A conspectus of Cyperus s.l. (Cyperaceae) in Europe (incl. Azores, Madeira and Canary Islands), with emphasis on non-native naturalized species

Abstract

The genus Cyperus s.l. (incl. Kyllinga and Pycreus) (Cypereae: Cyperaceae) is critically revised in the Flora Europaea area and the Canary Islands, with emphasis on the naturalized species. A modern classification scheme and a key for the identification of all taxa (43 in number, 20 of which are non-native, naturalized species) is presented. Notes on the taxonomy, chorology, frequency and degree of naturalization for all naturalized species in the study area are provided. Original line drawings for all are also presented. The following taxa are currently accepted as locally or widely naturalized: C. aggregatus, C. alternifolius (subsp. flabelliformis and subsp. textilis), C. brevifolioides, C. brevifolius, C. congestus, C. croceus, C. eragrostis, C. exaltatus (incl. C. dives), C. gracilis, C. lupulinus, C. microiria, C. odoratus, C. reflexus, C. retrorsus, C. rigens, C. schweinitzii, C. sesquiflorus (subsp. cylindrica), C. squarrosus and C. strigosus. The taxonomy of the C. alternifolius species complex is thoroughly discussed and, as a result, C. textilis is reduced to subspecies rank. Some other nomenclatural changes, all for supraspecific taxa, are also required: sect. Fasciculati, Stoloniferi, Umbellati. Cyperus congestus and C. exaltatus are reliably reported for the first time from Spain. Bulgarian and Serbian claims for “C. strigosus” turn out to be referable to C. odoratus. The latter is also reported for the first time from Italy although its presence for many decades could be demonstrated. “Cyperus ovularis” from the Azores belongs in fact to a related species, C. retrorsus, and the species formerly naturalized in Spain (Tordera river) and previously referred to as “C. auricomus” is ascribable to C. strigosus. Finally, the (at least historical) presence of C. rigens in Italy was also approved.

Full text

A conspectus of Cyperus s.l. (Cyperaceae) in Europe (incl. Azores, Madeira and Canary
Islands), with emphasis on non-native naturalized species
Filip Verloove*
Department Spermatophytes, Botanic Garden of Meise, Nieuwelaan, Meise, Belgium
(Received 20 July 2014; final version received 6 October 2014)
The genus Cyperus s.l. (incl. Kyllinga and Pycreus) (Cypereae: Cyperaceae) is critically revised in the Flora Europaea area
and the Canary Islands, with emphasis on the naturalized species. A modern classification scheme and a key for the identifi-
cation of all taxa (43 in number, 20 of which are non-native, naturalized species) is presented. Notes on the taxonomy,
chorology, frequency and degree of naturalization for all naturalized species in the study area are provided. Original line
drawings for all are also presented. The following taxa are currently accepted as locally or widely naturalized:
C. aggregatus,C. alternifolius (subsp. flabelliformis and subsp. textilis), C. brevifolioides,C. brevifolius,C. congestus,
C. croceus,C. eragrostis,C. exaltatus (incl. C. dives), C. gracilis,C. lupulinus,C. microiria,C. odoratus,C. reflexus,
C. retrorsus,C. rigens,C. schweinitzii,C. sesquiflorus (subsp. cylindrica), C. squarrosus and C. strigosus. The taxonomy
of the C. alternifolius species complex is thoroughly discussed and, as a result, C. textilis is reduced to subspecies rank.
Some other nomenclatural changes, all for supraspecific taxa, are also required: sect. Fasciculati, Stoloniferi, Umbellati.
Cyperus congestus and C. exaltatus are reliably reported for the first time from Spain. Bulgarian and Serbian claims for
“C. strigosus”turn out to be referable to C. odoratus. The latter is also reported for the first time from Italy although its
presence for many decades could be demonstrated. “Cyperus ovularis”from the Azores belongs in fact to a related species,
C. retrorsus, and the species formerly naturalized in Spain (Tordera river) and previously referred to as “C. auricomus”is
ascribable to C. strigosus. Finally, the (at least historical) presence of C. rigens in Italy was also approved.
Keywords: Cyperus; Europe; naturalization; nomenclature; revision; taxonomy
Introduction
Cyperus L. is a nearly cosmopolitan genus of the Cyper-
aceae family (Mabberley 2008). Its generic boundaries
have long been controversial. However, recent molecular
data strongly suggest a broad circumscription: segregate
genera like Kyllinga Rottb. and Pycreus P. Beauv., all
with representatives in the study area (either native or
naturalized), are best united with it (Larridon et al.
2014). In this new circumscription Cyperus is a large
genus with c. 950 species (Larridon et al. 2013). Despite
the existence of a worldwide monograph (Kükenthal
1935–1936), the genus remains notoriously difficult in
terms of taxonomy and identification.
Recent investigations by the author, in the field as well
as in the herbarium, already revealed interesting new data
about non-native Cyperus in parts of southern Europe
(mainly Italy, Spain and the Canary Islands, see for
instance: Verloove and Sánchez Gullón 2010; Verloove and
Soldano 2011; Verloove 2012; Verloove and Mesterházy
2013). However, further study is surely required. Relatively
little is known about the frequency and distribution of most
of the introduced species, their genuine degree of
naturalization, habitats occupied, etc. Moreover, many of
these xenophytes are still poorly known and some perhaps
hide as yet unrecognized look-alikes.
A number of species in this family are economically,
ethnobotanically or horticulturally important or are
significant weeds (Simpson and Inglis 2001; Bryson and
Carter 2008). Many of the latter are rapidly spreading
beyond their native ranges and have invaded the warm-
temperate and (sub-)tropical regions of the world, includ-
ing parts of southern Europe. Four of the world’s worst
weeds are in the genus Cyperus (Carter and Bryson
1996) and it is expected that many weedy species of the
Cyperaceae family will benefit from climate change.
Especially taxa with C
4
photosynthetic pathways, such as
Cyperus rotundus (“the world’s worst weed”) are poten-
tial winners that could considerably extend their distribu-
tions, also in Europe (Simpson et al. 2011). Hence, it is
important to correctly assess the identity of any Cyperus-
like plant because an accurate identification is a critical
first step in the management of new invasions. Correctly
identified pest species allow for immediate control and
management efforts without wasting valuable resources,
placing the discipline of taxonomy at the forefront of
invasive species research (e.g. Gotzek et al. 2012).
The present paper attempts to improve the knowledge
on naturalized Cyperus in the Flora Europaea area and
the Canary Islands in presenting a key for the identifica-
tion of all taxa (native as well as introduced). All natu-
ralized species are illustrated (original line drawings),
distributional data in the study area are provided, as well
as additional taxonomic, nomenclatural or general
remarks.
*Email. filip.verloove@br.fgov.be
© 2014 Dipartimento di Biologia, Università di Firenze
Webbia: Journal of Plant Taxonomy and Geography, 2014
Vol. 69, No. 2, 179–223, http://dx.doi.org/10.1080/00837792.2014.975013

Material and methods
A large number of herbarium specimens, all from public
herbaria relevant for this study, were examined (from
BCN, BEOU, BR, C, CGE, GENT, H, LG, MA,
MSNM, PAV and SO; abbreviations according to Thiers
2008+). Some supplementary private herbaria were also
consulted. Our understanding of the taxa was further
supplemented with field observations in France, Italy and
Spain (incl. Canary Islands). Additional information on
species and (type) specimens was obtained from numer-
ous literature sources.
Original line drawings for all taxa were prepared,
based on herbarium collections from the study area.
Taxonomy
The generic boundaries of Cyperus have long been con-
tested and the species included in this account have been
accommodated in several different segregate genera in
the past [e.g. Dichostylis P. Beauv. ex T. Lestib.,
Juncellus (Griseb.) C·B. Clarke, Kyllinga,Mariscus Vahl,
Pycreus and Torulinium Desv. ex Ham.]. Bruhl (1995)
and Goetghebeur (1998) provided new classifications for
the Cyperaceae family. Recent molecular phylogenetic
studies, however, shed new light on intergeneric relation-
ships within, among others, Cypereae (e.g. Larridon et al.
2013). A much broader generic concept corroborates with
these new data. In this chapter we briefly discuss why
three of the segregate genera that were, until recently,
widely accepted in the study area (i.e. Kyllinga,Mariscus
and Pycreus), should be abandoned.
The persistence of glumes on a readily deciduous ra-
chilla was the distinguishing factor most widely used to
separate Mariscus from Cyperus. Raynal (1981), while
discussing the identity of Cyperus strigosus in the study
area, strongly advocated the segregation of Mariscus
from Cyperus. However, O’Neill (1942) already demon-
strated that several species of Mariscus in fact showed
characteristics of both genera, some even more of Cype-
rus than of Mariscus. This was later confirmed by Lye
(1992). For the taxa of “Mariscus”relevant to this study
Cyperus aggregatus,C. croceus and C. retrorsus
unequivocally pertain to Mariscus, while C. hamulosus,
C. squarrosus,C. strigosus and the entire section
Laxiglumi share features with Cyperus. Molecular data
confirmed that Mariscus is completely nested in C
4
Cyperus and not worthy of generic recognition (e.g.
Larridon et al. 2011b).
Kyllinga is a strongly supported monophyletic clade
but it is also entirely nested in C
4
Cyperus. Moreover,
morphological peculiarities (deciduous glumes, laterally
flattened gynoecia, reduced number of flowers per spike-
let, etc.) also occur in other segregates of Cyperus.
Hence, there are no sufficient arguments to warrant gen-
eric status (Larridon et al. 2013).
Pycreus is polyphyletic and also completely nested in
Cyperus, although a core-Pycreus clade is discernible
that also includes C. laevigatus. Relationships are still
poorly resolved. Characters that traditionally separate
Pycreus also occur in other segregates of Cyperus; there-
fore, no generic status should be assigned to Pycreus
(Larridon et al. 2013; Reynders 2013).
Taxonomic arrangement of the species of Cyperus
(native and naturalized) in Europe
The following classification is mostly based on Larridon
et al. (2011a) and Reynders et al. (2011), see also
Table 1. [Note: although relationships in Cyperus s.l.
are mostly evident now, in some groups they remain
largely unresolved. Pycreus, for instance, is still critical.
Its type species, Pycreus polystachyos (Rottb.) P.
Beauv., belongs to an unresolved assemblage of species.
In the present account several different sections are
upheld but sections Flavescentes and Globosi are per-
haps better included in a broadly circumscribed section
Zonati, while section Vestiti could be accepted at sub-
section level (although nested in a paraphyletic, unre-
solved matrix of the other sections). For the species
here concerned this means that Cyperus flavescens,
C. flavidus and C. mundii are probably best accommo-
dated in section Zonati, while C. polystachyos belongs
to section Pycreus (comm. M. Reynders).] Taxa that are
believed to be non-native in the entire study area are
preceded by an asterisk. Taxa in italic are not yet
included in the Euro+Med Plantbase (Jiménez-Mejías
and Luceño 2011).
Cyperus L.
Subgenus Anosporum (Nees) C.B. Clarke in J. Linn.
Soc. Bot. 21: 34. 1884
Section Luzuloidei (Kunth) C.B. Clarke, J. Linn. Soc.,
Bot. 21: 115. 1884
*Cyperus eragrostis Lam., Tabl. Encycl. 1: 146. 1791
*Cyperus reflexus Vahl, Enum. Pl. 2: 299. 1805
Section Alternifolii (Kunth) C.B. Clarke, J. Linn. Soc.,
Bot. 20: 288. 1883
*Cyperus alternifolius L., Mant. Pl. 1: 28–29. 1767
subsp. flabelliformis Kük., Pflanzenr. IV.20 (Heft
101): 193. 1936
subsp. textilis (Thunb.) F. Verloove, comb. nov.
Section Fusci (Kunth) C.B. Clarke, J. Linn. Soc., Bot.
21: 131. 1884
Cyperus difformis L., Cent. Pl. 1: 6. 1756
Cyperus fuscus L., Sp. Pl.: 46. 1753
Section Graciles (Benth.) Kük., Pflanzenr. IV. 20 (Heft
101): 202. 1936
*Cyperus gracilis R. Br., Fl. Nov. Holland: 213. 1810
180 F. Verloove

