. 7.4).
≡ Courtoisina Soják (see Huygh & al., 2010).
Although the generic name Courtoisia Nees is an illegitimate later homonym, the subgeneric name Cyperus subg.
Courtoisia is legitimate with priority from its date of publication (1983) (Art. 58.1). Consequently, it is the correct name for
the taxon at subgeneric rank in Cyperus, although Courtoisina
Soják is the correct name at generic rank.
“Cyperus sect. Cuspidati Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 4. 1922, nom. nud. (Art. 32.1(d) and
36.1).
This intended name was based on C. cuspidatus Kunth.
Cyperus L. sect. Cyperus, autonym automatically established
by Nees in Linnaea 9: 284. 1834 (Art. 22.3) ≡ Cyperus L.,
Sp. Pl. 1: 44. 1753 (Huygh & al., 2010) ≡ Cyperus subg.
Cyperus, autonym automatically established in Petermann,
Deutschl. Fl. 11: 595. 1849 (Art. 22.3) ≡ C. subg. Pterocy
perus Peterm., Deutschl. Fl. 11: 595. 1849 ≡ Pterocyperus
(Peterm.) Opiz, Seznam: 80. 1852 ≡ C. sect. Solubiles C.B.
Clarke in J. Linn. Soc., Bot. 20: 291. 1883 ≡ C. subg. Chori
stachys C.B. Clarke in Hooker, Fl. Brit. India 6: 605. 1893 ≡
C. sect. Choristachys (C.B. Clarke) C.B. Clarke” in Urban,
Symb. Antill. 2: 22. 1900 (‘Choristachyae’) ≡ C. [unranked]
873
Larridon & al. • Names of subdivisions of Cyperus
Alati C.B. Clarke in Oliver, Fl. Trop. Afr. 8: 314. 1901 (‘Ala
tae’) ≡ Cyperus subsect. Cyperus, autonym automatically
established by Nakai in Bot. Mag. (Tokyo) 26: 188. 1912
(Art. 22.3) – Lectotype: Cyperus esculentus L. (lectotype
of Cyperus designated by Britton (1907) – lectotype of
autonyms Art. 7.6 – lectotypes of C. subg. Pterocyperus,
C. sect. Solubiles, C. subg. Choristachys, C. sect. Choris
tachys and C. [unranked] Alati designated here).
Because the lectotype of Cyperus was only designated
by Britton (1907), Art. 22.2 does not apply on the names of
subdivisions of Cyperus including C. esculentus published
before 1907. Kern (1974) was the first to effectively use the
autonyms of Cyperus at sectional and subgeneric rank. Väre
& Kukkonen (2006) erroneously indicated C. rotundus L. as
the type of C. sect. Cyperus. Five species were mentioned in
the original description of C. subg. Pterocyperus, Opiz (1852)
later established the genus Pterocyperus including only C. es
culentus (Huygh & al., 2010).
“Cyperus sect. Debilissimi Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 18. 1922, nom. nud. (Art. 32.1(d) and
36.1).
This intended name was based on C. debilissimus Baker.
Cyperus (subg. Mariscus) sect. Decidui Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 471. 1936 – Type: Cyperus
deciduus Boeck. (Art. 22.6).
“Cyperus sect. Denudati Cherm.” in Ann. Mus. Colon. Marseille,
ser. 3, 10(1): 12. 1922, nom. nud. (Art. 32.1(d) and 36.1).
This intended name was based on C. denudatus L. f.
Cyperus sect. Dichostylis (P. Beauv. ex T. Lestib.) Baill., Hist.
Pl. 12: 338. 1893 (see Dichostylis P. Beauv. ex T. Lestib.;
Huygh & al., 2010).
“Cyperus sect. Dichroostachyi Cherm.” in Ann. Mus. Colon.
Marseille, ser. 3, 10(1): 9. 1922, nom. nud. (Art. 32.1(d)
and 36.1).
This intended name was based on C. dichrostachyus
Hochst. ex A. Rich.
Cyperus sect. Diclidium (Schrad. ex Nees) Griseb., Fl. Brit.
W. I.: 565. 1864 ≡ Diclidium Schrad. ex Nees in Martius,
Fl. Bras. 2, 1: 51. 1842 ≡ C. subg. Diclidium (Schrad. ex
Nees) C.B. Clarke in J. Linn. Soc., Bot. 21: 191. 1884 ≡
Mariscus subg. Torulinium (Desv. ex Ham.) C.B. Clarke in
Hooker, Fl. Brit. India 6: 624. 1893 ≡ C. subg. Torulinium
(Desv. ex Ham.) Kük. in Engler, Pflanzenr. IV, 20 (Heft
101): 614. 1936 ≡ C. (subg. Torulinium) sect. Feraces Kük.
in Engler, Pflanzenr. IV, 20 (Heft 101): 614. 1936 – Type:
Cyperus ferax Rich. [= Cyperus odoratus L. subsp. odo
ratus] (lectotype of Diclidium and Torulinium designated
by Koyama (1961) – type of C. sect. Feraces Art. 22.6).
Although Torulinium Desv. ex Ham. has priority at generic
level (see Huygh & al., 2010), at subgeneric rank the epithet
Diclidium has priority.
874
TAXON 60 (3) • June 2011: 868–884
Cyperus sect. Difformes C.B. Clarke in J. Linn. Soc., Bot. 20:
289. 1883, nom. superfl. ≡ Cyperus (“sect. Eucyperus”)
[unranked] Nigricantes Benth., Fl. Austral. 7: 255. 1878
– Type: Cyperus difformis L. (type of C. sect. Difformes
Art. 22.6 – lectotype of C. [unranked] Nigricantes designated here).
When published, Cyperus sect. Difformes included the
type of C. sect. Irioidei Nees (1834) and is therefore superfluous. Bentham’s older name of unspecified rank does not have
priority (Art. 35.3). Although most of the epithets in Bentham’s
named subdivisions of Cyperus in Flora Australiensis appear
to be derived from species epithets and from the context some,
such as C. [unranked] Compressi (above) clearly are, there
seems no possibility that Bentham intended his C. [unranked]
Nigricantes to include either C. nigricans (L.) With. (1796) (≡
Schoenus nigricans L.) or C. nigricans Steud. (1842), nom.
illeg. (≡ Pycreus nigricans C.B. Clarke).
Cyperus sect. Diffusi (Kunth) C.B. Clarke in J. Linn. Soc., Bot.
21: 124. 1884 ≡ C. [unranked] Diffusi Kunth, Enum. Pl. 2:
25. 1837 ≡ C. subg. Pycnostachys C.B. Clarke in Hooker,
Fl. Brit. India 6: 597. 1893 ≡ C. sect. Pycnostachys (C.B.
Clarke) C.B. Clarke in Urban, Symb. Antill. 2: 22. 1900
(‘Pycnostachyae’) ≡ C. (Diffusi) [unranked] Eudiffusi C.B.
Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 98. 1908
(‘Eu-Diffusae’) – Type: Cyperus diffusus Vahl. (type of
C. sect. Diffusi Art. 22.6 – lectotype of C. subg. Pycno
stachys (≡ C. sect Pycnostachys) designated by Tucker
(1987) – type of C. [unranked] Eudiffusi Art. 22.6).
Clarke (1908) mistakenly describes two sections XII: the
Diffusi and the Eudiffusi, the latter containing the type, C. dif
fusus. Possibly, this error can be blamed on the posthumously
publishing of this work and Eudiffusi was meant as a group of
unspecified rank under sectional rank to contrast with C. [unranked] Simplices and C. [unranked] Angustifolii, which have
either a simple inflorescence or narrower leaves. Although the
epithet in this subdivision begins with the prefix Eu-, Art. 21.3
does not apply here.
Cyperus [unranked] Digitati J.V. Suringar, Cyperus: 133. 1898
– Type: Cyperus digitatus Roxb. (Art. 22.6).
Cyperus [unranked] Digitati includes the type of C. sect.
Exaltati (Kunth) C.B. Clarke (1883), which is the correct name
at sectional rank.
Cyperus sect. Distantes C.B. Clarke in J. Linn. Soc., Bot. 20:
290. 1883 ≡ C. sect. Longispicati J.V. Suringar, Cyperus:
123. 1898, nom. illeg. (Art. 52.1) ≡ C. [unranked] Distantes
(C.B. Clarke) J.V. Suringar, Cyperus: 123. 1898 – Type:
Cyperus distans L. f. (Art. 22.6).
Suringar (1898: 123) included C. sect. Distantes as a group
of unspecified rank in his new, but illegitimate C. sect. Long
ispicati in which he published two more groups of unspecified
rank based on the species C. digitatus Roxb. and C. pilosus Vahl.
Cyperus sect. Elegantes C.B. Clarke in J. Linn. Soc., Bot. 20:
288. 1883 – Type: Cyperus elegans L. (Art. 22.6).
TAXON 60 (3) • June 2011: 868–884
“Cyperus sect. Esculenti Kük.” in Engler, Pflanzenr. IV, 20
(Heft 101): 116. 1935, nom. inval. (Art. 22.2).
“Cyperus subg. Eucyperus Griseb.”, Spic. Fl. Rumel. 2: 420.
1846, nom. inval. (Art. 21.3) – “Cyperus sect. Eucyperus
(Griseb.) Boeck.” in Linnaea 35: 493. 1868, nom. inval.
(Art. 21.3).
Cyperus (Diffusi) [unranked] Eudiffusi C.B. Clarke (see Cype
rus sect. Diffusi (Kunth) C.B. Clarke).
Cyperus sect. Eukyllinga Kük. (see Kyllinga sect. Brevifoliae
(Kük.) Kukkonen; Reynders & al., 2011).