Subgenus Cyperus
Section Papyrus (Willd.) Thouars, in Griseb., Fl. Brit. W.
I.: 566. 1864
Cyperus papyrus L., Sp. Pl.: 47. 1753
Section Exaltati (Kunth) C.B. Clarke, J. Linn. Soc., Bot.
20: 293. 1883
*Cyperus exaltatus Retz., Observ. Bot. 5: 11. 1789
Section Alopecuroidei Nees, Linnaea 9: 285. 1834
Cyperus alopecuroides Rottb., Descr. Pl. Rar.: 20.
1772
Section Cyperus, Linnaea 9: 284. 1834
Cyperus cyprius Post, in Mém. Herb. Boiss. 18:
102. 1900 [Note: The exact placement of this enig-
matic species is uncertain. Kükenthal (1935–1936)
placed it in section Fusci but it is obviously much
more closely related to C. longus and probably
belongs in section Cyperus.]
Cyperus esculentus L., Sp. Pl.: 45. 1753
Cyperus longus L., Sp. Pl.: 45. 1753
Cyperus rotundus L., Sp. Pl.: 45. 1753
Section Distantes C.B. Clarke, J. Linn. Soc., Bot. 20:
290. 1883
Cyperus glomeratus L., Cent. Pl. 1: 5. 1756
Section Irioidei Nees, Linnaea 9: 285. 1834
*Cyperus microiria Steud., Syn. Pl. Glumac. 2: 23.
1855
Section Compressi Nees, Linnaea 9: 2 84. 1834
Cyperus glaber L., Mant. Pl. Alt.: 179. 1771
Section Galilea (Parl.) T.V. Egorova, Novosti Sist. Vyssh.
Rast. 34: 21. 2002
Cyperus capitatus Vand., Fasc. Pl.: 5. 1771
Section Monocephali C.B. Clarke in Oliver, Fl. Trop.
Afr. 8: 310, 1901
Cyperus rubicundus Vahl, Enum. Pl. 2: 308, 1806
Section Pygmaei Cherm. ex Y.L. Chang, Fl. Pl. Herb.
Chin. Bor.-Or. 11: 204. 1976
Cyperus michelianus (L.) Link, Hort. Berol. 1: 303.
1827
Cyperus pygmaeus Rottb., Descr. Pl. Rar.: 20. 1772
Section Stoloniferi (Nakai) F. Verloove, comb. nov.
Cyperus serotinus Rottb., Descr. Icon. Rar. Pl.: 31.
1773
Section Juncellus Griseb., Fl. Brit. W. I.: 562. 1864
Cyperus distachyos All., Fl. Pedem.: 48. 1785
Cyperus laevigatus L., Mant. Pl. Alt.: 179. 1771
Cyperus pannonicus Jacq., Fl. Austriac. 5: 29. 1778
[“Pycreus”]
Section Globosi (C.B. Clarke) Kük., Pflanzenr. IV.20
(Heft 101): 352. 1936
Cyperus flavidus Retz., Obs. Bot. 5: 13. 1788
Section Pycreus (P. Beauv.) Griseb., Spic. Fl. Rumel. 2:
419. 1846
Cyperus polystachyos Rottb., Descr. Pl. Rar. 21. 1772
Section Vestiti (C.B. Clarke) J. Kern, Fl. Males., ser. 1,
7(3): 646. 1974
Cyperus mundii (Nees) Kunth, Enum. Pl. 2: 17.
1837 [Note: The spelling of the epithet of this spe-
cies is still very controversial. The species was
named by Nees (1836) in honour of J.L. Mund (vari-
ation: J.L. Mundt, though only long after his death,
so not at the time of publication). According to the
ICBN (art. 60.1) the original spelling of a name or
epithet is to be retained; hence the correct spelling
would then be “mundtii”. However, all hinges on
whether “Mundt”is a customary spelling (Rec.
60C.4). According to Glen et al. (2003) it is not as
Johannes always wrote his name as “Mund”,so
“Mundt”must be considered an error and not a cus-
tomary spelling. Art. 60.12, referring to Rec.
60C.1.(b), would then apply and “mundii”would be
the correct (obligatory) formation of the epithet.]
Section Flavescentes Kük., Pflanzenr. IV.20 (Heft 101):
395. 1936
Cyperus flavescens L., Sp. Pl. 1: 46. 1753
[“Mariscus”]
Section Strigosi Kük., Pflanzenr. IV.20 (Heft 101): 404.
1936
*Cyperus strigosus L.
Section Spicati (Boeckeler) C.B. Clarke in J. Linn. Soc.
Bot. 20: 294. 1883
*Cyperus rigens J. Presl et C. Presl, in Reliq. Haenk.
1: 170. 1830
Section Fasciculati (Boeckeler) F. Verloove, comb. nov.
*Cyperus congestus Vahl, Enum. Pl. 2: 358. 1805
Section Laxiglumi (C.B. Clarke) Kük., Pflanzenr. IV.20
(Heft 101): 457. 1936
*Cyperus lupulinus (Sprengel) Marcks, Trans. Wis-
consin Acad. Sc. 62: 271. 1974
*Cyperus schweinitzii Torrey, Ann. Lyceum Nat.
Hist. New York 3: 276. 1836
Section Aristati Nees, Linnaea 9: 284. 1834
Cyperus hamulosus M. Bieb., Fl. Taur.-Caucas. 1:
35. 1808
*Cyperus squarrosus L., Cent. Pl. 2: 6. 1756
Section Umbellati (C.B. Clarke) F. Verloove, comb. nov.
*Cyperus aggregatus (Willd.) Endl., Cat. Horti Vin-
dob. 1: 93. 1842
*Cyperus croceus Vahl, Enum. Pl. 2: 357. 1805
*Cyperus retrorsus Chapm., in Bot. Gaz. 3: 175.
1878
Webbia: Journal of Plant Taxonomy and Geography 181

[“Kyllinga”]
Section Kyllinga (Rottb.) Kern, Fl. Males., ser. 1, 7(3):
655. 1974
*Cyperus brevifolius (Rottb.) Endl. ex Hassk., Cat.
Hort. Bot. Bogor. 24. 1844
*Cyperus brevifolioides Thieret et Delahouss. Sida 3
(3): 129. 1967
*Cyperus sesquiflorus (Torr.) Mattf. & Kük., Pflan-
zenr. IV.20 (Heft 101): 591. 1936
[“Torulinium”]
Section Diclidium (Schrad. ex Nees) Griseb., Fl. Brit. W.
I.: 565. 1864
*Cyperus odoratus L., Sp. Pl. 1: 46. 1753
Residence status
Residence status (native versus non-native) for many
species of Cyperus in Europe is open to question. In the
present paper, only species that are believed to be non-
native in the entire study area are taken into account.
Species that are considered native in at least one area
(even though they may be much more widespread as an
introduced species elsewhere in the study area) are not
dealt with. Below follows a concise, alphabetical over-
view of supposedly native species for which residence
status is in fact uncertain:
Cyperus alopecuroides: known for quite a long time
from the Canary Islands (Gran Canaria, La Palma and
Tenerife; e.g. Webb and Berthelot 1840) and considered
“possibly native”by Acebes Ginovés et al. (2009). This
is debatable: in the climatologically more favourable
Cape Verde Islands, it is believed to be an introduced
species (Sánchez-Pinto et al. 2005). A recently detected
population in Sicily was qualified as “probably autoch-
thonous”(Brullo and Sciandrello 2006).
Cyperus difformis: residence status hard to assess (see
Kükenthal 1935–1936). Probably originally native in
tropical Africa and Asia and possibly also in the warmest
parts of the study area. However, considered non-native
in France (e.g. Marnotte et al. 2006), Italy (Conti et al.
2005), Romania (Cioĉarlan 2009), etc. In Spain its resi-
dence status is unclear (Castroviejo 2007a).
Cyperus esculentus: probably only originally native in the
warmest parts of the study area but much expanding fur-
ther north in recent times. Some of the infraspecific taxa
(e.g. the more or less widely naturalized North American
var. leptostachyus Boeckeler) are doubtlessly non-native.
Cyperus glomeratus: probably only native in southeast-
ern Europe. In Italy (e.g. Fiori 1923) naturalized for
quite a long time. Recently also occurring further to the
west, for instance in France and Spain (e.g. Pascal 1978;
Castroviejo 2007a), doubtlessly as an introduction.
Cyperus mundii: probably native in Spain (Castroviejo
2007c). Recently introduced in the Canary Islands
(Tenerife; Siverio Núñez et al. 2013).
Cyperus papyrus: introduced in the warmest parts of the
study area in ancient times. In Spain it was reported as
an escape near Valencia (Peña et al. 2003) but consid-
ered a mere relic of cultivation by Castroviejo (2006,
2007a). In Sicily it is accepted as a native species by
Conti et al. (2005; see also Giardina et al. 2007),
although it is –evidently –at most an archaeophyte.
Cyperus polystachyos: is considered native in two locali-
ties in Italy (Conti et al. 2005), both near hot springs,
where it is critically endangered (e.g. Merola 1957;
Crosti and Fabrini 2010). Canarian populations (La
Gomera) are “probably introduced”(Acebes Ginovés
et al. 2009).
Cyperus rotundus: probably only originally native in the
warmest parts of the study area but much expanding fur-
ther north in recent times.
Excluded species
In the past years several different species of Cyperus
have been erroneously reported from the study area.
Claims from Spain of C. bellus Kunth (also as C. rubi-
cundus Vahl) and C. cyperoides (L.) Kuntze (Castroviejo
2005,2006,2007a) were already previously corrected
and these records are, in fact, referable to C. glaber and
C. retrorsus respectively (Verloove and Sánchez Gullón
2010; Verloove and Mesterházy 2013).
Castroviejo (2005,2007a) also reported about
C. distans L. f., a species recorded on the exposed banks
of a water reservoir near Negreira in 2000 (prov. A
Coruña, Spain). A population of this plant is still present
and a herbarium collection, with the following details,
was kindly provided by J. Amigo:
A Coruña, Negreira, a orillas del Tambre, junto a la
desembocadura del rio Barcala. En herbazal de Cypero-
Bidentetum frondosae sobre los depósitos limosos que se
forman en el anillo de inundación por efecto del embalse
Barrié de la Maza (UTM 0521428/4749806), 145 m,
02.09.2013, J. Amigo (SANT 68614, dupl. BR)
However, this collection does not belong to C. distans
but represents one of the many forms of C. esculentus.
Outlook
A lot of species of Cyperus are significant weeds (e.g.
Bryson and Carter 2008) or are otherwise of importance,
for instance as ornamentals (Simpson and Inglis 2001).
In addition to the taxa that are currently naturalized in
the study area, more are to be expected. The African
Cyperus albostriatus Schrad. is widely cultivated as an
ornamental and was recently recorded as an escape in
southern Spain (E. Sánchez Gullón, August 2014 pers.
182 F. Verloove

comm.); a future naturalization in the Mediterranean area
is very likely (see also Rosen et al. 2012). And in a plant
nursery near to Valencia, also in Spain, several supple-
mentary species of Cyperus (C. compressus,C. distans,
C. iria and C. pumilus; see Verloove et al. 2014) have
been recorded as local, probably ephemeral, weeds.
Artificial key to Cyperus species (native as well as
naturalized) in Europe, including Azores, Madeira
and Canary Islands (non-native taxa in bold face)
1. Nutlet predominantly or exclusively biconvex to
almost rounded or plano-convex, never all trigonous.
Style branches predominantly 2 (sometimes 3 in some
flowers) ........................................................................... 2
All nutlets trigonous. Style branches always 3 ........... 15
2. Spikelets with only one fertile (bisexual) floret.
Glumes deciduous. Fresh subterranean parts with very
distinct citronella-like scent (“Kyllinga”) ....................... 3
Spikelet with 2 to many fertile florets. Glumes persistent.
Subterranean parts without citronella-like scent ............ 5
3. Rhizomes short, plant tufted. Spikelets whitish. Spikes
1–3, the central cylindrical. Nutlet black at maturity
…………………………………….Cyperus sesquiflorus
Rhizomes long-creeping. Spikelets greenish. Spike usu-
ally solitary (rarely 3), spherical. Nutlet brownish at
maturity …...…………………………………………... 4
4. Glume keel with scattered glassy spines. Anther 1
.................................................................... . C. brevifolius
Glume keel smooth. Anthers 2 or 3
……… ...…………………………… C. brevifolioides
5. Nutlets plano-convex (dorsiventrally compressed),
with face towards rachilla .............................................. 6
Nutlet biconvex (bilaterally compressed), i.e. with edge
(angle) facing rachilla (“Pycreus”) ............................... 12
6. Inflorescence terminal ................................................ 7
Inflorescence apparently lateral: (main) involucral bract
leaf-like and strictly erect as a continuation of the culm
………………………………………………………... 10
7. Robust, erect, rhizomatous perennial, culm 30–250 cm.
Inflorescence compound with several spikes on distinct
stalks ………………………………………………….. 8
Small annual, often with decumbent culms usually not
exceeding 30 cm. Inflorescence a dense, sessile, ovoid to
globose head ..............................................…………….9
8. Nutlet 0.9 mm long. Culm up to 250 cm. Inflores-
cence with bracts 20–100 cm long. Spikelets 50–100,
+/–quadrangular to slightly compressed, golden. Anthers
2 ....... ................................................... .. C. alopecuroides
Nutlet 1.4–1.6 mm long. Culm rarely exceeding 100 cm.
Inflorescence with bracts 5–25 cm long. Spikelets 7–14,
distinctly flattened, dark reddish brown. Anthers 3
........................................................................ C. serotinus
9. Glumes distichously arranged .............. .. C. pygmaeus
Glumes spirally arranged ......................... C. michelianus
10. Spikelets obtuse at apex, broadly lanceolate to ovate
in outline. Nutlet nearly as long as glume. Involucral
bracts dilated at base ............................... C. pannonicus
Spikelets acute at apex, oblong or oblong-lanceolate in
outline. Nutlet at most two-thirds as long as glume.
Involucral bract not or hardly dilated at base
..........................................................………………….. 11
11. Glumes stramineous. Spikelets usually less than
10 mm long, numerous (up to 40 ) ........... .. C. laevigatus
Glumes dark purplish to nearly black. Spikelets usually
more than 10 mm long, few (up to 6) ....... C. distachyos
12. Glumes with distinct groove on both sides of the
keel. Stoloniferous perennial, extensively rooting at
nodes ............... ……………………………….. C. mundii
Glumes not grooved. Annual or short-lived perennial,
without long stolons .....................................………… 13
13. Glume apex acute. Spikelets tapering towards apex.
Nutlet narrowly oblong. Rachilla narrowly winged
….............................…………………….C. polystachyos
Glume apex obtuse. Spikelets parallel-sided, not tapering
towards apex. Nutlet oblong to obovate. Rachilla not
winged …………………………………………………14
14. Nutlet surface with distinct network of nearly isodia-
metric cells. Glumes usually brownish. Annual to short-
lived perennial, up to 60 cm tall ….. ……….. C. flavidus
Nutlet surface with transverse, whitish undulations and
indistinct longitudinally elongate cells. Glumes always
golden yellow. Always annual, rarely exceeding 30 cm
…............................. ……………………….C. flavescens
15. Leaf blade absent (reduced to basal sheaths). Robust
perennials with woody rhizomes ……...……………... 16
Plant with leaves. Annuals or perennials ………......... 18
16. Inflorescence with 100–200 rays. Culm very robust,
200–500 cm. Involucral bracts much shorter than inflo-
rescence ………………....................………... C. papyrus
Inflorescence with at most 25 rays. Culm less robust, up
to c. 150 cm. Involucral bracts much longer than inflo-
rescence ……..............................……………………... 17
Webbia: Journal of Plant Taxonomy and Geography 183

17. Inflorescence bracts 8–12, rarely exceeding 10 cm,
pungent at apex. Culm terete and perfectly smooth below
inflorescence ….......... ….. C. alternifolius subsp. textilis
Inflorescence bracts 11–25, up to 40 cm long, acute but
not pungent at apex. Culm terete but becoming distinctly
ribbed on drying, mostly scabrous below inflorescence
…………… ……… C. alternifolius subsp. flabelliformis
18. Glumes with strongly excurved mucro c. 1 mm long.
Small annuals with distinct smell of cumarin when dry
……….........................………………………………... 19
Glumes muticous or mucronate but never with a long ex-
curved mucro. Perennials or annuals, without distinct
smell of cumarin ……………………………………... 20
19. Glumes distichously (tidily) arranged, laterally green-
ish to brownish and with 7–9 veins. Spikelets 10–20
flowered. Nutlet distinctly shorter than glume (excl.
mucro) ……....... ………………………… C. squarrosus
Glumes spirally arranged (appearing very untidy),
laterally red-tinged and with 5 veins. Spikelets 6–8
flowered. Nutlet subequal to glume (excl. mucro)
…................................................................. C. hamulosus
20. Glumes 1–5. Rachilla deciduous, falling as a whole.
Spikelets always arranged along a distinct rachis. Caespi-
tose, shortly rhizomatous perennials ……… ................ 21
Glumes in normally developed spikelets 6–40. Rachilla
persistent or deciduous. Spikelets digitately arranged or
along a distinct rachis. Annuals or perennials ............. 23
21. Spike solitary or several, quite sessile, cylindrical,
c. trice as long as wide. Nutlet broadly ellipsoid, c. twice
as long as wide ………..... ……………… C. aggregatus
Spikes several and on distinct peduncles, spherical to
oblong-ovoid, at most twice as long as wide. Nutlet
oblong to ellipsoid, c.3–4× as long as wide ………... 22
22. Spikes oblong-ovoid, distinctly longer than wide,
dense …… .................................. ……………C. retrorsus
Spikes spherical, about as long as wide (or only slightly
longer than wide), usually loose …....... …….C. croceus
23. Annual. Glumes 0.5–1.3 mm long. Rachilla not
winged ……………… ...........................……………... 24
Perennial or annual. Glumes usually longer (except in
the clearly perennial C. exaltatus). Rachilla winged or
not ……… ..........……………………………………... 25
24. Spikelets 30–120 per head, 2–8 × 1 mm, with 10–40
glumes. Glumes 0.5–0.7 mm long, obovate to orbicular,
obtuse at apex. Nutlet yellowish ….........….C. difformis
Spikelets 3–12 per head, 3–6×1–2 mm, with 10–20
glumes. Glumes 1–1.3 mm long, ovate, usually with
mucro c. 0.1 mm. Nutlet white …… ........... ….C. fuscus
25. Nutlet stipitate and apiculate. Glumes bicarinate
proximally, the two keels merging in distal half to form
a single central keel ……..........................…………… 26
Nutlet stipitate or not. Glumes not bicarinate
………................................................................……… 27
26. All bracts horizontal to ascending. Spikelets golden
brown at maturity. Nutlet 1.2–1.4 mm long, with distinct
stipe. Spikes globose, 20–40 mm diameter
...................................................................…C. eragrostis
Longest bract erect, as a continuation of the culm. Spik-
elets reddish at maturity. Nutlet 0.8–0.9 mm long, with
indistinct stipe. Spikes ovoid, 9–15 mm diameter
……….……………………………………….C. reflexus
27. Spikelets digitately or stellately arranged, i.e. borne
at practically the same level upon a much shortened
rachis, or in umbellate or glomerulate heads. Rachilla
not winged (except in the very rare C. cyprius)
………………..........................................…………….. 28
Spikelets arranged at some distance from one another
upon an elongated rachis. Rachilla winged (except in
C. microiria)……… ............................................…….33
28. Stoloniferous perennial. Glumes 6–7 mm long. Plant
confined to coastal sandy places …………… ....... ………
…….........................................................…... C. capitatus
Caespitose perennial or annual. Glumes 2.5–4 mm long.
Plants not restricted to coastal sandy habitats
………….............................................………………... 29
29. Rachilla winged. Spikelets reddish to purplish
……………..........................………………….C. cyprius
Rachilla not winged. Spikelets variably colored
……………….............................……………………... 30
30. Inflorescence capitate. Spikes all sessile (rays absent).
Slender annual or perennial. Nutlet c. 1.2–1.5 mm long,
less than half as long as glumes ……………………... 31
Inflorescence capitate or umbellate. Spikes sessile and
often also on distinct rays. Coarse perennial with stout
culms and cormose, knotted rhizomes. Nutlet 1.7–
2.4 mm long, more than half as long as glumes
……………………………............................………… 32
31. Spikelets whitish-green, c. 2 mm wide, 1–3(-7) in
number. Glumes c. 2.5 mm long. Slender annual or
(more often) perennial with arching culms (often vivipa-
rous) …...................................……………….. C. gracilis
184 F. Verloove

Spikelets reddish-brown, c.4–6 mm wide, 3–17 in num-
ber. Glumes c. 3.5–4 mm long. Tussocky, tufted annual
…………………………………………….C. rubicundus
32. Glumes with distinct mucro 0.3–1 mm. Anthers 0.8–
1.4 mm long. Culm usually scabridulous, especially
below inflorescence. Rays of inflorescence 3–5, 15–80
(-150) mm long. Longest inflorescence bract erect
……....................................... ……….C. schweinitzii p.p.
Glumes with short mucro 0.05–0.2 mm long. Anthers
0.3–0.6 mm long. Culms smooth. Rays of inflorescence
usually absent (rarely 1–4, 10–60 mm long). Longest
inflorescence bract horizontal to reflexed
……...........................………………….C. lupulinus p.p.
33. Rachilla distinctly winged with revolute margins,
tightly clasping the nutlet and becoming thick and corky
at maturity, eventually disarticulating above each glume
in several units ………....................... ……... C. odoratus
Rachilla internodes different, never becoming corky
…....................................……………………………….34
34. Rhizomatous perennial, rhizomes long-creeping or
emitting stolons, often tuberiferous. Spikelets laxly
arranged (inflorescence axis clearly visible), never den-
sely congested …………...........……………………… 35
Annuals or more frequently caespitose perennials; if rhi-
zomatous, then always with short, cormose rhizomes.
Spikelets often densely congested, obscuring the inflores-
cence axis …………… .............……………………… 37
35. Tubers absent. Culms usually stout, up to 160 cm tall
……………….....................................………... C. longus
Tubers present (but not always easily uprooted and often
lacking in herbarium specimens!). Culms always shorter,
rarely exceeding 60 cm ……… .............................. …...36
36. Glumes laterally reddish with distinct green keel,
each flank with 1 or 2 often rather indistinct veins close
to the keel, glumes often 3 mm or longer. Tubers without
grey tomentum ……………………………... C. rotundus
Glumes laterally yellowish or golden with less distinct
green keel, each flank with 3 or more distinct and more
or less equally spaced veins, glumes often less than
3 mm long. Tubers, at least with age, covered with grey
tomentum ….............................................. . C. esculentus
37. Shortly rhizomatous perennial, cormose at base with
knotty rhizomes ……… ....................................……… 38
Annuals or caespitose perennials, without knotty rhi-
zomes ……… ...................................…………………..39
38. Glumes with distinct mucro 0.3–1 mm. Anthers 0.8–
1.4 mm long. Culm usually scabridulous, especially
below inflorescence. Rays of inflorescence 3–5, 15–80
(-150) mm long. Longest inflorescence bract erect
………....................................... …….C. schweinitzii p.p.
Glumes with short mucro 0.05–0.2 mm long. Anthers
0.3–0.6 mm long. Culms smooth. Rays of inflorescence
usually absent (rarely 1–4, 10–60 mm long). Longest
inflorescence bract horizontal to reflexed
……………...........................………….C. lupulinus p.p.
39. Rachilla deciduous as a whole (with the glumes per-
sistent) …………..................................……………….40
Rachilla persistent with deciduous glumes
…………....................………………………………… 42
40. Spikelets oblong-lanceolate (c. 5× as long as wide),
c. 3 mm wide. Glumes yellowish to reddish
……......................................... ………………... C. rigens
Spikelets linear (c. 10× as long as wide), c. 1.5 mm wide
or narrower. Glumes greenish yellow, golden or red to
maroon ……………………………………………….. 41
41. Glumes yellowish green, eventually golden. Nutlet
narrowly oblong, purplish brown at maturity. Culm usu-
ally short, rarely exceeding 40 cm …...... ….C. strigosus
Glumes eventually red to maroon. Nutlet obovoid, nearly
black at maturity. Culm usually tall and robust, (much)
exceeding 40 cm …… ..........……………… C. congestus
42. Annual. Glumes obovate-orbiculate. Nutlet nearly as
long as subtending glume. Rachilla not winged. Spikelets
laxly arranged ……… ......... ……………….C. microiria
Perennial (but often flowering first year!) or annuals.
Glumes linear to linear-lanceolate or elliptical-ovate.
Nutlet about half as long as subtending glume or slightly
longer (nearly as long in C. glomeratus but then glumes
linear-lanceolate). Rachilla narrowly or broadly winged.
Spikelets in dense clusters …….........………………... 43
43. Inflorescence axis short (spikelets seemingly digi-
tately arranged). Glumes 2–2.5 mm long. Nutlet obovoid.
Annual (rarely perennial) ……… ..…………….C. glaber
Spikelets arranged along a distinct axis. Glumes less than
2 mm long. Nutlet linear-oblong or obovoid. Perennial
(rarely annual) …………… ........…………………….. 44
44. Glumes pale to reddish brown, linear-lanceolate, with
very indistinct mucro. Nutlet linear-oblong, nearly as
long as subtending glume …............. …... C. glomeratus
Glumes golden, ovate with short but distinct mucro. Nut-
let obovoid, trigonous, about half as long as subtending
glume…………........................…………….. C. exaltatus
Species sequence follows Larridon et al. (2011a) and
Reynders et al. (2011).
Webbia: Journal of Plant Taxonomy and Geography 185