Cyperus [unranked] Eupycreus Boeck. (see Pycreus P. Beauv.
ser. Pycreus; Reynders & al., 2011)
Cyperus sect. Exaltati (Kunth) C.B. Clarke in J. Linn. Soc.,
Bot. 20: 293. 1883 ≡ C. [unranked] Exaltati Kunth, Enum.
Pl. 2: 70. 1837 ≡ C. subsect. Exaltati (Kunth) Nakai in
Bot. Mag. (Tokyo) 26: 192. 1912 – Type: Cyperus exaltatus
Retz. (Art. 22.6).
Bentham (1878: 257) adopted Kunth’s epithet Exaltati for
a group of unspecified rank under C. sect. Papyrus. Nakai
(1912) included C. subsect. Exaltati in his illegitimate C. sect.
Corymbosi Nakai.
“Cyperus sect. Exiles Cherm.” in Ann. Mus. Colon. Marseille,
ser. 3, 10(1): 33. 1922, nom. nud. (Art. 32.1(d) and 36.1).
This intended name was based on C. exilis Willd. ex Kunth.
Cyperus [unranked] Fasciculati Boeck. in Linnaea 36: 342.
1870 – Lectotype (designated here): Cyperus congestus
Vahl.
The characters of C. congestus fit very well the description
of this rankless group (e.g., “spiculae … elongatae lineares …
Rhacheola flexuosa compressa angusta, internodiis elongates
squamis decurrentibus alatis”). Although this name is validly
published under Art. 35.3, its epithet has not been adopted in
any ranked subdivision of Cyperus.
Cyperus sect. Fastigiati Kük. in Engler, Pflanzenr. IV, 20 (Heft
101): 53. 1935 – Type: Cyperus fastigiatus Rottb. (Art. 22.6).
Cyperus sect. Feraces Kük. (see Cyperus sect. Diclidium
(Schrad. ex Nees) Griseb.).
Cyperus (subg. Torulinium) sect. Filiformes Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 624. 1936 – Type: Cyperus
filiformis Sw. (Art. 22.6).
Cyperus subg. Fimbricyperus Lye in Nordic J. Bot. 1: 60. 1981
– Type: Cyperus subparadoxus Kük. [≡ Alinula paradoxa
(Cherm.) Goetgh. & Vorster].
Cyperus sect. Flabelliformes (C.B. Clarke) J. Kern in Fl.
Males., ser. 1, 7(3): 638. 1974 ≡ Mariscus sect. Flabelli
Larridon & al. • Names of subdivisions of Cyperus
formes C.B. Clarke in Hooker, Fl. Brit. India 6: 624. 1893
– Type: Mariscus microcephalus J. Presl & C. Presl. [=
Cyperus compactus Retz.].
Mariscus microcephalus was the only species included
in M. sect. Flabelliformes by Clarke (1893). Kern (1974) considers C. sect. Subulati C.B. Clarke as a synonym of C. sect.
Flabelliformes (C.B. Clarke) J. Kern and in consequence erroneously indicated C. subulatus R. Br. as the type of C. sect.
Flabelliformes (C.B. Clarke) J. Kern (Art. 7.4).
Cyperus sect. Flavescentes Kük. (see Pycreus ser. Zonati (C.B.
Clarke) C.B. Clarke; Reynders & al., 2011).
Cyperus (sect. Pseudohaspani) subsect. Flavidi Nakai in Bot.
Mag. (Tokyo) 26: 188. 1912 – Type: Cyperus flavidus Retz.
[= Pycreus flavidus (Retz.) T. Koyama] (Art. 22.6).
Nakai (1912) misinterpreted C. flavidus as being closely
related to C. haspan L. and C. tenuispica Steud.; he mentioned
the type of his C. sect. Pseudohaspani, i.e., C. pseudohaspan
Makino [= C. tenuispica Steud.] as a synonym under C. flavi
dus. This subsection has not been combined in Pycreus.
Cyperus (subg. Pycreus) sect. Fontinales Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 340. 1936 – Type: Cyperus
fontinalis (Cherm.) Kük. [≡ Pycreus fontinalis Cherm.]
(Art. 22.6).
This epithet has not been adopted for any ranked subdivision of Pycreus.
Cyperus (“sect. Eucyperus”) [unranked] Fulvi Benth., Fl.
Austral. 7: 255. 1878 – Type: Cyperus fulvus R. Br. (Art.
22.6).
Bentham (1878) published this name without mention of
rank. This epithet has not been adopted in any ranked subdivision of Cyperus.
Cyperus sect. Fusci (Kunth) C.B. Clarke in J. Linn. Soc., Bot.
21: 131. 1884 ≡ C. [unranked] Fusci Kunth, Enum. Pl. 2:
37. 1837 ≡ C. subsect. Fusci (Kunth) Nakai in Bot. Mag.
(Tokyo) 26: 190. 1912 – Type: Cyperus fuscus L. (Art.
22.6).
Steudel (1854) synonymised C. sect. Irioidei Nees (1834)
and C. [unranked] Fusci Kunth (1837) under the rankless group
“Iriae et fusci ”. Nakai (1912) included C. subsect. Fusci in his
illegitimate C. sect Corymbosi Nakai.
Cyperus sect. Galilea (Parl.) T.V. Egorova in Novosti Sist.
Vyssh. Rast. 34: 21. 2002 ≡ Galilea Parl., Fl. Palerm. 1:
297. 1845 (Huygh & al., 2010) – Type: Galilea mucronata
(L.) Parl. (Art. 7.4) [= Cyperus capitatus Vand.].
Cyperus sect. Globosi (C.B. Clarke) Kük. (see Pycreus sect.
Globosi C.B. Clarke; Reynders & al., 2011).
Cyperus sect. Glomerati (Kunth) C.B. Clarke in J. Linn. Soc.,
Bot. 21: 140. 1884, nom. superfl. ≡ C. [unranked] Glom
erati Kunth, Enum. Pl. 2: 75. 1837 ≡ Cyperus ([unranked]
875
Larridon & al. • Names of subdivisions of Cyperus
Leptolepides) [unranked] Umbellati Boeck. in Linnaea 35:
591. 1868 ≡ Cyperus subsect. Glomerati (Kunth) Nakai in
Bot. Mag. (Tokyo) 26: 191. 1912 – Type: Cyperus glomeratus
L. (type of C. [subdivisions] Glomerati Art. 22.6 – lectotype
of C. [unranked] Umbellati designated here).
= Cyperus sect. Distantes C.B. Clarke.
When published, Cyperus sect. Glomerati included the
type of C. sect. Distantes C.B. Clarke (1883) and is therefore
superfluous. The older name of unspecified rank does not have
priority (Art. 35.3).
Böckeler (1868) distinguishes C. [unranked] Umbellati
in his C. [unranked] Leptolepides and included four species:
C. semiradiciflorus Boeck. [= C. glomeratus L.], C. kappleri
Hochst. ex Steud., C. glomeratus L. and C. prolixus Kunth.
Nakai (1912) included C. subsect. Glomerati in his illegitimate C. sect. Corymbosi Nakai.
Cyperus sect. Graciles (Benth.) Kük., in Engler, Pflanzenr. IV,
20 (Heft 101): 202. 1936 ≡ Cyperus (“sect. Eucyperus”)
[unranked] Graciles Benth., Fl. Austral. 7: 254. 1878 –
Type: Cyperus gracilis R. Br. (Art. 22.6).
Cyperus sect. Graciles C.B. Clarke in Oliver, Fl. Trop. Afr.
8: 311. 1901, nom. illeg. (Art. 35.3, Art. 53.4), non C. [unranked] Graciles Benth. (1878) – Type: Cyperus cancel
latus Ridl. [= C. haspan L.].
= Cyperus sect. Haspani (Kunth) C.B. Clarke.
Cyperus sect. Haspani (Kunth) C.B. Clarke in J. Linn. Soc., Bot.
20: 287. 1883 ≡ C. [unranked] Haspani Kunth, Enum. Pl. 2:
34. 1837 – Type: Cyperus haspan L. (Art. 22.6).
Cyperus sect. Humiles (Kunth) J. Kern in Fl. Males., ser. 1, 7(3):
631. 1974, nom. superfl. ≡ C. [unranked] Humiles Kunth,
Enum. Pl. 2: 51. 1837 – Type: Cyperus humilis Kunth (Art.
22.6).
When published, Cyperus sect. Humiles included the type
of C. sect. Rupestres C.B. Clarke (1908) and is therefore superfluous. The older name of unspecified rank does not have
priority (Art. 35.3).
Kern (1974) considers C. sect. Rupestres as a synonym of
C. [unranked] Humiles and in so incorrectly indicated C. ru
pestris Kunth as the type species of C. sect. Humiles (Kunth)
J. Kern.
Cyperus sect. Hymenolepides Nees in Linnaea 9: 284. 1834 ≡
C. [unranked] Platystachyi Kunth, Enum. Pl. 2: 44. 1837 ≡
C. sect. Platystachyi (Kunth) C.B. Clarke in J. Linn. Soc.,
Bot. 21: 105. 1884 ≡ C. sect. Nivei C.B. Clarke in Hooker,
Fl. Brit. India 6: 601. 1893 – Type: Cyperus niveus Retz.
(lectotype of C. sect. Hymenolepides designated by Väre
& Kukkonen (2006) – lectotype of C. sect. Platystachyi
designated here – type of C. sect. Nivei Art. 22.6).
Although Kern (1974) lectotypified C. sect. Platystachyi
with C. leucocephalus Retz., Kunth (1837) did not include this
species in his C. [unranked] Platystachyi (but in his C. [unranked] Kyllingioides) and consequently it is not eligible as
876
TAXON 60 (3) • June 2011: 868–884
type. Instead we select C. niveus. Cyperus niveus was included
in the original circumscription of C. [unranked] Platystachyi
given by Kunth (1837), and its characters fit very well the
description of this group, in contrast to C. leucocephalus.