(I) Subgenus Anosporum
Section Luzuloidei
Cyperus eragrostis Lam., Tabl. Encycl. 1: 146. 1791.
(Figure 1)
(=) Cyperus vegetus Willd., Species Plantarum. Editio
quarta 1(1): 283. 1797
Native: South America. Naturalized in Europe, Australia,
Asia and Africa.
Figure 1. Cyperus eragrostis: Habitus (1a, 1b), spike (2), spikelet (3), glume (4), nutlet + style (5).
186 F. Verloove

The invasion history of C. eragrostis in Europe was
thoroughly documented by Petřík (2003). Additional,
more recent information was provided for Belgium
(Verloove 2006b), the Netherlands (Bremer 2006) and
Slovenia (Dakskobler and Vreš2009).
Cyperus eragrostis grows in a wide range of (at least
temporarily) wet habitats. These may range from highly
disturbed (e.g. rough ground, agricultural fields) to natu-
ral, mostly wetlands, exposed river banks and pond mar-
gins (see also Petřík 2003). It locally behaves like an
invasive species.
There seems to be fairly little variation in C. eragrostis
in Europe although plants may range in stature from deli-
cate to very robust, with flowering stems ranging from 15
to 100 cm respectively. Likewise, inflorescences may be
either sessile with primary peduncles absent or composed
with primary peduncles up to 15 cm long. Delicate plants
with compact inflorescences may be distinguished as var.
compactus (Desv.) Kük. (beautifully illustrated in Barros
1960; see also Kükenthal 1935–1936; Denton 1978) but
this taxon is probably of little taxonomic value. Nutlets
also seem to be very homogenous (about twice as long as
wide and with a distinct stipe and apical beak). Worth
mentioning is a single collection from St Aygulf (France;
see “Specimens examined”) in which nutlets are slightly
longer and narrower than is usually the case. However, in
all other respects, this collection does not differ from
C. eragrostis. A few collections (e.g. R. Otto 20,634 from
La Palma, Canary Islands) are characterized by remark-
ably declined glumes, a feature considered typical of
C. ochraceus Vahl (Denton 1978; Tucker et al. 2002).
However, glume declination occasionally occurs in
C. eragrostis as well (and perhaps in other members of the
Luzulae group) and is not an exclusive character of
C. ochraceus. Earlier authors (e.g. Kükenthal 1935–1936;
Barros 1960) distinguished these species on very different
grounds, without taking into account glume declination.
Several related species from the Luzulae group are
worth mentioning. Cyperus surinamensis Rottb., a weedy
species widely distributed in the Antilles, Mexico, Central
and South America and the southernmost parts of North
America, is increasingly reported as an introduction in the
Old World (e.g. Chen et al. 2009; Dai et al. 2010a; Viji
et al. 2013). This species was collected in the nineteenth
century in Greece [Graecia, communic. Landerer &
Sartoni 1835 ex herb. Martii, BR; sub C. eragrostis] but it
is unclear whether or not this plant was collected in the
wild. In general appearance C. surinamensis is much rem-
iniscent of C. eragrostis but it usually has stems that are
scabrous or scabrellate (at least below the involucral
bracts) (versus stems smooth), smaller glumes (c.1–
1.5 mm long versus 2–2.3 mm) and a narrower nutlet. It
may have been overlooked so far in Europe.
Another relative of C. eragrostis is C. luzulae (L.)
Rottb. ex Retz. (incl. C. entrerianus Boeckeler). It is origi-
nally native in Mexico, Central and South America but
increasingly naturalized in North America (e.g. Carter
1990; Carter and Jones 1991) and it readily became an
invasive environmental and agricultural weed there (Carter
and Bryson 1996; Rosen et al. 2006; Bryson and Carter
2012). Its successful naturalization is linked to its rapid
growth, morphological plasticity, high survival and
generalized habitat requirements (King et al. 2012). It is
not easily eradicated (“deeprooted sedge”). In Europe, at
least to our knowledge, it has not yet been recorded in the
wild but it is offered for sale as an ornamental plant in the
nursery trade, for instance in Germany (R. Otto, 2013
pers. comm.; identity confirmed by the author). Like
C. eragrostis it has smooth stems but its nutlets are nar-
rowly trigonous (four to five times longer than wide, ver-
sus two times longer than wide) and the medial part of the
glumes tend to be paler (whitish to stramineous).
For the sake of completeness these three species are
opposed in the following identification key:
1. Culms scabrous with minute retrorse prickles, at least
below involucral bracts. Spikelets multi-flowered, with
20–58(-72) glumes …......... …….. Cyperus surinamensis
Culms smooth throughout. Spikelets less densely flow-
ered, with c.6–30 glumes …......................................…2
2. Nutlet narrowly trigonous, c.4–5 times longer than
wide. Glumes 1–1.5 mm long, with whitish stramineous
medial part …… .............................………….. C. luzulae
Nutlet broadly trigonous, c. twice as long as wide.
Glumes 2–2.3 mm long, with light brown medial part
………...................................……………… C. eragrostis
Selection of specimens examined (chronologically per
geographical unit)
AZORES –Flores: 07.1842, H·C. Watson 264 (BR; sub
C. polystachyos); Corvo: Vila do Corvo, near the town,
12.06.1964, P. Dansereau et al. 600 (LG); Santa Maria:
Pico(s), near the top, 28.07.1970, A. Hansen s.n. (C);
Santa Maria: Vila do Porto, nearly dry brooklet-bed near
the harbour, 28.08.1970, A. Hansen s.n. (C); Santa
Maria: São Lourenço (east), 29.08.1970, A. Hansen s.n.
(C); São Miguel: Ponta Delgada, waste place at the air-
port, 28.06.1971, A. Hansen 14 (C); São Miguel: Pedras
de Galega, ditch, 29.06.1971, A. Hansen 37 (C); São
Miguel: Furnas, town, near the “boilers”, 29.06.1971, A.
Hansen 46 (C); São Miguel: Sete Cidades, at the shore
of Lagoa Azul, 30.06.1971, A. Hansen 120 (C); São
Miguel: Sete Cidades, at the shore of Lagoa Azul,
02.07.1971, A. Hansen 202 (C); Terceira: Angra, waste
place, 02.07.1972, A. Hansen 156 (C); Terceira: São Se-
bastião, 13.03.1973, B. Gonçalves 4869 (C); Faial:
Horta, on waste place, 02.07.1973, A. Hansen 25 (C);
Graciosa: Luz, c. 130 m, 21.08.1973, B. Gonçalves 5340
(C); Pico: Madalena, on roadside, 04.06.1974, A. Hansen
79 (C); São Miguel: 2 km west of Furnas, 27.06.1979, F.
Skovgaard s.n. (C); São Miguel: Ribeira Quente (S of
Furnas), near the road tunnel, 04.04.1984, A. Hansen
1035 (C); Pico: Cais do Pico, in waste place,
Webbia: Journal of Plant Taxonomy and Geography 187

05.07.1987, A. Hansen 1374 (C). CANARY
ISLANDS –Tenerife: Tejina, fossé, 05.1962, L. Delvos-
alle s.n. (BR); Tenerife: Anaga-Gebirge, Ruderalflur am
Strassenrand, 20.04.1970, R. Otto 5591 (priv. herb. R.
Otto); Tenerife: entre San Andrés et Taganana, près de
l’Emisora forestal, lisière de la laurisilve, 03.01.1972, J.
Duvigneaud 72Can273bis (BR); Tenerife: San Andrés,
fossé le long de la route vers El Bailadero, 03.01.1972,
J. Duvigneaud 72Can245 (BR); Tenerife: Igueste, bord
de l’eau dans le barranco au pied du village, 03.01.1972,
J. Duvigneaud 72Can229 (BR); Tenerife: entre Cruz del
Carmen et Pico del Inglés, sur la route La Laguna/Las
Mercedes/Pico del Inglés, dans un fossé, à droite de la
route, 11.07.1984, G. Van Buggenhout s.n. (LG); La
Palma: Los Sauces, Barranco del Agua, Ruderalfläche
auf unbefestigtem Parkplatz, 25.08.1998, R. Otto 1274
(BR); La Palma: Santa Cruz, Avenida Bajamar, frisch
angesäte Rasenrabatte bei Hafeneinfahrt, 11.08.2007, R.
Otto 13392 (BR); Tenerife: Playa de Las Americas, bor-
der of TF1 motorway, plantation, 1x, 08.05.2009, F. Ver-
loove 7557 (BR); Gran Canaria: Cueva de las Niñas,
embalse, pond margin, common, 08.11.2011, F. Verloove
9140 (BR); La Palma: Los Sauces, Ortseingang von Los
Tilos kommend, Wasserrinne an LP 105, 24.05.2013, R.
Otto 20,216 (BR); La Palma: Los Sauces, Parkplatz an
LP 105, 300 m vor Los Tilos, 24.05.2013, R. Otto
20217 (BR); La Palma: Los Sauces, Uferpromenade
zwischen Chargo Azul und San Andrés, Brache,
26.05.2013, R. Otto 20258 (BR); La Palma: Los Sauces,
San Andrés, Graben an LP 104, 26.05.2013, R. Otto
20267 (BR); La Palma: Los Sauces, San Andrés, feuch-
ter Graben am Straßenrand LP 104 unter überrieseltem
Felsen, 26.05.2013, R. Otto 20268 (BR); La Palma: Bre-
na Alta, ruderalisierte Brache bei Poligono Industrial el
Molino, 06.06.2013, R. Otto 20386 (BR); Gran Canaria:
Cercados de Araña, embalse de Chira, gravelly bank, fre-
quent, 14.09.2013, F. Verloove 10512 (BR); La Palma:
San Andrés, feuchter Graben am Strassenrand unter
überrieseltem Felsen, Population kleinwüchsiger Pflanzen
trotz gutter Wasserversorgung, 20.09.2013, R. Otto
20634 (BR). FRANCE (incl. Corsica) –Dep. Gironde,
Bacalan, près Bordeaux, fossés des marécages de la rive
gauche de la Garonne, 20.08.1859, L. Motelay s.n. (BR);
Dep. Gironde: Bordeaux, allée du Boutaut, naturalisé
dans les fossés, 07.1862, L. Motelay s.n. (BR); Dep. Gir-
onde: Bordeaux, chemin de Labarde, 15.09.1863, E.
Gaedefroy s.n. (BR); Dep. Gironde: Bordeaux, fossés à
Bacalan, 19.09.1863,(LG); Dep. Gironde: Bordeaux,
près du fleuve en aval, rive gauche, naturalisé en abon-
dance, apporté sans doute par le lest des navires,
08.1867, (LG); Dep. Pyrénées-Atlantiques, aux environs
de Bayonne, complètement naturalisé, 07.1880, Blanchet
s.n. (GENT); Dep. Pyrénées-Atlantiques, Bayonne, par-
ties marécageuses de l’ancien arsenal maritime près de
l’Adour, 14.07.1881, Blanchet s.n. (BR, LG);Dep. Héra-
ult: Lattes (Maurin), 09.1882 (BR); Dep. Pyrénées-Atlan-
tiques: Hendaye, embouchure de la Bidassoa, 07.1890,
A. Lefebvre s.n. (BR); Dep. Pyrénées-Orientales: Perpi-
gnan, bords de la Têt, 27.09.1894 (BR, sub C. globulo-
sus); Dep. Hérault: Bédarieux à Hérépian, bords de
l’Orb, 18.08.1897 (BR); Dep. Hérault: Lamalou, bords
de l’Orb, 09.1910, P. Fourès s.n. (BR); Dep. Hérault:
Hérépian, bords de l’Orb, naturalisé (provient des laines
de Bédarieux), 26.07.1917, J.B. Renaud s.n. (BR); Dep.
Hérault: Béziers, bords de l’Orb, 25.09.1920, J.B. Re-
naud s.n. (BR); Dep.: Gironde: Bordeaux, vases de la
Garonne (rive gauche), au-bas du Pont du chemin de fer,
16.08.1927, J. Jallu 27305 (LG); Dep. Alpes-Maritimes:
Cannes, colline de Super-Cannes, dans un petit chemin
humide débouchant en face de la plage de Golfe Juan,
13.08.1937, J. Callé 3212 (BR); Dep. Pyrénées-Orien-
tales: Collioure, 07.1948, A. Lefebvre s.n. (BR); Dep.
Landes: Aire-sur-Adour, bord de l’Adour, 11.08.1949,
Baily s.n. (BR); Dep. Pyrénées-Orientales: Banyuls, fossé
le long de la route Banyuls-Cerbère, 50 m, 20.07.1950,
W. Mullenders 6499 (BR); Dep. Pyrénées-Orientales: Sa-
inte-Marie-Plage, 20.07.1950, J.-M. Warlet 126/601
(BR); Dep. Pyrénées-Atlantiques: Bayonne, fossés et
bas-cotés de la route de Boucau, entre l’Adour et la voie
ferrée, 21.07.1952, J. Jallu 5295 (BR, LG); Dep. Corse-
du-Sud: Ajaccio, chemin du Salario, lieux herbeux,
28.07.1954, Isaacson s.n. (BR); Dep. Pyrénées-Orien-
tales: Sainte-Marie, im Fluss Têt bei der Brücke der
D.11, 18.07.1955, E. Berger s.n. (LG); Dep. Alpes-Mari-
times: entre Cannes et Vallauris, versant Sud de l’Esterel,
07.1956, N. Adam s.n. (LG); Dep. Corse-du-Sud: Ajac-
cio, bord d’une route, 07.1957, C. Pelgrims s.n. (BR);
Dep. Landes: Seignosse, bords piétinés de l’Etang Blanc,
25.08.1965, C. Vanden Berghen s.n. (BR); Dep. Seine-
et-Marne: Seine-Port, adventice, 24.09.1966, N. Hallé
s.n. (BR); Dep. Landes: Sanguinet, chemin humide,
03.07.1967, C. Vanden Berghen s.n. (BR); Dep. Pyré-
nées-Orientales: Banyuls-sur-Mer (nabij Frans-Spaanse
grens), camping municipal, droge gracht, 21.07.1968, L.
Vanhecke 1020 (BR); Dep. Alpes-Maritimes, Cannes, au
lieu-dit Observatoire (Super Cannes), fossé, 20.06.1969,
G. André s.n. (BR); Dep. Corse-du-Sud: Golfe de Sa-
gone, embouchure du Liamone, 11.07.1970, J. Duvig-
neaud 70C632 (BR); Dep. Pyrénées-Orientales: Argelès-
sur-Mer, route N114, entre le village et le Racou, fossé,
06.08.1971, A. Lawalrée 16756 (BR); Dep. Var: Le Can-
net de Maures, Basse Verrerie, im Aillebett, 19.07.1972,
E. Berger 4475 (LG); Dep. Landes: Beyres, ten noorden
van Bayonne, strandje langs Lac d’Irieu, 07.07.1973, J.-
E. De Langhe 268/73 (BR); Dep. Gironde, Langon,
08.1973, P. Sotiaux s.n. (BR); Dep. Landes: Dax, rive de
l’Adour, 08.1973, P. Sotiaux s.n. (BR, LG); Dep. Var:
Le Cannet de Maures, Aillebett bei der Brücke der
N.558, 26.09.1973, E. Berger 4738 (LG, MSNM
36889); Dep. Landes: Saubusse, bord de l’Adour,
18.07.1975, N. Cnops 75/112 (BR); Dep. Pyrénées-Ori-
entales: Banyuls-sur-Mer, dans le lit à sec du Baillaury
dans le village, 16.06.1975, E. Jacques 9655 (BR); Dep.
Var: St.-Aygulf, N-helling van een 10 m diepe holle weg
tussen 2 rotswanden met kleiige aarde tussen de rotsen,
op natte plaats met in onderste zone vooral grassoort
188 F. Verloove