Simpson (1990) clearly states the differences between C. sect.
Leucocephali Cherm. ex Kük. and C. sect. Platystachyi. Kern’s
choice of type is based on Clarke (1884), who incorrectly
placed C. leucocephalus in C. sect. Platystachyi based on the
presence of C. sphaerocephalus var. leucocephalus Kunth
[= C. niveus var. leucocephalus (Kunth) Fosberg] in Kunth’s
Platystachyi. Furthermore, Kunth’s Platystachyi and C. sect.
Leucocephali form natural groups in Cyperus and following the subgeneric classification of Goetghebeur (1998) they
respectively belong in the C. subg. Cyperus (C4 photosynthesis—chlorocyperoid anatomy) and C. subg. Anosporum
(C3 photosynthesis—eucyperoid anatomy) and are in so not
closely related.
Cyperus sect. Incurvi Kük. in Engler, Pflanzenr. IV, 20 (Heft
101): 216. 1936 – Lectotype (designated here): Cyperus
disjunctus C.B. Clarke.
The characters of C. disjunctus fit very well the description
of this section; especially the curved glumes, dilated filaments
and well-developed connective of the anthers are clearly visible
in this species. Kükenthal (1936) also added an illustration of
this species showing the typical characteristics.
Cyperus sect. Irioidei Nees in Linnaea 9: 285. 1834 ≡ C. subg.
Caespitosi Nakai in Bot. Mag. (Tokyo) 26: 188. 1912 ≡
C. sect. Corymbosi Nakai in Bot. Mag. (Tokyo) 26: 190.
1912, nom. illeg. (Art. 53.4), non C. sect. Corymbosi
(Kunth) C.B. Clarke (1884) ≡ C. subsect. Irioidei (Nees)
Nakai in Bot. Mag. (Tokyo) 26: 191. 1912 (‘Iriae’) – Type:
Cyperus iria L. (type of C. [subdivisions] Irioidei Art.
22.6 – type of C. sect. Corymbosi Nakai, nom. illeg. Art.
7.5 – lectotype of C. subg. Caespitosi designated here).
Väre & Kukkonen (2006) remarked that this sectional
name has often been incorrectly assigned to Kunth (1837).
Kunth (1837: 38) published this name as C. [unranked] Iriae.
Steudel (1854) synonymised C. sect. Irioidei Nees (1834) and
C. [unranked] Fusci Kunth (1837) under the rankless group
Iriae et fusci.
Cyperus iria is the type of the name C. subsect. Irioidei
and C. sect. Corymbosi Nakai, nom. illeg. which are included
in C. subg. Caespitosi. Cyperus sect. Corymbosi Nakai (1912)
is illegitimate under Art. 52.1 since it includes the type of the
older name C. sect. Irioidei Nees (1834) (Art. 11.4). Nakai (1912)
included Nees’s Irioidei as a subsection in his illegitimate section Corymbosi. Like most authors he incorrectly assigned the
section based on C. iria to Kunth (1837).
Cyperus (subg. Pycreus) [unranked] Isodiametrici Kük. in
Engler, Pflanzenr. IV, 20 (Heft 101): 327. 1936 – Lectotype
(designated here): Cyperus polystachyos Rottb. [≡ Pycreus
polystachyos (Rottb.) P. Beauv.].
Cyperus polystachyos is the type of the name C. subg.
Pycreus (P. Beauv.) J. Carey in which C. [unranked] Isodia
TAXON 60 (3) • June 2011: 868–884
metrici is included. Kükenthal (1936) used this name at a level
between subgenus and section to replace Pycreus [unranked]
Puncticulati C.B. Clarke and P. subg. Reticulati C.B. Clarke
(both characterised by nutlets with isodiametric epidermal
cells). Although this name is validly published under Art. 35.3,
its epithet has not been adopted in any ranked subdivision of
Pycreus.
Cyperus sect. Ixiocarpi S.T. Blake in Proc. Roy. Soc. Queens
land 58: 37. 1947 – Type: Cyperus ixiocarpus F. Muell.
(Art. 22.6).
Cyperus sect. Juncellus Griseb., Fl. Brit. W. I.: 562. 1864 ≡
C. subg. Juncellus (Griseb.) C.B. Clarke in J. Linn. Soc.,
Bot. 20: 282. 1883 ≡ Juncellus (Griseb.) C.B. Clarke in
Hooker, Fl. Brit. India 6: 594. 1893 ≡ J. sect. Juncellus,
autonym automatically established by Nakai in Bot. Mag.
(Tokyo) 26: 199 1912 (Art. 22.3) – Type: Cyperus mu
cronatus Rottb. [= C. laevigatus subsp. distachyos (All.)
Maire & Weiller].
Cyperus subg. Kyllinga (Rottb.) J.V. Suringar, Cyperus (subg.
Kyllinga) sect. Kyllinga (Rottb.) J. Kern (see Kyllinga
Rottb. sect. Kyllinga; Reynders & al., 2011).
Cyperus (subg. Mariscus) subsect. Kyllingiformes Kük. in
Engler, Pflanzenr. IV, 20 (Heft 101): 553. 1936 – Lectotype
(designated here): Cyperus dubius Rottb.
The characters of C. dubius fit very well the description
of this subsection (“Anthela capitato contracta”).
Cyperus sect. Kyllingioides (Kunth) J. Kern in Fl. Males., ser.
1, 7(3): 643. 1974 ≡ C. [unranked] Kyllingioides Kunth,
Enum. Pl. 2: 94. 1837 – Type: Cyperus kyllingioides Vahl
[= C. dubius Rottb.] (Art. 22.6).
Cyperus (subg. Juncellus) sect. Laevigati Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 319. 1936 ≡ C. (sect. Py
creus) [unranked] Pseudopycreus Boeck. in Linnaea 35:
485. 1868 – Type: Cyperus laevigatus L. (type of C. sect.
Laevigati Art. 22.6 – lectotype of C. [unranked] Pseudopy
creus Boeck. designated here).
= Cyperus sect. Juncellus Griseb.
Cyperus [unranked] Pseudopycreus Boeck. (1868) is different from C. sect. Pseudopycreus (C.B. Clarke) Kük., nom.
illeg. (1936).
Cyperus sect. Lancei Kük., nom. superfl. (see Pycreus sect.
Lancei L.K. Dai; Reynders & al., 2011).
Cyperus sect. Latespicati Kük. (see Pycreus sect. Latespicati
(Kük.) L.K. Dai; Reynders & al., 2011).
Cyperus sect. Latifolii C.B. Clarke in Bull. Misc. Inform. Kew,
Addit. Ser. 8: 99. 1908 (‘Latifoliae’), nom. superfl. – Type:
Cyperus latifolius Poir. (Art. 22.6).
= Cyperus sect. Proceri (Kunth) C.B. Clarke.
Larridon & al. • Names of subdivisions of Cyperus
When published, Cyperus sect. Latifolii included the type
of C. sect. Proceri (Kunth) C.B. Clarke (1883) and is therefore
superfluous.
Cyperus (subg. Mariscus) sect. Laxiglumi (C.B. Clarke)
Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 457. 1936
≡ Mariscus (“sect. Eumariscus”) subsect. Laxiglumi
C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8:
103. 1908 (‘Laxiglumae’) – Lectotype (designated here):
Mariscus ischnos (Schltdl.) C.B. Clarke [≡ Cyperus
ischnos Schltdl.].
The characters of C. ischnos fit very well the original description of this subsection (Clarke, 1908).
Cyperus [unranked] Leptolepides Boeck. (see Cyperus sect.
Leucocephali Cherm. ex Kük.).
Cyperus sect. Leptostachyi (Boeck.) C.B. Clarke in J. Linn.
Soc., Bot. 20: 282. 1883 ≡ C. [unranked] Leptostachyi
Boeck. in Linnaea 36: 353. 1870 – Lectotype (designated
here): Cyperus nitidulus Boeck. [= C. odoratus L. subsp.
odoratus].
= Cyperus sect. Diclidium (Schrad. ex Nees) Griseb.
This name is not based on C. leptostachyus Griff. [= C. ro
tundus L.]. Böckeler included 19 species in this section; Kükenthal (1936) later divided them among nine of his sections
in the subgenera Mariscus and Torulinium. Clarke (1883)
published the name on sectional level including only C. ferax
Rich. [= C. odoratus L. subsp. odoratus]. Böckeler (1870: 353)
included this species in C. [unranked] Diclidium, but included
C. nitidulus, a synonym of C. ferax in C. [unranked] Lepto
stachyi. Cyperus sect. Diclidium (Schrad. ex Nees) Griseb.
had already been published in 1864 with C. odoratus as type.
Clarke (1883) clearly remarked on the relationship of the latter
species with C. ferax, and definitely considered both taxa as
belonging to the same section, writing: “C. odoratus, Vahl; …
paullo differt spiculis 14–21-floris, sed mihi varietas videtur”.
Cyperus sect. Leucocephali Cherm. ex Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 276. 1936 ≡ C. [unranked]
Leptolepides Boeck. in Linnaea 35: 588. 1868 ≡ C. ([unranked] Leptolepides) [unranked] Capitati Boeck. in Linnaea 35: 588. 1868 – Type: Cyperus leucocephalus Retz.
(type of C. sect. Leucocephali Art. 22.6 – lectotype of
C. [unranked] Leptolepides and C. [unranked] Capitati
designated here).
Cyperus sect. Leucocephali was first published as a nomen nudum by Chermezon (1931), Kükenthal (1936) added a
valid description. The characters of C. leucocephalus fit very
well the description of C. [unranked] Leptolepides (e.g., “Inflorescentia saepissime capitato-contracta”). Böckeler (1868)
distinguished two lower groups in C. [unranked] Leptolepides,
i.e., C. [unranked] Capitati (incl. C. leucocephalus) and C. [unranked] Umbellati. Although these names are validly published
under Art. 35.3, they do not have priority. Consequently, the
correct name of this taxon at sectional level is C. sect. Leuco
cephali Cherm. ex Kük. (1936) (Art. 11.4).
877
Larridon & al. • Names of subdivisions of Cyperus
Cyperus sect. Limosi T.V. Egorova in Novosti Sist. Vyssh. Rast.
34: 27. 2002 – Type: Cyperus limosus Maxim. (Art. 22.6).
Cyperus subg. Lipocarpha (R. Br.) T. Koyama in Quart. J.
Taiwan Mus. 14: 163. 1961 ≡ Lipocarpha R. Br. in Tuckey,
Narr. Exped. Zaire: 459. 1818 – Type: Cyperus lipocarpha
T. Koyama [= Lipocarpha chinensis (Osbeck) J. Kern] (Art.
7.4).
≡ Lipocarpha R. Br. (see Huygh & al., 2010).
Cyperus sect. Longispicati J.V. Suringar, nom. illeg. (see Cype
rus sect. Distantes C.B. Clarke).
Cyperus (subg. Mariscus) subsect. Longistyli Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 507. 1936 – Lectotype (designated here): Cyperus cyperoides (L.) Kuntze.
We designate C. cyperoides as lectotype of this subsection
as it is a common species and it fits very well to the description
given by Kükenthal (1936).
Cyperus (sect. Papyrus) [unranked] Lucidi Benth., Fl. Austral.
7: 256. 1878 – Type: Cyperus lucidus R. Br. (Art. 22.6).
Bentham (1878) published this name without mention of
rank. This epithet has not been adopted in any ranked subdivision of Cyperus.
Cyperus sect. Luzuloidei (Kunth) C.B. Clarke in J. Linn. Soc.,
Bot. 21: 115. 1884 ≡ C. [unranked] Luzuloidei Kunth,
Enum. Pl. 2: 39. 1837 (‘Luzuloideae’) ≡ C. [unranked]
Bracteati Boeck. in Linnaea 35: 549. 1868, nom. illeg. (Art.
52.1) – Type: Cyperus luzulae (L.) Retz. (type of C. sect.
Luzuloidei Art. 22.6 – type of C. [unranked] Bracteati
Art. 7.5).
Kunth (1837) was the first to describe this group (as C. [unranked] Luzuloidei). Böckeler (1868) referred to the same group
as C. [unranked] “Bracteati s. Luzuloidei ” but excluding several
species. According to Art. 34.2 both of Böckeler’s names are
validly published, but C. [unranked] Bracteati Boeck. is nomenclaturally superfluous and illegitimate (Art. 52.1).
Cyperus sect. Mariscoidei Nees in Linnaea 9: 285. 1834 –
Type: Cyperus mariscoides Nutt., nom. illeg. [= C. fili
culmis Vahl] (Art. 22.6).
Steudel (1854) incorrectly assigned this section to Kunth
(1837: 84 as ‘Cyperus [unranked] Mariscoides’).
Cyperus subg. Marisculus (Goetgh.) Lye in Lidia 3, 4: 132.
1994 ≡ Marisculus Goetgh., Bull. Jard. Bot. Natl. Belg.
47: 444. 1977 (Huygh & al., 2010) – Type: Alinula peteri
(Kük.) Goetgh. & Vorster (Ascolepis peteri Kük.) (Art. 7.4)
[≡ Cyperus microaureus Lye].
= Alinula J. Raynal (see Huygh & al., 2010).
Cyperus sect. Mariscus (Vahl) Griseb., Fl. Brit. W. I.: 566. 1864
≡ Mariscus Vahl, Enum. Pl. 2: 372. 1805, nom. cons. ≡ C.
subg. Mariscus (Vahl) C.B. Clarke in J. Linn. Soc., Bot.
20: 296. 1883 ≡ M. subg. Mariscus, autonym automatically
878
TAXON 60 (3) • June 2011: 868–884
established by C.B. Clarke in Hooker, Fl. Brit. India 6: 619.
1893 (Art. 22.3) ≡ M. sect. Mariscus, autonym automatically established by C.B. Clarke in Hooker, Fl. Brit. India 6:
620. 1893 (Art. 22.3) ≡ M. ser. Mariscus, autonym automatically established by C.B. Clarke in Urban, Symb. Antill.
2: 39. 1900 (Art. 22.3) ≡ M. subsect. Mariscus, autonym
automatically established by C.B. Clarke in Bull. Misc.
Inform. Kew, Addit. Ser. 8: 103. 1908 (Art. 22.3) – Typ.
cons.: Mariscus capillaris (Sw.) Vahl (Schoenus capillaris
Sw.) [= Cyperus nanus Willd. var. nanus] (Art. 7.4).
Endlicher (1836: 119) was the first to use Mariscus as an infrageneric taxon of Cyperus as C. [unranked] Mariscus (Vahl)
Endl. In this group, Böckeler (1870: 372) distinguished several
unnamed subgroups based on the inflorescence morphology
from a single capitate inflorescence (group a), suborbiculate
spikes (b), an umbel with sessile spikes (group b1), pedunculated spikes (b2), cylindrical spikes (c), simple umbel (c1),
compound with sessile umbels (c2) and compound umbel with
stalked umbels (c3).
Cyperus subg. Micheliani (C.B. Clarke) Kukkonen in Ann.
Bot. Fenn. 32, 3: 161. 1995 ≡ Scirpus sect. Micheliani C.B.
Clarke in Hooker, Fl. Brit. India 6: 662. 1893 – Type: Scir
pus michelianus L. (Art. 22.6) [≡ Cyperus michelianus (L.)
Delile subsp. michelianus].
Clarke (1893) described only two species in S. sect.
Micheliani, S. michelianus and S. kyllingioides (A. Rich.)
Boeck. [= Kyllingiella microcephala (Steud.) R.W. Haines &
Lye] the type of the genus Kyllingiella. In 1902, Clarke added a
third species S. steudneri Boeck. [= Kyllinga ambigua Steud.].
Cyperus subg. Micromariscus Lye in Nordic J. Bot. 3: 215.
1983 – Type: Cyperus micromariscus Lye (Art. 22.6).
Cyperus (subg. Juncellus) sect. Minuti Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 325. 1936 – Type: Cyperus
minutus (C.B. Clarke) Kük, nom. illeg., non J.F. Gmel.
(Art. 22.6).
The type specimen of C. minutus, nom. illeg. clearly shows
affinities to C. meeboldii Kük. and relatives which are treated
by Kükenthal (1936) in C. sect. Rupestres which is characterised by a tendency towards monostigmatic gynoecia, distinctive obovoid nutlets and swollen culm bases. The exact status
of C. minutus, nom. illeg. is unclear and needs further attention.
Cyperus sect. Monocephali C.B. Clarke in Oliver, Fl. Trop.
Afr. 8: 310. 1901 (‘Monocephalae’) ≡ C. sect. Rupestres
C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8:
96. 1908 – Type: Cyperus rupestris Kunth (lectotype of
C. sect. Monocephali designated here – type of C. sect.
Rupestres Art. 22.6).
The characters of C. rupestris fit very well the description
of C. sect. Monocephali, especially in having a constant single
head of spikelets.
Cyperus (subg. Pycreus) sect. Muricati Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 394. 1936, nom. superfl.
TAXON 60 (3) • June 2011: 868–884
Larridon & al. • Names of subdivisions of Cyperus
– Type: Cyperus muricatus Kük. [≡ Pycreus muricatus
(Kük.) Napper] (Art. 22.6).
When published, Cyperus sect. Muricati included the type
of Pycreus sect. Tuberculati Cherm. (1919) and is therefore
superfluous. However, Reynders & Goetghebeur (2010) misinterpreted Art. 52.3 in stating that the name C. sect. Muricati
can be used when C. divulsus Ridl. [≡ Pycreus divulsus (Ridl.)
C.B. Clarke] is excluded.
Cyperus sect. Pennati (Kunth) J. Kern in Fl. Males., ser. 1,
7(3): 635. 1974, nom. superfl. ≡ C. [unranked] Pennati
Kunth, Enum. Pl. 2: 79. 1837 – Type: Cyperus pennatus
Lam. (Art. 22.6).
= Cyperus (subg. Mariscus) sect. Turgiduli (C.B. Clarke) Kük.
When published, Cyperus sect. Pennati included the type
of C. sect. Turgiduli (C.B. Clarke) Kük. (1936) and is therefore
superfluous.
Cyperus sect. Nivei C.B. Clarke (see Cyperus sect. Hyme
nolepides Nees).
Cyperus (subg. Mariscus) sect. Phleoidei Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 426. 1936 – Type: Cyperus
phleoides (Nees ex Kunth) H. Mann. (Art. 22.6).
“Cyperus sect. Nudicaules Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 3. 1922, nom. nud. (Art. 32.1(d) and
36.1).
This intended name was based on C. nudicaulis Poir. [=
C. pectinatus Vahl].
Cyperus (subg. Mariscus) sect. Ochrocephali Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 537. 1936 – Type: Cyperus
ochrocephalus C.B. Clarke, nom. illeg. [= C. rhynchospor
oides Kük.] (Art. 22.6).
Cyperus (“sect. Eucyperus”) [unranked] Ornati Benth., Fl.
Austral. 7: 256. 1878 – Type: Cyperus ornatus R. Br. (Art.