Pennisetum villosum, 01.08.1976, K. De Waele 1063
(GENT); Dep. Pyrénées-Orientales: nord-ouest de l’Ile-
sur-Têt, le long de la route D2 près Belvédère, petit
fossé à côté de la route plein de gravier fin, 20.07.1978,
P. Van der Veken 12823 (BR, GENT); Dep. Corse-du-
Sud: Porto, embouchure du ruisseau de Porto, gravillons
au bord du ruisseau, 12.06.1979, J. Lambinon 79/Co/435
et J. Rousselle (LG); Dep. Alpes-Maritimes: St.-Laurent-
du-Var, sur sable vaseux dans l’embouchure du Var,
19.08.1979, H.G. Rabyns 6103 (BR); Dep. Pyrénées-At-
lantiques: Orthez, usines du Bois Béarnais, c. 50 m, ter-
rain vague graveleux, temporairement inondé,
21.06.1985, J. Lambinon 85/F/123 et J. Vivant (LG);
Dep. Haute-Corse: Base militaire dite de Solenzara, au
nord de l’embouchure du Travo (côte orientale), endroit
argileux humide mais asséché en été le long de la piste
d’aviation, 02.06.1986, J. Lambinon 86/Co/164 (BR,
LG); Dep. Hérault: Aniane, bord de l’Hérault à l’hauteur
de la source Saint-Pierre, 07.1987, D. Geerinck-Coutrez
4203 (BR); Dep. Lozère: Pied-de-Borne, river Chassezac,
c. 550 m, muddy shore of artificial river basin, with fluc-
tuating water level (+/–1.5 m !), 01.08.1990, P. Goetg-
hebeur 7425 (BR, GENT, LG, MSNM 31599); Dep.
Ardèche: Borne, ruisseau, 12.09.1991, R. Dechamps
10448 (BR; sub C. fuscus); Dep. Ardèche: Notre-Dame
de Bon Secours (Lablachère), bord de ruisseau,
31.08.1992, R. Dechamps 10590 (BR; sub C. longus);
Dep. Aveyron: Millau, aux Ondes (Pont de Raujolles),
grindstrandjes langs de Tarn, 08.08.1993, F. Verloove
Fr.283 (BR); Dep. Gard: Rousson, bord de ruisseau entre
Brissac et Pont d’Avène, 07.1994, D. Geerinck-Coutrez
7366 (BR); Dep. Var: Cogolin, près de Saint-Tropez,
bord de la route, fossé, 08.08.1994, F. Verloove Fr.443
(BR); Dep. Gard: Collias, near Pont-du-Gard, streambed
of Gardon river, abundant, 14.08.1998, F. Verloove 3199
(BR); Dep. Gironde: Bordeaux, rive droite de la Gironde,
bord de la Gironde, quelques plantes, 08.2001, F. Verloo-
ve 5018 (BR); Dep. Gard: Sylvéréal, RD202 towards
Saint-Gilles (Camargue), ditch, locally, 13.09.2006, F.
Verloove 6509 (BR). ITALY –Liguria, Prov. Savona: Pi-
etra Ligure, lungo il torrente Maremola, c. 10 s.l.m.,
greto di torrente, 12.07.2004, G. Galasso s.n. (MSNM
39087, 39088); Liguria, Prov. Savona: Pietra Ligure, vi-
ale G. Moretti, c. 10 s.l.m., aiuola, 25.07.2005, G. Ga-
lasso s.n. (MSNM 40063, 40067); Liguria, Prov.
Savona: Pietra Ligure, torrente Maremola, destra idrogra-
fica, c. 10 s.l.m., greto di torrente, 26.07.2005, G. Galas-
so s.n. (MSNM 40070, 40071); Lombardia, Prov.
Milano: Milano, piazzale V. Gambara, tra via A.T. Triv-
ulzio e via P. Fornari, c. 120 s.l.m., lato esterno del
marciapiede, 25.05.2007, G. Ceffali s.n. (MSNM 41943,
41944); Lombardia, Prov. Pavia: Viale Lungo Ticino
Sforza, left bank of river Ticino (WGS84:
45°10’51.45" N, 09°09’17–18" E), abandoned flowerbed
(previously flooded), on wet sandy soil, 66 m,
25.09.2013, N. Ardenghi and P. Cauzzi s.n. (BR); Pie-
monte, Prov. Vercelli: Arborio, left bank of river Sesia,
exposed river bank, few plants, 06.09.2014, F. Verloove
10983 (BR). Lombardia, Prov. Pavia: Pancarana, river
Po, sandy river bank, a single individual, 10.09.2014, F.
Verloove 10998 (BR); Emilia-Romagna, Prov. Piacenza:
Castelvetro Piacentino, località Maginot, river Po, muddy
river bank, 14.09.2014, F. Verloove 11021 (BR); Lom-
bardia. Prov. Cremona: Casalmaggiore, river Po west of
railway bridge, frequent, 14.09.2014, F. Verloove 11028
(BR). MADEIRA –Machico, bord de mer, en arrière de
la plage de galets, fossé, 1975, J. Duvigneaud 75M544
(BR); Loreto (32°43’35" N, 17°08’41" W), 18.08.2014,
Q. Groom s.n. (BR). PORTUGAL –Prov. Coimbra:
Montemor-o-Velho, sur les bords des rizières dans la pla-
ine alluviale du Mondego, 03.08.1959, J. Duvigneaud
s.n. (BR, LG); Prov. Beira Litoral: Pessegueiro do Vou-
ga, in arenosis humidis ad rivo Vouga, c. 50 m s.m.,
05.08.1962, A.R. Pinto da Silva 6852 (LG). SPAIN –
Prov. Vizkaya: environs de Lekeitio (vers Bermeo), mar-
écage près d’une fontaine, 28.07.1955, J. Duvigneaud
s.n. (BR); Prov. Girona: entre Lloret de Mar et Tossa de
Mar, bord de la route, 26.07.1966, N. Cnops 66.79 (BR);
Prov. Girona: Blanes, Rio Tordera, sables de la rivière,
29.07.1966, N. Cnops 66.93 (BR); Prov. Gipuzkoa: langs
de weg Aya (Aia) naar Orio, in slijk, 08.1972, H.G. Rab-
yns 2670 (BR); Prov. La Coruña: Santiago, Burgo de las
Naciones, cunetas humedas, 28.06.1976, Losa, Quintana
et al. s.n. (LG); Prov. Barcelona: Malgrat de Mar, bancs
de sable et rives du Rio Tordera, 5 m, 22.06.1978, J.-J.
Symoens E2209 (BR, GENT, LG); Prov. Barcelona: bai-
xados de Peralada, 12.06.1981, T. Casasayas Fornell s.n.
(BCN 68666); Prov. Gipuzkoa: Donostia, Miramon
(UTM 30TWN8294), 100 m, zonas encharcadas en pis-
tas forestales, sustrato margoso, 18.10.1992, P. Garin
s.n. (BR, LG, MSNM 33,713).
Cyperus reflexus Vahl, Enum. Pl. 2: 42. 1805 (Figure 2)
Native: South America (Argentina, Bolivia, Brazil, Chile,
Paraguay, Uruguay), Central America, Mexico and USA
(Texas). Naturalized in Europe.
Cyperus reflexus is a regular but usually ephemeral alien
in several parts of Europe. It was known, for instance, as
an alien associated with the former wool-processing indus-
try in Belgium and the British Isles (e.g. Verloove 2006a;
Clement 1984). In France it was recorded in 1959 in
Eysines, near Bordeaux, but not confirmed afterwards
(Vivant 1970). However, a population discovered in a
ditch in Le Muy in 1977 (Auriault 1979) survived and
now C. reflexus occurs in a second locality in this area
(see “Specimens examined”). Still in the Var department it
is also naturalized in the valley of riverlet Môle near Bor-
mes-les-Mimosas (comm. H. Michaud, April 2013). In
2001 it was furthermore discovered in the valley of the
river Dordogne between Tauriac and Prudhomat in the Lot
department and soon afterwards on several different
occasions in the same area between Tauriac and Pinsac
(Felzines and Loiseau 2003; Felzines 2004).
Webbia: Journal of Plant Taxonomy and Geography 189

Cyperus reflexus is usually found in sandy or grav-
elly, temporarily wet habitats, mostly on exposed river
banks and pond margins.
All populations currently known from France have
been ascribed to var. fraternus (Kunth) Kuntze (syn.
C. fraternus Kunth) (Felzines and Loiseau 2003; J.-M.
Figure 2. Cyperus reflexus: Habitus (1), spike (2), spikelet (3), glume (4), nutlet + style (5).
190 F. Verloove