22.6).
Cyperus [unranked] Ornati includes the type of C. sect.
Irioidei Nees (1834), which is the correct name at sectional
rank.
“Cyperus sect. Pandanophylli Cherm.” in Ann. Mus. Colon.
Marseille, ser. 3, 10(1): 27. 1922, nom. nud. (Art. 32.1(d)
and 36.1).
This intended name was based on C. pandanophyllum C.B.
Clarke.
Cyperus sect. Papyrus (Willd.) Thouars in Griseb., Fl. Brit.
W. I.: 566. 1864 ≡ Papyrus Willd., Abh. Königl. Akad.
Wiss. Berlin 1812–1813: 70. 1816 (Huygh & al., 2010) –
Type: Cyperus papyrus subsp. antiquorum (Willd.) Kük.
[= C. papyrus L. subsp. papyrus] (Art. 7.4).
Kunth (1837: 64) was the first to treat Papyrus as an infrageneric taxon of Cyperus as C. [unranked] Papyrus (Willd.)
Kunth (‘Papyri’).
Cyperus subsect. Parvuli Kük. (see Kyllinga subsect. Parvulae
(Kük.) Tang & F.T. Wang; Reynders & al., 2011).
“Cyperus sect. Paucispiculati Cherm.” in Ann. Mus. Colon.
Marseille, ser. 3, 10(1): 17. 1922, nom. nud. (Art. 32.1(d)
and 36.1).
This intended name was based on C. paucispiculatus
Cherm., nom. illeg. [≡ C. limiticola Larridon & Reynders].
Recently, Larridon & al. (2008) proposed a new name for
C. paucispiculatus Cherm. since the latter is an illegitimate
later homonym of C. paucispiculatus Boeck. [= Pycreus di
vulsus (Ridl.) C.B. Clarke].
Cyperus sect. Pilosi (J.V. Suringar) Nakai in Bot. Mag. (Tokyo)
26: 187. 1912, nom. superfl. ≡ C. [unranked] Pilosi J.V.
Suringar, Cyperus: 128. 1898 – Type: Cyperus pilosus Vahl
(Art. 22.6).
= Cyperus sect. Proceri (Kunth) C.B. Clarke.
When published, Cyperus sect. Pilosi included the type
of C. sect. Proceri (Kunth) C.B. Clarke (1883) and is therefore
superfluous.
Cyperus subsect. Pingues Kük. (see Kyllinga subsect. Pingues
(Kük.) Tang & F.T. Wang; Reynders & al., 2011).
Cyperus (subg. Mariscus) sect. Pinnati Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 451. 1936 – Lectotype (Kern,
1974): Cyperus angustatus R. Br.
“Cyperus sect. Platyphylli Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 59. 1922, nom. nud. (Art. 32.1(d) and 36.1).
This intended name was based on C. platyphyllus Roem.
& Schult.
Cyperus sect. Platystachyi (Kunth) C.B. Clarke (see Cyperus
sect. Hymenolepides Nees).
Cyperus sect. Polystachyi (C.B. Clarke) Kük., nom. illeg. (see
Pycreus P. Beauv. ser. Pycreus; Reynders & al., 2011).
Cyperus sect. Proceri (Kunth) C.B. Clarke in J. Linn. Soc., Bot.
20: 291. 1883 ≡ C. [unranked] Proceri Kunth, Enum. Pl.
2: 72. 1837 ≡ C. [unranked] Marginati Boeck. in Linnaea
35: 598. 1868 ≡ C. sect. Marginati (Boeck.) C.B. Clarke in
J. Linn. Soc., Bot. 21: 146. 1884, nom. illeg. (Art. 52.1) ≡
C. subg. Rhizomati Nakai in Bot. Mag. (Tokyo) 26: 186.
1912 – Type: Cyperus procerus Rottb. (type of C. sect. Pro
ceri Art. 22.6 – type of C. sect. Marginati Art. 7.5 – lectotype of C. subg. Rhizomati designated here).
Kunth (1837) cited the species as C. procerus Vahl and
Böckeler (1868) considered the Wallich 3355a material as mixed,
partly being C. pilosus Vahl, and partly being C. heynei Boeck.
(with reference to C. procerus Kunth). According to Art. 7.4 this
has no influence on C. procerus Rottb. being the type of Kunth’s
group of unspecified rank. Clarke (1884) and Suringar (1898)
considered the descriptions and illustrations as sufficiently resembling each other and assigned them all to C. procerus Rottb.
879
Larridon & al. • Names of subdivisions of Cyperus
Cyperus sect. Propinqui (C.B. Clarke) Kük. (see Pycreus sect.
Propinqui C.B. Clarke; Reynders & al., 2011).
Cyperus subg. Protocyperus Lye in Nordic J. Bot. 1: 57. 1981
– Type: Cyperus difformis L.
= Cyperus subg. Pycnostachys C.B. Clarke (by lectotypification).
According to Lye (1981a), his C. subg. Protocyperus includes all species of Cyperus with an eucyperoid culm-anatomy.
This includes C. diffusus Vahl, the lectotype (Tucker, 1987) of
C. subg. Pycnostachys C.B. Clarke (1893).
Cyperus sect. Pseudanosporum C.B. Clarke in J. Linn. Soc.,
Bot. 21: 117. 1884 ≡ C. sect. Natantes C.B. Clarke in
Hooker, Fl. Brit. India 6: 597. 1893, nom. illeg. (Art. 52.1)
– Type: Cyperus platystylis R. Br.
Cyperus sect. Pseudohaspani (‘Pseudo-Haspani ’) Nakai in
Bot. Mag. (Tokyo) 26: 188. 1912 – Type: Cyperus pseudo
haspan Makino [= C. tenuispica Steud.] (Art. 22.6).
Nakai (1912) mentioned C. pseudohaspan as a synonym
under C. flavidus Retz. [= Pycreus flavidus (Retz.) T. Koyama].
Cyperus sect. Pseudomariscus C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 99. 1908 (‘Pseudo-Mariscus’)
– Lectotype (designated here): Cyperus schweinitzii Torr.
The characters of C. schweinitzii fit very well the description of this section (Clarke, 1908). Clarke (1908) included
C. sect. Pseudomariscus in C. subg. Choristachys as its species have subcaduceus spikelets, in contrast to the completely
caduceus spikelets in Mariscus (which he treats as a separate
genus).
Cyperus (sect. Pycreus) [unranked] Pseudopycreus Boeck. (see
Cyperus sect. Laevigati Kük.).
Cyperus sect. Pseudopycreus (C.B. Clarke) Kük., nom. illeg.
(see Kyllinga subg. Pseudopycreus C.B. Clarke; Reynders
& al., 2011).
Cyperus subg. Pterocyperus Peterm. (see Cyperus L. sect.
Cyperus).
Cyperus sect. Pulchrae C.B. Clarke in Oliver, Fl. Trop. Afr.
8: 312. 1901 – Lectotype: Cyperus apricus Ridl. [C. semi
trifidus Schrad.].
Cyperus sect. Pumili Kük. (see Pycreus sect. Pumili (Kük.)
L.K. Dai; Reynders & al., 2011).
TAXON 60 (3) • June 2011: 868–884
Cyperus sect. Pycreus (P. Beauv.) Griseb., Cyperus subg.
Pycreus (P. Beauv.) J. Carey (see Pycreus P. Beauv. ser.
Pycreus; Reynders & al., 2011).
Cyperus sect. Pygmaei Cherm. ex Y.L. Chang in Fl. Pl. Herb.
Chin. Bor.-Or. 11: 204. 1976 ≡ Juncellus sect. Caespitosi
Nakai in Bot. Mag. (Tokyo) 26: 200. 1912 – Type: Cype
rus pygmaeus Rottb. [≡ C. michelianus (L.) Delile subsp.
pygmaeus (Rottb.) Asch. & Graebn.] (Art. 22.6).
The epithet of the name Juncellus sect. Caespitosi Nakai
(1912), which has the same type being the only species referred
to in its protologue, has priority under Art. 11.4. However, at
the same time Nakai (1912) also published a C. subg. Caespitosi
in Cyperus with a different type, i.e., C. iria L. Consequently,
Chang’s name is the correct name for this section when included in Cyperus.
Cyperus (subg. Kyllinga) sect. Queenslandiella (Domin) J. Kern
in Fl. Males., ser. 1, 7(3): 654. 1974 ≡ Queenslandiella
Domin in Biblioth. Bot. 85, 4: 415. 1915 ≡ C. subg. Queens
landiella (Domin) Govind. in Reinwardtia 9: 194. 1975
– Type: Queenslandiella hyalina (Vahl) Ballard (Art. 7.4)
[≡ Cyperus hyalinus Vahl].
≡ Queenslandiella Domin (see Huygh & al., 2010).
Cyperus sect. Radiantes J.V. Suringar, Cyperus: 84. 1898 –
Type: Cyperus radians Nees & Meyen ex Kunth (Art.
22.6).
Cyperus sect. Remirea (Aubl.) T. Koyama in Quart. J. Taiwan
Mus. 14: 162. 1961 ≡ Remirea Aubl., Hist. Pl. Guiane 1:
44. 1775 ≡ C. subg. Remirea (Aubl.) Lye in Nordic J. Bot.
3: 230. 1983 – Type: Remirea maritima Aubl. (Art. 7.4).
≡ Remirea Aubl. (see Huygh & al., 2010).
The name Remirea maritima Aubl. has not been combined
in Cyperus; Lye (1983) used the heterotypic synonym: Cyperus
pedunculatus (R. Br.) J. Kern.
Cyperus sect. Remotiglumi C.B. Clarke (see Cyperus sect.
Subimbricati C.B. Clarke).