Tison, 2013 pers. comm.) on behalf of their stiffly erect
involucral bracts, many-headed inflorescences and small
nutlets (Barros 1960). The taxonomic value of this variety
is questionable. In a monographic study of the Luzulae
group of Cyperus, Denton (1978) accepted both varieties
but on slightly different grounds (achene versus scale
length, achene form, scale length, etc.). Other authors
reject the existence of any infraspecific taxa (e.g. Tucker
and McVaugh 1993; Tucker 1994; Tucker et al. 2002).
However, recent Brazilian taxonomists tend to maintain
these two taxa. According to Araujo and Longhi-Wagner
(1996) both are best distinguished on inflorescence char-
acters: a monocephalous, globose head in var. reflexus and
an anthelodium in var. fraternus. The latter also seems to
be confined to rather damp habitats while var. reflexus is
relatively drought tolerant. All French populations indeed
tend to have an anthelate inflorescence and exclusively
occur in at least temporarily damp habitats. However,
inflorescence shape does not seem to be correlated with
glume and nutlet characters. Therefore, C. reflexus is here
accepted in a broad sense so as to include C. fraternus.
Selection of specimens examined
FRANCE –Dep. Var: Le Muy, Château du Rouet,
09.1992, J.M. Tison s.n. (priv. herb. J.-M. Tison); Dep.
Lot: La Treyne (Pinsac), border of the river Dordogne,
close to RD 43, muddy border of river, one specimen
(but recently becoming naturalized in this area),
07.09.2003, F. Verloove 5478 (BR); Dep. Var: Le Muy,
Château du Rouet, bord de la route, 09.07.2013, H. Mi-
chaud s.n. (BR); Dep. Var: Le Muy, La Roquette, fossés
en bord de route, 09.07.2013, H. Michaud s.n. (BR);
Dep. Var: Bormes-les-Mimosas, aux Campaux, mares
artificielles dans la vallée de la Môle, 03.08.2013,
H. Michaud s.n. (BR).
Section Alternifolii
The taxonomy of section Alternifolii is very complex
and largely unresolved. Kükenthal (1935–1936; as sec-
tion Vaginati) accepted nine species, most of them
confined to South Africa and Madagascar. Especially the
so-called “umbrella sedges”, i.e. the species related to
C. alternifolius, form a critical group (see for instance:
Baijnath 1975; Kukkonen 1990; Chen et al. 2008;
Gordon-Gray and Ward 2010). These plants, readily rec-
ognized by their leafless culms bearing an apical tuft of
leaf-like bracts among which the inflorescences are
borne, are popular ornamentals and some are also used
for weaving or mat-making. As such, they are widely
introduced in warm-temperate and tropical regions of the
world and often escape. Morphological relationships are
close within the complex and species boundaries are
weak. Distinguishing features are based, among others,
on details of culm shape and texture and not always eas-
ily assessed. In this account two taxa are accepted
(C. alternifolius subsp. flabelliformis and subsp. textilis).
Their taxonomic rank is debatable. However, species
rank for both seems to be too elevated (see below).
Molecular data could probably shed new light on the
relationships within this species complex. Cyperus alter-
nifolius s.str. and C. sexangularis Nees, the latter with
clearly six-angled, scabrid culms (doubtlessly also best
subsumed under C. alternifolius), may have been over-
looked in our area.
Cyperus alternifolius L., Mant. Pl. 1: 28–29. 1767
subsp. flabelliformis Kük., Pflanzenr. IV. 20(Heft 101):
193. 1936. (Figure 3)
(≡)Cyperus flabelliformis Rottb., Descr. Icon. Rar. Pl.
42, pl. 12, f. 2. 1773
(=) Cyperus involucratus Rottb., Descr. Pl. Rar. 22. 1772
(=) Cyperus alternifolius L. var. obtusangulus (Boeckeler)
T. Koyama, J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 8: 120.
1961
Native: East Africa. Naturalized in North America, Asia,
Australia and Europe.
This taxon is perhaps better known as C. involucratus.It
has long been confused with C. alternifolius s.str. but is
more frequently cultivated as an ornamental and also
more common as an escape, although genuine C. alter-
nifolius is also increasingly found outside its area of ori-
gin (Madagascar and Réunion) (e.g. Chen et al. 2008).
Several authors have discussed their separation (e.g.
Koyama 1961; Baijnath 1975; Kukkonen 1990; Chen
et al. 2008; Gordon-Gray and Ward 2010) but not all
features provided by these authors proved to be reliable
and some were in fact inconsistent or simply erroneous.
Both seem to be best opposed by their nutlets. Subsp.
flabelliformis has smaller, obovoid nutlets c. 0.4–0.8 mm
long and less than half the glume length (versus elliptic
nutlets c. 1.1–1.4 mm long and more than half the glume
length. Also, glumes tend to be broadly ellipsoid in
subsp. flabelliformis but lanceolate in subsp. alternifo-
lius. Nutlet versus scale length is an important distin-
guishing feature, according to Koyama (1961), perhaps
the only reliable one. Culm characters can be indicative
as well: culms of subsp. flabelliformis are often, at least
when young and undamaged, scabrous (versus always
smooth in subsp. alternifolius). Both subspecies have
subterete stems when fresh but these become shallowly
ridged-striate after drying in subsp. flabelliformis while
they remain more or less smooth and terete in subsp. alt-
ernifolius. Further anatomical characters are emphasized
by Baijnath (1975). Other features (e.g. degree of
rigidness of involucral bracts; spikelet length; shape and
colour of glumes and width of their hyaline margin;
nutlets stipitate or not; etc.) are probably of a lesser or
no taxonomic value at all. Chen et al. (2008) refer to
inflorescence bracts in subsp. flabelliformis that have
“margins with unconspicuous teeth”, while those of
subsp. alternifolius have margins with “conspicuous
Webbia: Journal of Plant Taxonomy and Geography 191

Figure 3. Cyperus alternifolius subsp. flabelliformis / subsp. alternifolius: subsp. flabelliformis: Habitus (1a, 1b), culm section (2a),
spike (3), spikelet (4), glume (5), nutlet + style (6a) / subsp. alternifolius: Culm section (2b), nutlet (6b).
192 F. Verloove

teeth”. However, both Baijnath (1975) and Kukkonen
(1990) give “glabrous”(as opposed to “scabrid”) bract
margins for subsp. alternifolius and subsp. flabelliformis,
respectively. This is congruent with Rottboell’s extensive
protologue of C. flabelliformis:“(…) involucrum univer-
sale (…) margine & carinis retrorsum scabra (…)”
(Rottboell 1773). Baijnath (1975) and Kukkonen (1990)
state that the anthela rays are “rough or occasionally
rough”in subsp. flabelliformis. However, in all speci-
mens seen of both subsp. alternifolius and subsp. flabell-
iformis, the rays of the anthela are smooth.
For convenience, major distinguishing characters for
both taxa are summarized in the following couplet:
1. Nutlet obovoid, only slightly longer than wide, c. 0.4–
0.8 mm long ( less than half glume length). Culm below
inflorescence slightly to definitely scabrous, ridged-striate
when dry, rarely smooth and remaining more or less terete
…....................................................... subsp. flabelliformis
Nutlet elliptic, obviously longer than wide, c. 1.1–
1.4 mm long (more than half glume length).
Culm below inflorescence always perfectly smooth,
more or less terete and unridged when dry
…..................................... ………… subsp. alternifolius
The taxonomy of C. alternifolius and related taxa has
always been very controversial. Subsp. flabelliformis was
accepted as such by Kükenthal (1935–1936) but was only
given varietal rank by Koyama (1961). Some authors even
considered both as merely conspecific (e.g. Hooper 1972).
Baijnath (1975) demonstrated that these taxa should be
separated at species rank and his point of view was fol-
lowed by most recent authors (e.g. Tucker et al. 2002;
Dai et al. 2010a). However, as shown above and in the
identification key, distinguishing features are very subtle
and quantitative rather than qualitative. The range of mor-
phological variation of the entire C. alternifolius complex
is uncertain (see also Kukkonen 1990). In the absence of
qualitative characters for their separation both are proba-
bly best accepted as subspecies of one variable species.
Cyperus alternifolius s.str. is a critical species. It was
long thought to be very rare and confined to Madagascar
and Réunion and not or hardly cultivated outside its area of
origin (Baijnath 1975). However, in recent times it became
evident that it is also, though much less widely and fre-
quently than subsp. flabelliformis,cultivatedasanorna-
mental in warm-temperate and tropical regions of the world
(e.g. Dai et al. 2010a). In the study area a few populations
from Corsica and the Azores (São Miguel) (listed sepa-
rately below) deviate from the other populations of subsp.
flabelliformis in having culms that are terete and perfectly
smooth below the inflorescence, a character usually associ-
ated with subsp. alternifolius. However, these plants do not
differ from subsp. flabelliformis in nutlet characters. This
observation seems to corroborate with Koyama’sstatement
that nutlet versus scale length is the only reliable character
for the separation of both taxa (Koyama 1961).
Cyperus alternifolius subsp. flabelliformis grows in
shallow water in drainage channels, wetlands and other (at
least temporarily) wet habitats. However, in the study
area it is most extensively naturalized and locally very
invasive in dried-out riverbeds and ravines (“barrancos”).
Selection of specimens examined (typical)
AZORES –Terceira: Angra, near the fortress,
05.07.1972, A. Hansen 375 (C). BALEARIC
ISLANDS –Mallorca: Soller, au Puente de las Animas,
dans le barranco, 13.08.1971, J. Duvigneaud 71 E 512
(BR). CANARY ISLANDS –Tenerife: Sauzal, Los
Angeles, abords des urbanisations, échappé de culture,
26.03.1972, J. Duvigneaud 72 Can 307 (BR); Tenerife:
Santa Ursula, fosse à sec, 26.04.1972, L. Delvosalle s.n.
(BR); Tenerife: Igueste de Candelaria, north of Candelar-
ia, ditch near the village, 24.03.2004, F. Verloove 5570
(BR); La Palma: Fuencaliente, Böschung neben Hotel
Prinzess de la Palma, Palmenfuss, 14.08.2010, R. Otto
17172 (BR); La Palma: San Andrés, Barranco del Agua,
im Bachbett c. 100 m vor Mündung, 08.05.2012, R. Otto
19157 (BR); Gran Canaria: La Sorrueda, embalse de Ti-
rajana, exposed pond margin, common, 07.11.2012, F.
Verloove 9911 (BR); La Palma, Los Sauces, Puerto Es-
pindola, unter Felswand am Hafen, 24.05.2013, R. Otto
20212 (BR); La Palma: Los Sauces, Puerto Espindola,
Wasserrinne neben Straße, unter überrieselter Felswand,
24.05.2013, R. Otto 20228 (BR); La Palma, Los Sauces,
San Andrés, Bco. del Agua, 150 m vor Mündung,
Bachbett, 26.05.2013, R. Otto 20261 (BR); La Palma,
Los Llanos, Argual, nahe Straßenkreuz vor Casa Mas-
sieu, Ablaufrinne, 29.05.2013, R. Otto 20,337 (BR);
Gran Canaria: Maspalomas towards Ayagaures, barranco
de Ayagaures, gravelly riverbed, commonly naturalized,
15.09.2013, F. Verloove 10,584 (BR); Tenerife: Igueste
de San Andrés, barranco de Igueste, close to the village,
gravelly riverbed, very invasive, 16.03.2014, F. Verloove
10632 (BR); Tenerife: Golf del Sur, barranco del Salta-
dero, dry, gravelly riverbed, scattered populations,
20.03.2014, F. Verloove 10633 (BR); Tenerife: San Juan
de La Rambla, Las Aguas, roadside ditch, 22.03.2014, F.
Verloove 10645 (BR); Tenerife: Los Abrigos, Blas (nat-
ure reserve), lake margin, invasive, 22.06.2014, F. Verlo-
ove 10810 (BR); Tenerife: Playa Paraiso, barranco del
Pinque, dry riverbed, near sewage work, 23.06.2014, F.
Verloove 10829 (BR); Tenerife: Callao Salvaje, barranco
de las Barandas, dry gravelly riverbed, 24.06.2014, F.
Verloove 10833 (BR); Tenerife: Tejina, barranco Aguas
de Dios, damp area, ruderalized, 27.06.2014, F. Verloove
10851 (BR); Tenerife: Santa Cruz de Tenerife, Valle
Seco, barranco de Valle Seco, riverbed, locally very
invasive, 29.06.2014, F. Verloove 10874 (BR); Tenerife:
Los Silos, barranco de Cuevas Negras o del Agua, river-
bed, 30.06.2014, F. Verloove 10863 (BR); Tenerife: Los
Silos, barranco de Cuevas Negras o del Agua, riverbed,
depauperate plants, among normally developed
individuals, 30.06.2014, F. Verloove 10864 (BR);
Webbia: Journal of Plant Taxonomy and Geography 193

Figure 4. Cyperus alternifolius subsp. textilis: Habitus (1a, 1b), stem (section) (2), spike (3), spikelet (4), glume (5), nutlet + style (6).
194 F. Verloove

Tenerife: Tabaiba Alta, barranco de las Higueras x TF-
28, gravelly riverbed, 01.07.2014, F. Verloove 10897
(BR). FRANCE –Dep. Landes: Dax, rive de l’Adour,
08.1973, A. Sotiaux s.n. (BR, LG). ITALY –Liguria,
Prov. Savona: in Regione Legna da Cà, strada carroza-
bile dirimpetto all’ex Ospedale Militare, 25.05.19[…],
C·V. Nam s.n. (PAV; sub C. textilis); Emilia-Romagna,
Prov. Parma: Salsomaggiore, 03.08.1955, A. Piazzoli
6774 (MSNM).
Selection of specimens examined (atypical, with culms
terete and smooth)
AZORES –São Miguel: Furnas lake, northern side,
29.06.1971, A. Hansen 69 (C); Pico: Madalena, Quinta das
Roses, cult., 20.05.1980, B. Gonçalves 6191 (C). FRANCE
(incl. Corsica) –Dep. Haute-Corse: Bastia, sortie sud de la
ville (bord de la route de corniche, non loin de la petite
station ferroviaire de Lupino), talus rudéralisé en bordure
d’une ancienne propriété, 19.03.1984, J. Lambinon 84/Co/
15 (LG); Dep. Corse-du-Sud: Golfe de Santa Manza, plage
de Maora, large fossé à Typha en arrière-plage sableuse,
naturalisé en vastes populations, 02.06.1997, J. Lambinon
97/Co/121 et G. Van Den Sande (LG).
subsp. textilis (Thunb.) F. Verloove, comb. nov.
(Figure 4)
(≡)Cyperus textilis Thunb., Prodr. Pl. Cap.: 18. 1794.
Native: South Africa. Naturalized in Europe.
From the study area this taxon was probably first
reported from a lake margin in the Azores (island São
Miguel: Sete Cidades), where it was thought to be a
recent introduction in the early 1940s (Cedercreutz 1941;
Table 1. Modern sectional placement of Cyperus in the study area as compared with Kükenthal (1935–1936).
Cyperus species Laridon et al. (2011), Reynders et al. (2011) Kükenthal (1935–1936)
C. aggregatus Umbellati Umbellati
C. alopecuroides Alopecuroidei Exaltati
C. alternifolius subsp. flabelliformis Alternifolii Vaginati
C. alternifolius subsp. textilis Alternifolii Vaginati
C. brevifolioides Brevifoliae Kyllinga
C. brevifolius Brevifoliae Kyllinga
C. capitatus Galilea Bobartia
C. congestus Fasciculati Thunbergiana
C. croceus Umbellati Umbellati
C. cyprius Cyperus (?) Fusci
C. difformis Fusci Fusci
C. distachyos Juncellus Laevigati
C. eragrostis Luzuloidei Luzuloidei
C. esculentus Cyperus Esculenti
C. exaltatus Exaltati Exaltati
C. flavescens Zonati Flavescentes
C. flavidus Globosi Globosi
C. fuscus Fusci Fusci
C. glaber Compressi Compressi
C. glomeratus Distantes Distantes
C. gracilis Graciles Graciles
C. hamulosus Aristati Aristati
C. laevigatus Juncellus Laevigati
C. longus Cyperus Rotundi
C. lupulinus Laxiglumi Laxiglumi
C. michelianus Pygmaei Dichostylis
C. microiria Irioidei Iriae
C. mundii Vestiti Sulcati
C. odoratus Diclidium Feraces
C. pannonicus Juncellus Laevigati
C. papyrus Papyrus Papyrus
C. polystachyos Pycreus Polystachyi
C. pygmaeus Pygmaei Dichostylis
C. reflexus Luzuloidei Luzuloidei
C. retrorsus Umbellati Umbellati
C. rigens Spicati Thunbergiana
C. rotundus Cyperus Rotundi
C. rubicundus Monocephali Rupestres
C. schweinitzii Laxiglumi Laxiglumi
C. serotinus Stoloniferi Serotini
C. sesquiflorus Brevifoliae Kyllinga
C. squarrosus Aristati Aristati
C. strigosus Strigosi Strigosi
Webbia: Journal of Plant Taxonomy and Geography 195

Palhinha 1966). It is here also reported for the first time
from the island of Corfu. While it is still present and
locally naturalized in the Azores (“persistent”, according
to Schäfer 2003), it probably disappeared from Corfu as
even for C. alternifolius, with which species it has been
confused, there seem to be no recent records (e.g.
Jiménez-Mejías and Luceño 2011).
Cyperus textilis is part of the C. alternifolius complex
and is, like C. involucratus, poorly distinguished from it.
Most authors separate it without difficulty on behalf of
its involucral bracts: these are usually c. 12 in number
(as in Thunberg’s original diagnosis: “involucro dodeca-
phyllo”) and short (rarely more than 10 cm long) with
acute, pungent apices. However, their distinction is rarely
straightforward and merely based on quantitative
characters. Especially depauperate specimens of
C. alternifolius, with which species it shares the terete,
smooth upper culms, are much reminiscent and perhaps
indistinguishable. According to Gordon-Gray (1995)
these distinguishing features need confirmation because
for Natal plants they are not reliable and she even won-
dered if both could be conspecific. Other features that
are sometimes claimed appear to be of no worth or are
simply erroneous.
Careful examination of numerous collections, from
within the study area as well as from Africa, has
demonstrated that C. textilis and C. alternifolius s.str.
are indeed very similar. Typical plants of both are
readily told apart but others are hard to assign to
either. Thunberg’s type collections (no. 1506 and
1507, preserved in UPS) also add to the confusion:
while no. 1507 more or less corresponds with the cur-
rent concept of C. textilis (relatively short and few
involucral bracts) is no. 1506 very atypical (with c.15
bracts that are much longer than they should be). This
collection hardly differs from C. alternifolius s.str. As
both species seem to be mere extremes of one vari-
able species, a lower taxonomic rank seems to be
more appropriate. Surprisingly, C. textilis never seems
to have been combined under C. alternifolius before.
For reasons of consistency (cf. subsp. flabelliformis), it
is here reduced to subspecies rank. Koyama (1961)
erroneously accommodated C. textilis in subgenus
Chloristachys C.B. Clarke (with spikelets arranged
along a distinct spike), while C. alternifolius (incl. var.
obtusangulus; syn.: subsp. flabelliformis) was placed in
subgenus Pycnostachys C.B. Clarke (with spikelets
digitately arranged).
In the study area, C. alternifolius subsp. textilis is
only known from exposed pond margins.
Selection of specimens examined
AZORES –São Miguel: Furnas, naturalized at the lake,
01.09.1970, A. Hansen s.n. (C); São Miguel: Sete Cid-
ades, Lagoa Azul, 30.06.1971, A. Hansen 132 (C); São
Miguel: Sete Cidades, at the shore of Lagoa Azul,
02.07.1971, A. Hansen 201 (C); São Miguel: SV-siden
af Lagoa Azul, 09.08.2002, M. Thornberg s.n. (C).
GREECE –Corfu: bord d’étang, 15.05.1931, H. Rijmers
2676 (BR; sub C. alternifolius).
Section Graciles
Cyperus gracilis R. Brown, Prodr.: 213. 1810 (Figure 5)
Native: Australia. Naturalized in USA and Europe
(Canary Islands).
Cyperus gracilis was discovered as a lawn weed in a
public garden in La Orotava in Tenerife in 2010 and
subsequently confirmed (Siverio Núñez et al. 2013). It is
obviously well established and has extended since its
original discovery but is still limited to the garden.
Similar records are available from California (Tucker
et al. 2002) and Hawaii (Hughes 1995; Oppenheimer
and Bartlett 2002; Starr et al. 2003; Starr and Starr
2011). At least in Hawaii it was initially introduced on
purpose (as groundcover).
Selection of specimens examined
CANARY ISLANDS –Tenerife, La Orotava, centro ciu-
dad (parque), lawn weed, foot of Grevillea tree, small,
established population, 10.09.2010, F. Verloove 8444
(BR); Tenerife, La Orotava, Casa Chana, aggressive
weed in lawns, frequent but not extending beyond gar-
den limits, 02.07.2014, F. Verloove 10871 (BR, ORT).
(II) Subgenus Cyperus
Section Exaltati
Cyperus exaltatus Retz., Observ. Bot. 5: 11. 1789
(Figure 6)
(incl. C. dives Delile, Descr. Égypte, Hist. Nat.: 149.
1812)
Native: Old World tropics. Rarely introduced in the New
World (e.g. Barros 1938).
Cyperus exaltatus was first discovered along the river
Ripoll near Sabadell in Spain (prov. Barcelona) in 2003.
A single, young individual was tentatively ascribed to
C. imbricatus Retz. although it did not fully correspond
with the latter, especially in having wider involucral
bracts, thicker culms and longer inflorescence branches
(Verloove 2005). In 2013 numerous additional plants, in
various stages of development, were seen along the river
Ripoll in the same area (from north of Sabadell up to its
confluence with the river Besòs in Montcada, a stretch
of c. 15 km long). Mature plants turn out to belong to
C. exaltatus, not to C. imbricatus (see also below). This
species is obviously naturalized but apparently long
196 F. Verloove

Figure 5. Cyperus gracilis: Habitus (1), spike (2), spikelet (3), glume (4), nutlet + style (5).
Webbia: Journal of Plant Taxonomy and Geography 197

remained undetected. Like C. congestus, with which spe-
cies it is found growing in close proximity, it may be a
remnant of the alien flora associated with the former
wool-processing industry in the vicinity of Sabadell. It is
also sometimes cultivated as an ornamental and might
represent an escape from cultivation as well.
Along the river Ripoll (and to some extent along the
river Besòs too) C. exaltatus grows on temporarily
Figure 6. Cyperus exaltatus: Habitus (1a, 1b), spike (2), spikelet (3), glume (4), nutlet + style (5).
198 F. Verloove