Cyperus subg. Rhizomati Nakai (see Cyperus sect. Proceri
(Kunth) C.B. Clarke).
Cyperus sect. Rhizomatosi Kük. (see Pycreus sect. Rhizoma
tosi (Kük.) J. Raynal; Reynders & al., 2011).
Cyperus sect. Rotundi C.B. Clarke in Hooker, Fl. Brit. India 6:
614. 1893 – Type: Cyperus rotundus L. (Art. 22.6).
= Cyperus L. sect. Cyperus (by lectotypification).
“Cyperus sect. Pustulati Cherm.” in Arch. Bot. Mem. 4, Mém.
7: 19. 1931, nom. nud. (Art. 32.1(d) and 36.1).
This intended name was based on C. pustulatus Vahl.
Cyperus sect. Rupestres C.B. Clarke (see Cyperus sect. Mono
cephali C.B. Clarke).
Cyperus subg. Pycnostachys C.B. Clarke (see Cyperus sect.
Diffusi (Kunth) C.B. Clarke).
Cyperus sect. Serotini Kük., nom. illeg. (see Juncellus sect.
Stoloniferi Nakai; Reynders & al., 2011).
880
TAXON 60 (3) • June 2011: 868–884
Cyperus (sect. Diffusi) [unranked] Simplices C.B. Clarke in
Bull. Misc. Inform. Kew, Addit. Ser. 8: 98. 1908 (‘Dif
fusae’) – Type: Cyperus simplex Kunth (Art. 22.6).
This epithet has not been adopted in any ranked subdivision of Cyperus.
Cyperus sect. Solubiles C.B. Clarke (see Cyperus sect. Cype
rus).
Cyperus subg. Sorostachys (Steud.) Lye in Nordic J. Bot. 3:
230. 1983 ≡ Sorostachys Steud., Syn. Pl. Glumac. 2: 71.
1854 (Huygh & al., 2010) – Type: Sorostachys kyllingioides
Steud. (Art. 7.4). [= Cyperus pulchellus R. Br.].
“Cyperus sect. Sphacelati Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 39. 1922, nom. nud. (Art. 32.1(d) and
36.1).
This intended name was based on C. sphacelatus Rottb.
Cyperus [unranked] Sphaerolepides Boeck. in Linnaea 35:
595. 1868 – Type: Cyperus sphaerolepis Boeck. (Art. 22.6).
On first sight, Böckeler (1868) included only three species in C. [unranked] Sphaerolepides: C. iria L., C. eleusi
noides Kunth [≡ C. nutans var. eleusinoides (Kunth) R.W.
Haines] and C. nutans Vahl. However, Böckeler’s description
of C. [unranked] Sphaerolepides ends in “Inflorescentia interdum (in C. spectabili—C. filiculmi—C. leucolepidi) ad
capitulum solitarum redacta” but these species are only mentioned under his next group C. [unranked] Marginati. Under
C. [unranked] Marginati, Böckeler (1868) also published the
new species C. sphaerolepis Boeck. on which C. [unranked]
Sphaerolepides was probably based. Most likely, these species
were erroneously placed under C. [unranked] Marginati while
editing Böckeler’s notes. Although C. [unranked] Sphaerolepi
des is validly published under Art. 35.3, its epithet has not been
adopted in any ranked subdivision of Cyperus.
Cyperus sect. Spicati (Boeck.) C.B. Clarke in J. Linn. Soc.,
Bot. 20: 294. 1883 ≡ C. [unranked] Spicati Boeck. in Linnaea 36: 322. 1870 – Lectotype (designated here): Cyperus
ligularis L.
Originally, Böckeler (1870) included 21 species in C. [unranked] Spicati ; Clarke (1897) and Kükenthal (1936) later
placed 15 of these species in C. sect. Thunbergiani.
Cyperus (“sect. Eucyperus”) [unranked] Squarrosi Benth. (see
Cyperus subg. Aristomariscus Lye).
Cyperus (subg. Mariscus) sect. Strigosi Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 404. 1936 ≡ Mariscus ser.
Multif lori C.B. Clarke in Urban, Symb. Antill. 2: 40.
1900 (‘Multiflorae’) ≡ M. sect. Multiflori (C.B. Clarke)
C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8:
103. 1908 (‘Multiflorae’) ≡ M. (sect. Multiflori) subsect
Oblongispiculosi C.B. Clarke in Bull. Misc. Inform. Kew,
Addit. Ser. 8: 103. 1908 (‘Oblongi-spiculosae’) ≡ M. (sect.
Multiflori subsect. Oblongispiculosi) ser. Neogei C.B.
Larridon & al. • Names of subdivisions of Cyperus
Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 104.
1908 (‘Neogeae’) – Type: Cyperus strigosus L. (type of
C. sect. Strigosi Art. 22.6 – lectotype of M. sect. Multi
flori, M. subsect. Oblongispiculosi and M. ser. Neogei
designated here).
Mariscus sect. Multiflori includes species with manyflowered spikelets in contrast to Clarke’s other Mariscus sections, which only have a few nutlets ripening in each spikelet.
Clarke (1908) further divided this section into two subsections
and two series based on the shape of the spikelets and their new
vs. old world origin. Kükenthal (1936) used the epithet Strigosi,
instead of Multiflori when he published a section with a similar
circumscription in Cyperus.
Cyperus ([unranked] Corymbosi) [unranked] Subaphylli Boeck.
(see Cyperus sect. Corymbosi (Kunth) C.B. Clarke).
Cyperus [unranked] Subexalati C.B. Clarke (see Cyperus sect.
Compressi Nees).
Cyperus sect. Subimbricati C.B. Clarke in Hooker, Fl. Brit.
India 6: 612. 1893 (‘Subimbricatae’) ≡ C. sect. Remotiglumi
C.B. Clarke in Oliver, Fl. Trop. Afr. 8: 314. 1901 (‘Remoti
glumae’) ≡ C. sect. Zollingeriani C.B. Clarke in Kew Bull.
Addit. Ser. 8: 100. 1908 (‘Zollingerianae’) (Art. 22.6) ≡
C. sect. Subquadrangulares Kük. in Engler, Pflanzenr. IV,
20 (Heft 101): 127. 1935 – Lectotype: Cyperus zollingeri
Steud. (lectotype of C. sect. Subimbricati designated
by Kern (1974) – lectotype of C. sect. Remotiglumi and
C. sect. Subquadrangulares designated here).
Clarke (1893) included three species in C. sect. Subim
bricati: C. macer C.B. Clarke, C. tegetum Roxb. [= C. pan
gorei Rottb.] and C. zollingeri. Clarke (1908) placed all tree
species in different sections: C. macer into C. sect. Brevifo
liati, C. tegetum was put into the new C. sect. Tegetales and
C. zollingeri in the new C. sect. Zollingeriani. Kükenthal
(1935) included the first two species in C. sect. Brevifoliati,
nom. illeg. and the third species in his new C. sect. Subquad
rangulares.
Cyperus sect. Subquadrangulares Kük. (see Cyperus sect.
Subimbricati C.B. Clarke).
Cyperus sect. Subulati C.B. Clarke in Bull. Misc. Inform. Kew,
Addit. Ser. 8: 100. 1908 (‘Subulatae’) – Type: Cyperus
subulatus R. Br. (Art. 22.6).
Cyperus (subg. Pycreus) sect. Sulcati Kük., nom. illeg. non
C. [unranked] Sulcati Boeck. (see Pycreus sect. Vestiti
C.B. Clarke; Reynders & al., 2011).
Cyperus sect. Tegetales C.B. Clarke in Kew Bull. Addit. Ser. 8:
100. 1908 – Type: Cyperus tegetum Roxb. [= C. pangorei
Rottb.] (Art. 22.6).
Cyperus sect. Tenelli C.B. Clarke (see Cyperus sect. Capitu
ligeri Nees).
881
Larridon & al. • Names of subdivisions of Cyperus
Cyperus (subg. Mariscus) sect. Tetragoni Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 486. 1936 – Type: Cyperus
tetragonus Elliott (Art. 22.6).
Cyperus sect. Textiles C.B. Clarke in Oliver, Fl. Trop. Afr. 8:
312. 1901, nom. superfl. – Type: Cyperus textilis Thunb.
(Art. 22.6).
= Cyperus sect. Alternifolii (Kunth) C.B. Clarke.
When published, Cyperus sect. Textiles included the type
of C. sect. Alternifolii (Kunth) C.B. Clarke (1883) and is therefore superfluous.
Cyperus (subg. Mariscus) sect. Thunbergiani (C.B. Clarke)
Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 430. 1936 ≡
Mariscus sect. Thunbergiani C.B. Clarke in Dyer, Fl. Cap.
7: 186. 1897 – Type: Mariscus thunbergii (Vahl) Schrad.
(Art. 7.4, Art. 22.6) [≡ Cyperus thunbergii Vahl].
Cyperus subg. Torulinium (Desv. ex Ham.) Kük. (see Cyperus
subg. Diclidium (Schrad. ex Nees) C.B. Clarke).
Cyperus sect. Tunicati C.B. Clarke (see Cyperus sect. Corym
bosi (Kunth) C.B. Clarke).
Cyperus (subg. Mariscus) sect. Turgiduli (C.B. Clarke) Kük. in
Engler, Pflanzenr. IV, 20 (Heft 101): 473. 1936 ≡ Mariscus
sect. Turgiduli C.B. Clarke in Hooker, Fl. Brit. India 6:
623. 1893 – Type: Mariscus albescens Gaudich. [= C. ja
vanicus Houtt.].
Cyperus [unranked] Umbellati Boeck. (see Cyperus sect. Dis
tantes C.B. Clarke).
Cyperus sect. Umbellati (C.B. Clarke) Kük., nom. illeg. (see
Mariscus sect. Umbellati C.B. Clarke; Reynders & al., 2011).
Cyperus sect. Uncinati C.B. Clarke in Oliver, Fl. Trop. Afr.
8: 311. 1901 (‘Uncinatae’), nom. superfl. – Type: Cyperus
uncinatus Poir. [= C. cuspidatus Kunth] (Art. 22.6).
= Cyperus sect. Amabiles C.B. Clarke.
When published, Cyperus sect. Uncinati included the type
of C. sect. Amabiles C.B. Clarke (1893) and is therefore superfluous.
Cyperus sect. Vaginati (Boeck.) Kük. in Engler, Pflanzenr. IV,
20 (Heft 101): 187. 1936, nom. superfl. ≡ C. [unranked]
Vaginati Boeck. in Linnaea 35: 565. 1868 – Type: Cyperus
vaginatus R. Br. (Art. 22.6).
= Cyperus sect. Alternifolii (Kunth) C.B. Clarke.
When published, Cyperus sect. Vaginati included the type
of C. sect. Alternifolii (Kunth) C.B. Clarke (1883) and is therefore superfluous. The older name of unspecified rank does not
have priority (Art. 35.3).
Cyperus sect. Vegeti C.B. Clarke in Bull. Misc. Inform. Kew,
Addit. Ser. 8: 98. 1908 (‘Vegetae’), nom. superfl. – Type:
Cyperus vegetus Willd. [= C. eragrostis Lam.] (Art. 22.6).
882
TAXON 60 (3) • June 2011: 868–884
= Cyperus sect. Luzuloidei (Kunth) C.B. Clarke.
When published, Cyperus sect. Vegeti included the type of
C. sect. Luzuloidei (Kunth) C.B. Clarke (1883) and is therefore
superfluous.
Cyperus (subg. Mariscus) subsect. Vestiti Kük. in Engler,
Pflanzenr. IV, 20 (Heft 101): 540. 1936 – Type: Cyperus
vestitus Hochst. ex C. Krauss (Art. 22.6).
Kükenthal (1936) created this subsection in C. sect. Bulbo
caules C.B. Clarke (Kük.). Cyperus subsect. Vestiti Kük. (1936)
is different from C. sect. Vestiti (C.B. Clarke) J. Kern, nom.
illeg. (1974).
Cyperus sect. Vestiti (C.B. Clarke) J. Kern, nom. illeg. (see
Pycreus sect. Vestiti C.B. Clarke; Reynders & al., 2011).
Cyperus (sect. Pseudohaspani) subsect. Virides Nakai in Bot.
Mag. (Tokyo) 26: 189. 1912 – Type: Cyperus hakonensis
Franch. & Sav.
Cyperus sect. Viscosi C.B. Clarke in J. Linn. Soc., Bot. 21: 114.
1884 ≡ C. [unranked] Sulcati Boeck. in Linnaea 35: 547.
1868 ≡ C. [unranked] Glutinosi Boeck. in Linnaea 35: 547.
1868 ≡ C. sect. Glutinosi (Boeck.) Kük. in Engler, Pflanzenr.
IV, 20 (Heft 101): 163. 1936 – Type: Cyperus viscosus Sw.
[= C. elegans L. subsp. elegans] (type of C. sect. Viscosi
Art. 22.6 – lectotype of C. [unranked] Glutinosi (≡ C. sect.
Glutinosi) and C. [unranked] Sulcati designated here).
= Cyperus sect. Elegantes C.B. Clarke.
As Clarke (1884) is a local treatment of Cyperus (India),
he only described C. viscosus and C. oxylepis Nees ex Steud.
in C. sect. Viscosi. Kükenthal (1936) chose the epithet Gluti
nosi when he treated Böckeler’s rankless group “C. [unranked]
Sulcati s. Glutinosi Boeck.” at sectional rank. According to
Art. 34.2 both of Böckeler’s names are validly published, but
C. [unranked] Sulcati Boeck. is nomenclaturally superfluous
and illegitimate (Art. 52.1). The older names of unspecified
rank do not have priority (Art. 35.3). Böckeler (1868) included
C. viscosus, but not C. elegans. Kükenthal (1936) included
C. viscosus in synonymy under C. elegans.
Cyperus subg. Xerocyperus Lye in Nordic J. Bot. 3: 214. 1983
– Type: Cyperus rubicundus Vahl.
Cyperus sect. Zollingeriani C.B. Clarke (see Cyperus sect.
Subimbricati C.B. Clarke).
Cyperus (subg. Pycreus) [unranked] Zonati (see Pycreus ser.
Zonati (C.B. Clarke) C.B. Clarke; Reynders & al., 2011).
Acknowledgements
Nicole Hanquart (Library, National Botanic Garden of Belgium)
and the staff of the library of the Royal Botanic Gardens, Kew are
gratefully acknowledged to help with tracing several historical publications. Many thanks also to the editor John McNeill (Royal Botanic
TAXON 60 (3) • June 2011: 868–884
Garden, Edinburgh, Scotland, U.K.) and the reviewers for the careful
and critical review of the manuscript. Specimens were studied from
the following herbaria: B, BM, BR, GENT, K, P, UPS. This study
financed by the Special Research Fund (BO5622, BO7418, BOF, Ghent
University), and the Department of Biology, Ghent University.
Literature Cited
Bentham, G. 1878. Flora australiensis, vol. 7. London: Reeve.
Bentham, G. 1881. Notes on Cyperaceae. J. Linn. Soc., Bot. 18: 281–360.
Böckeler, O. 1868. Die Cyperaceen des Königlichen Herbariums zu
Berlin. 1. Theil. Die Cypereen, Scirpeen und Hypolytreen. Lin
naea 35: 397–612.
Böckeler, O. 1868–77. Die Cyperaceen des Königlichen Herbariums
zu Berlin. Linnaea 35: 397–612, 36: 271–512, 691–768, 37: 1–142,
520–647, 38: 223–544, 39: 1–152, 40: 327–452, 41: 145–356.
Böckeler, O. 1870. Die Cyperaceen des Königlichen Herbariums zu
Berlin. 1. Theil. Die Cypereen, Scirpeen und Hypolytreen. Lin
naea 36: 271–512.
Britton, N.L. 1907. The sedges of Jamaica (Cyperaceae). Bull. Dept.
Agric. Jamaica 5, Suppl. 1: 1–19.
Chermezon, H. 1919. Révision des Cypéracées de Madagascar. Ann.
Mus. Colon. Marseille, ser. 3, 7(2): 29–87.
Chermezon, H. 1922. Révision des Cypéracées de Madagascar. Ann.
Mus. Colon. Marseille, ser. 3, 10(1): 1–62.
Chermezon, H. 1931. Les Cypéracées de Haut-Oubangui. Arch. Bot.
Mem. 4, Mém. 7: 1–56.
Chermezon, H. 1937. Cypéracées. Pp. 1–335 in: Humbert, H. (ed.),
Flore de Madagascar, vol. 29. Tananarive: Imprimerie officièle.
Clarke, C.B. 1883. Cyperus. Pp. 279–296 in: Baker, J.G. (ed.), Contributions to the flora of Madagascar. — Part 3. J. Linn. Soc., Bot.
20: 237–304.
Clarke, C.B. 1884. On the Indian species of Cyperus; with remarks on
some others that specially well illustrate the subdivisions ot the
genus. J. Linn. Soc., Bot. 21: 1–202, pl. 1–4.
Clarke, C.B. 1893. Cyperaceae. Pp. 585–672 in: Hooker, J.D. (ed.), The
flora of British India, vol. 6, part 19. London: Reeve.
Clarke, C.B. 1897. Cyperaceae. Pp. 149–192 in: Thiselton-Dyer, W.T.
(ed.), Flora capensis, vol. 7. London: Reeve.
Clarke, C.B. 1900. Cyperaceae. Pp. 8–169 in: Urban, I. (ed.), Symbolae
antillanae, vol. 2, part 1. Leipzig: Fratres Borntraeger.
Clarke, C.B. 1901–02. Cyperaceae. Pp. 266–524 in: Oliver, D. (ed.).
Flora of tropical Africa, vol. 8. London: Reeve.
Clarke, C.B. 1908. New genera and species of Cyperaceae. Bull. Misc.
Inform. Kew, Addit. Ser. 8: 1–196.
Druyts-Voets, E. 1970. Types van stengel- en bladstrukturen in het
genus Cyperus L. Natuurw. Tijdschr., Antwerp. 52: 28–49.
Endlicher, S.F.L. 1836. Genera plantarum secundum ordines naturales
disposita, vol. 2. Vienna: Fr. Beck.
Goetghebeur, P. 1989. Studies in Cyperaceae 9. Problems in the lectotypification and infrageneric taxonomy of Cyperus. Bull. Soc.
Roy. Bot. Belgique 122: 103–114.
Goetghebeur, P. 1998. Cyperaceae. Pp. 141–190 in: Kubitzki, F., Huber,
H., Rudall, P.J., Stevens, P.S. & Stützel, T. (eds), The families and
genera of vascular plants, vol. 4. Berlin: Springer.
Govaerts, R., Simpson, D.A., Goetghebeur, P., Wilson, K.L.,
Egorova, T. & Bruhl, J. 2007. World checklist of Cyperaceae:
Sedges. Kew Publishing, Royal Botanic Gardens, Kew.
Govaerts, R., Simpson, D.A., Goetghebeur, P., Wilson, K.L.,
Egorova, T. & Bruhl, J. 2010. World checklist of Cyperaceae.
The Board of Trustees of the Royal Botanic Gardens, Kew. http://
www.kew.org/wcsp/monocots/ (accessed 20 May 2010).
Grisebach, A.H.R. 1846. Spicilegium florae Rumelicae, vol. 2. Brunswick: F. Vieweg et filium.
Larridon & al. • Names of subdivisions of Cyperus
Grisebach, A.H.R. 1864. Flora of the British West Indian Islands.
London: Reeve.
Haines, R.W. & Lye, K.A. 1983. The sedges and rushes of East Africa.
Nairobi: East African Natural History Society.
Hooper, S.S. 1985. Cyperaceae. Pp. 1682–1710 in: Meikle, R.D. (ed.),
Flora of Cyprus, vol. 2. Kew: Bentham-Moxon Trust.
Huygh, W., Larridon, I., Reynders, M., Muasya, A.M., Govaerts,
R., Simpson, D.A. & Goetghebeur, P. 2010. Nomenclature and
typification of names of genera and subdivisions of genera in Cy
pereae (Cyperaceae): 1. Names of genera in the Cyperus clade.
Taxon 59: 1883–1890.
Jaubert, H.F. & Spach, E. 1844. Illustrationes plantarum orientalium,
ou Choix de plantes nouvelles ou peu connues de l’Asie occiden
tale. Paris: Roret.
Kern, J.H., 1974. Cyperaceae. Pp. 435–753 in: Van Steenis, C.G.G.J.
(ed.), Flora Malesiana, ser. 1, vol. 7, part 3. Leiden: National Herbarium.
Koyama, T. 1961. Classification of the family Cyperaceae 3. Quart. J.
Taiwan Mus. 14: 159–194.
Koyama, T. 1985. Cyperaceae. Pp. 125–405 in: Dassanayake, M.D. &
Fosberg, F.R. (eds), A revised handbook to the flora of Ceylon, vol.
5. New Delhi: Amerind Publishing.
Kükenthal, G. 1935–36. Cyperaceae—Scirpoideae—Cypereae. Pp.
1–671 in: Engler, A. (ed.), Das Pflanzenreich, vol. 4, 20. Stuttgart:
Engelmann (J. Cramer).
Kunth, C.S. 1837. Enumeratio plantarum, vol. 2. Stuttgart & Tübingen:
Sumtibus J.G. Cottae.
Larridon, I., Reynders, M. & Goetghebeur, P. 2008. Cyperus limiti
cola, a new name for a Madagascan Cyperus (Cyperaceae). Novon
18: 187–188.
Larridon, I., Reynders, M., Huygh, W., Bauters, K., Van de Putte,
K., Muasya, A.M., Boeckx, P., Simpson, D.A., Vrijdaghs, A. &
Goetghebeur P. In press. Affinities in C3 Cyperus lineages (Cyperaceae) revealed using molecular phylogenetic data and carbon
isotope analysis. Bot. J. Linn. Soc.
Linnaeus, C. 1753. Species plantarum. Stockholm: Impensis Laurentii
Salvii.
Lye, K.A. 1981a. Studies in African Cyperaceae 18. Tho new subgenera
of Cyperus. Nordic J. Bot. 1: 57–61.
Lye, K.A. 1981b. Studies in African Cyperaceae 19. The genera Ano
sporum Nees and Sorostachys Steudel. Nordic J. Bot. 1: 186–191.
Lye, K.A. 1981c. Studies in African Cyperaceae 20. New taxa and
combinations in Pycreus Beauv. Nordic J. Bot. 1: 617–622.
Lye, K.A. 1982. Studies in African Cyperaceae 21. New taxa and combinations in Kyllinga Rottb. Nordic J. Bot. 1: 741–747.
Lye, K.A. 1983. Studies in African Cyperaceae 25. New taxa and combinations in Cyperus L. Nordic J. Bot. 3: 213–232.
Lye, K.A. 1992. The history of the genus Mariscus (Cyperaceae). Lidia
3: 37–72.
McNeill, J., Barrie, F.R., Burdet, H.M., Demoulin, V., Hawksworth,
D.L., Marhold, K., Nicolson, D.H., Prado, J., Silva, P.C., Skog,
J.E., Wiersema, J.H. & Turland, N.J. (eds). 2006. International
code of botanical nomenclature (Vienna Code): Adopted by the
Seventeenth International Botanical Congress Vienna, Austria,
July 2005. Regnum Vegetabile 146. Ruggell, Liechtenstein:
Gantner.
Metcalfe, C.R. 1971. Anatomy of the Monocotyledons, vol. 5, Cypera
ceae. Oxford: Clarendon Press.
Muasya, A.M., Simpson, D.A. & Chase, M.W. 2002. Phylogenetic
relationships in Cyperus L. s.l. (Cyperaceae) inferred from plastid
DNA sequence data. Bot. J. Linn. Soc. 138: 145–153.
Muasya, A.M., Simpson, D.A. & Smets, E. 2006. Isolepis tenella, a
new combination in Cyperaceae. Novon 16: 89–90.
Muasya, A.M., Simpson, D.A. & Smets, E. 2007. Isolepis levynsiana,
a new name for Cyperus tenellus (Cyperaceae). Novon 17: 59.
Muasya, A.M., Simpson, D.A., Verboom, G.A., Goetghebeur, P.,
Naczi, R.F.C., Chase, M.W. & Smets, E. 2009a. Phylogeny of
883
Larridon & al. • Names of subdivisions of Cyperus
Cyperaceae based on DNA sequence data: Current progress and
future prospects. Bot. Rev. 75: 2–21.
Muasya, A.M., Vrijdaghs, A., Simpson, D.A., Chase, M.W., Goetghebeur, P. & Smets, E. 2009b. What is a genus in Cypereae:
Phylogeny, character homology assessment and generic circumscription in Cypereae. Bot. Rev. 75: 52–66.
Nakai, T. 1912. Cyperaceae-Cyperinae Japono-Coreanae. Bot. Mag.
(Tokyo) 26: 185–207.
Nees von Esenbeck, C.G. 1834. Uebersicht der Cyperaceengattungen.
Linnaea 9: 273–308.
Opiz, F.M. 1852. Seznam Rostlin Kveteny Ceske. Prague.
Raynal, J. 1973. Notes cypérologiques: 19. Contribution à la classification de la sous-famille des Cyperoideae. Adansonia, ser. 2,
13(2): 145–171.
Reynders M. & Goetghebeur P. 2010. Reestablishment of Pycreus
sect. Tuberculati Cherm. Blumea 55: 226–230.
Reynders, M., Huygh, W., Larridon, I., Muasya, A.M., Govaerts,
R., Simpson, D.A. & Goetghebeur, P. (2011). Nomenclature and
typification of names of genera and subdivisions of genera in Cy
pereae (Cyperaceae): 3. Names of subdivisions in segregate genera
of Cyperus. Taxon 60: 885–895.
Rikli, M. 1895. Beiträge zur vergleichenden Anatomie der Cyperaceen
mit besonderer Berücksichtigung der inneren Parenchymscheide.
Jahrb. Wiss. Bot. 27: 485–580.
Schischkin, B.K. 1935. Cyperaceae and Juncaceae. Pp. 1–578 in: Komarov, V.L. (ed.), Flora unionis rerumpublicarum sovieticarum
socialisticarum, vol. 3. Leningrad: Institutum Botanicum Academiae Scientiarum URSS.
Simpson, D.A. 1990. A revision of Cyperus sect. Leucocephali. Kew
Bull. 45: 485–501.
884
View publication stats
TAXON 60 (3) • June 2011: 868–884
Simpson, D.A. & Koyama, T. 1998. Cyperaceae. Pp. 345–387 in: Santisuk, T. & Larsen, K. (eds), Flora of Thailand. Bangkok: Royal
Forest Department.
Simpson, D.A., Muasya, A.M., Alves, M., Bruhl, J.J., Dhooge, S.,
Chase, M.W., Furness, C.A., Ghamkhar, K., Goetghebeur, P.,
Hodkinson, T.R., Marchant, A.D., Nieuborg, R., Reznicek, A.A.,
Roalson, E.H., Smets, E., Starr, J.R., Thomas, W.W., Wilson,
K.L. & Zhang, X. 2007 (publ. 2008). Phylogeny of Cyperaceae
based on DNA sequence data—a new rbcL analysis. Pp. 72–83 in:
Columbus, J.T., Friar, E.A., Porter, J.M., Prince, L.M. & Simpson,
M.G. (eds.), Monocots: Comparative biology and evolution: Poales.
Aliso 23. Claremont: Rancho Santa Ana Botanic Garden.
Soros, C.L. & Bruhl, J.J. 2000. Multiple evolutionary origins of C4
photosynthesis in the Cyperaceae. Pp. 629–636 in: Wilson, K.L.
& Morrison, D.A. (eds.), Monocots: Systematics and evolution.
Melbourne: CSIRO.
Steudel, E.G. von. 1854–1855. Synopsis plantarum glumacearum, vol.
2, Cyperaceae. Stuttgart: Metzler.
Strid, A. 1974. On the typification of Bobartia L. Taxon 23: 422–424.
Suringar, J.V. 1898. Het geslacht Cyperus. Leeuwarden: Hugo Suringar.
Tucker, G.C. 1983. The taxonomy of Cyperus (Cyperaceae) in Costa
Rica and Panama. Syst. Bot. Monogr. 2: 1–85.
Tucker, G.C. 1987. The genera of Cyperaceae in the SE United States.
J. Arnold Arbor. 68: 361–445.
Van der Veken, P. 1965. Contribution à l’embryographie systématique des Cyperaceae-Cyperoideae. Bull. Jard. Bot. État Brux
elles 35: 285–354.
Väre, H. & Kukkonen, I. 2006. Typification of names in Cyperus section Arenarii (Cyperaceae). Nordic J. Bot. 24: 279–294