exposed gravelly and sandy river banks, often
accompanied by other species of Cyperus like
C. congestus,C. eragrostis,C. esculentus,C. longus and
C. rotundus.
In the 1980s and 1990s C. exaltatus was also
recorded along the rivers Cervo and Elvo near Biella in
Italy (initially as C. imbricatus Retz.; Soldano and Sella
2000; Verloove and Soldano 2011). However, it was not
confirmed recently in this area.
Cyperus exaltatus is a member of the small but very
complex section Exaltati and its taxonomy is not undebat-
ed. The two other members of this section [C. dives (incl.
C. immensus C·B. Clarke) and C. imbricatus] are not eas-
ily told apart and the exact placement of a third species,
C. alopecuroides, is also controversial. Boulos (2005)
considers the latter to be conspecific with C. dives,as
characters separating these species are not constant. How-
ever, although C. alopecuroides superficially resembles
C. dives both are readily distinguished on glume, nutlet
and stigma characters. Moreover, both are currently
accommodated in different sections (Larridon et al.
2011a). According to Hoenselaar et al. (2010)C. dives
and C. exaltatus are differentiated on quantitative rather
than qualitative characters; these authors therefore
re-instated varietal rank for C. dives (see also Prasad and
Singh 2002). The separation of these species indeed
appears to be critical: C. dives has densely clustered spik-
elets that are obliquely inserted, while those of C. exalta-
tus are much laxer (individual spikelets and rachis readily
discerned) and inserted at almost right angles. However,
var. iwasakii (Makino) T. Koyama and var. megalanthus
Kük. of the latter (Kükenthal 1935–1936; Koyama 1961;
see also Dai et al. 2010a) are characterized by very den-
sely arranged, larger and multi-flowered spikelets, obscur-
ing the rachis and seem to link both species. The
populations currently found near Barcelona combine fea-
tures of both these species, e.g. turgid, golden yellow spik-
elets and relatively small glumes less than 1.5 mm long of
C. dives (Haines and Lye 1983; Prasad and Singh 2002),
while the untidily arranged spikes and spikelets are more
characteristic of C. exaltatus. Therefore, European
populations are here ascribed to C. exaltatus s.l., incl.
C. dives.
Finally, confusion is also likely with C. imbricatus.
All recent southern European claims of the latter (Soldano
and Sella 2000; Verloove 2005) are referable to C. exalta-
tus s.l. (incl. C. dives; see Verloove and Soldano 2011).
Cyperus imbricatus is a more slender species (rarely
exceeding 100 cm) with narrower leaves and spikes (resp.
4–8 mm versus at least 10 mm and 3–8 mm versus
6–15 mm in mature specimens of C. exaltatus), shorter
and paler glumes (0.8–1.2 mm versus 1.2–1.8 mm) with a
conspicuously excurved mucro (versus straight or incur-
ved mucro). See Verloove and Soldano (2011) for an
extended discussion.
Cyperus exaltatus is found in shallow water by rivers
and on gravelly, exposed river shores.
Selection of specimens examined
ITALY –Prov. Biella: a valle di Biella, rivoletto che
confluisce nel Cervo, 15.10.1989, A. Soldano 6777
(priv.herb. A. Soldano); Prov. Biella: ad ovest di Ma-
ghetto di Mongrando, sponda destra dell’Elvo, 09.1993,
A. Soldano 8983 (priv.herb. A. Soldano). SPAIN –Prov.
Barcelona: Barbera del Vallès (Santa Maria de Barbera),
north of Pont de Santiga, gravelly border of riu Ripoll,
one tall caespitose plant, 10.09.2003, F. Verloove 5485
(BR, GENT, MA; sub C. imbricatus); Prov. Barcelona:
Barbera del Vallès, riu Ripoll south of railway bridge,
gravelly river bank, 2 plants, 21.08.2013, F. Verloove
10415 (BR); Prov. Barcelona: Cerdanyoles del Vallès, riu
Ripoll at footbridge, gravelly river bank, two very robust
plants, 21.08.2013, F. Verloove 10425 (BR, BC, MA);
Prov. Barcelona: Ripolett, riu Ripoll, west of Carrer
Limit Poligon la Ferrer (bridge), gravelly river bank,
scattered individuals, 21.08.2013, F. Verloove 10426
(BR, BC); Prov. Barcelona: El Masrampinyo (Montca-
da), riu Ripoll, close to its junction with riu Besòs, river
border, rather frequent, several tens, 22.08.2013, F. Ver-
loove 10434 (BC); Prov. Barcelona: Montcada, riu
Besòs, close to its junction with riu Ripoll, sandy river
bank, several plants, 22.08.2013, F. Verloove 10446
(BR).
Section Irioidei
Cyperus microiria Steud., Syn. Pl. Glumac. 2: 23. 1855
(Figure 7)
Native: East Asia. Naturalized in U.S.A. and Europe.
Cyperus microiria was first collected in Italy in 1908 in
Porto di Malgrate (province of Como) (Camperio and
Fiori 1910). This record was from a garden so was not
related to its naturalization in rice fields, which clearly
represents a later, independent and more successful intro-
duction. The species was not yet known to Jacometti
(1912). Koch (1952) and Stucchi (1969,1972) probably
were the first to report about its occurrence as a rice field
weed in Italy, although Koch (1952) erroneously
ascribed it to the closely related C. amuricus Maxim. It
remained rare and local for quite a long time and early
records, for instance those by Becherer (1969), were
partly erroneous and referable to C. strigosus (Raynal
1977). Even by the end of the 1970s it was still very
rare and confined to the region of Vercelli (Raynal 1977,
1979) although a few years later several additional local-
ities (and a detailed chronological overview of the spe-
cies’naturalization history in Italy) were provided by
Soldano (1980). In the past decades it has quickly spread
further and now occurs widely in large parts of northern
Italy (e.g. Desfayes 2005; Banfiand Galasso 2010).
According to Celesti-Grapow et al. (2010) it is at present
known from the regions Calabria, Emilia-Romagna,
Friuli-Venezia Giulia, Lombardia, Piemonte and Veneto,
Webbia: Journal of Plant Taxonomy and Geography 199

in some of which it is considered an invasive species.
Especially in its original area of introduction it is locally
very common, e.g. along the river Sesia in the surround-
ings of Vercelli.
In Italy, C. microiria was initially mostly confined to
rice fields. At present it also frequently occurs on
exposed river banks and other temporarily wet and regu-
larly disturbed places.
Figure 7. Cyperus microiria: Habitus (1), spike (2), spikelet (3), glume (4), nutlet + style (5).
200 F. Verloove

In the Euro+Med Plantbase (Jiménez-Mejías and
Luceño 2011) the binomial C. microiria is considered to
be misapplied for C. iria L. Both species are indeed clo-
sely related but distinct (e.g. Kükenthal 1935–1936; Kern
1974; Tucker et al. 2002; Dai et al. 2010a). Raynal
(1977) demonstrated that the plants naturalized in Italy
belong to C. microiria, not to C. iria, nor C. amuricus.
Genuine C. iria was recently reliably recorded in a plant
nursery near Valencia in Spain (Verloove et al. 2014).
Selection of specimens examined
ITALY –Lombardia, Prov. Milano: Cuggiono, ghiaieti
del Ticino, 08.1950, C. Stucchi s.n. (MSNM 234); Lom-
bardia, Prov. Milano: Cuggiono, campi umidi, 09.1960,
C. Stucchi s.n. (MSNM 233); Lombardia, Prov. Pavia:
entro il perimetro delle mura della città di Pavia, via
Lungoticino Visconti, L. Pavan et al. s.n. (PAV); Lom-
bardia, Prov. Pavia: Pieve del Cairo, limoso, bordo
d’aqua fangoso, 25.10.1983, F. Bracco & al. s.n. (PAV;
sub C. glaber); Lombardia, Prov. Pavia: Bereguardo
(northwest of Pavia), lit du Tessin, 05.09.1987, M. Des-
fayes s.n. (GENT); Piemonte, Prov. Vercelli: Oldenico,
bord de rizière, 06.09.1987, M. Desfayes s.n. (GENT);
Piemonte, Prov. Vercelli: Greggio, 04.09.1989, M. Desfa-
yes s.n. (GENT); Piemonte, Prov. Torino: Rivarossa (near
Torino), Malone river, 28.07.1990, M. Desfayes 90–10
(GENT); Piemonte, Prov. Vercelli: Vercelli, rice fields,
17.09.1990, M. Desfayes 90–03A (GENT); Piemonte,
Prov. Vercelli: near Oldenico, Sesia river, summer bed,
17.09.1990, M. Desfayes 90–01 (GENT); Piemonte,
Prov. Novara: Galliate, south of Ponte Turbigo, left bank
of river Ticino, gravelly border of river, common in
this area, 04.09.2004, F. Verloove 5771 (BR); Emilia-
Romagna, Prov. Piacenza: Castelvetro Piacentino, c.31
s.l.m., greto del fiume Po a nord della Cascina Babina,
lanca sponda, 31.08.2006, L. Gariboldi s.n. (MSNM
40666, 40667); Lombardia, Prov. Pavia: Castello d’Ago-
gna, Centro Ricerche sul Riso, c. 105 s.l.m., incolto,
13.10.2007, G. Galasso s.n. (MSNM 41259, 41260);
Emilia-Romagna, Prov. Piacenza: Castelvetro Piacentino
(south Cremona), left bank of river Po, sandy river bank,
08.09.2008, F. Verloove 7346 (BR, LG); Piemonte, Prov.
Vercelli: Arborio, destra idrografica del fiume Sesia a
ovest del centro abitato (UTM ED50 32T04529.50381),
179 s.l.m., greto sabbioso, 12.09.2009, G. Galasso s.n.
(MSNM 43669, 43670); Lombardia, Prov. Pavia: Viale
Lungo Ticino Sforza, left bank of river Ticino (WGS84:
45°10’49" N, 09°09’22" E), sandy bank, 64 m,
25.09.2013, N. Ardenghi et P. Cauzzi s.n. (BR); Pie-
monte, Prov. Vercelli: Arborio, left bank of river Sesia,
exposed river bank, very common, 06.09.2014, F. Verlo-
ove 10982 (BR). Piemonte, Prov. Alessandria: Valenza,
river Po south of the bridge, sandy river bank,
07.09.2014, F. Verloove 10991 (BR); Lombardia, Prov.
Pavia: Pancarana, river Po, sandy river bank,
10.09.2014, F. Verloove 10997 (BR). Lombardia, Prov.
Pavia: Colonne (south Pavia), rice field, 10.09.2014,
F. Verloove 11010 (BR); Lombardia, Prov. Pavia: Bastida
Pancarana, SP 35 close to river Po, maize field, 11.09.2014,
F. Ve r l o o v e 1 1 0 0 5 (BR); Lombardia, Prov. Pavia:
Bereguardo, river Ticino close to the bridge, sandy river
bank, 11.09.2014, F. Ve r l o o ve 1 1 0 1 5 (BR); Lombardia.
Prov. Lodi: San Rocco al Porto, river Po close to SS 9,
muddy river bank, 13.09.2014, F. Ve rl o o v e 11 0 1 3 (BR);
Emilia-Romagna, Prov. Piacenza: Castelvetro Piacentino,
località Maginot, river Po, muddy river bank, 14.09.2014,
F. V e rl o o v e 11 0 2 3 (BR); Lombardia, Prov. Mantova:
Viadana, river Po west of the bridge, sandy river bank, few
specimens, 15.09.2014, F. Ve r l o o v e 11 0 3 0 (BR).
[Mariscus]
Section Strigosi
Cyperus strigosus L., Sp. Pl.: 47. 1753 (Figure 8)
(≡)Mariscus strigosus (L.) C·B. Clarke, Contr. U.S.
Natl. Herb. 10(6): 453. 1908
(=) M. elatus auct. ital. non Vahl, Enum. Pl. 2: 377. 1805
Native: USA. Naturalized in Europe.
Cyperus strigosus was apparently first collected in Italy
in 1896 near Garlasco (province of Pavia) (Mattirolo and
Fiori 1917). By 1923 it was also known from the Torino
province (Mattirolo 1915; Fiori 1923). In the past dec-
ades it was regularly confirmed from the rice-producing
areas in Piemonte (e.g. Cook 1973; Abbà 1975; Soldano
1976a; Raynal 1979,1981). At present it is still spread-
ing and known as a naturalized or invasive species in
the regions Emilia-Romagna, Lombardia, Piemonte, Tos-
cana and Veneto (e.g. Desfayes 2005; La Rosa et al.
2008; Banfiand Galasso 2010; Celesti-Grapow et al.
2010; Masin and Scortegagna 2012). In 2014 the genu-
ine presence of C. strigosus in the regions Emilia-Ro-
magna, Lombardia and Piemonte was confirmed (see
“Specimens examined”). However, many records surely
require confirmation since it has widely been confused
with C. odoratus in northern Italy (see also under that
species). Cyperus strigosus was locally temporarily
established in Spain in the twentieth century. It was dis-
covered in 1939 in the Girona province near Hostalric
(Pla d’en Marqués) alongside a small canal (Llensa de
Gelcén 1941). Soon afterwards it was also seen at sev-
eral places on the exposed banks of the river Tordera
between Hostalric and its estuary near Blanes (e.g.
Llensa de Gelcén 1945; Montserrat 1964; Casasayas
1989). These plants were initially ascribed to the African
C. auricomus Sieber ex Spreng. [syn.: C. digitatus Roxb.
subsp. auricomus (Sieber ex Spreng.) Kük.] and were
included under this name in the Flora Europaea
(DeFilipps 1980). However, they are readily
distinguished from the latter in having wider, flattened
spikelets that fall as a whole (not subterete and
c. 0.6–0.9 mm wide and with deciduous glumes), longer
Webbia: Journal of Plant Taxonomy and Geography 201

Figure 8. Cyperus strigosus: Habitus (1a, 1b), spike (2), spikelet (3), glume (4), nutlet + style (5).
202 F. Verloove

glumes (c. 3.5 mm versus 1.9–2.4 mm) and shorter
anthers (c. 0.3–0.5 mm versus 0.9–1.6 mm). Castroviejo
(2005), while preparing the account for Cyperus in the
Flora Iberica, found out that they in fact belong to the
similar but American C. strigosus. Unfortunately, shortly
afterwards, this determination was incorrectly altered to
C. odoratus (Castroviejo 2006). Only populations from
the rivers Segre and Ebro belong to the latter (see under
that species). According to Vilar (1987)C. strigosus has
disappeared from the river Tordera, possibly already in
the 1950s, more precisely after the cold winter of 1956.
Recent claims of Cyperus strigosus from Bulgaria
(Tzonev et al. 2003) and Serbia and Montenegro
(Stevanovićet al. 2005) turn out to be in error for
C. odoratus (see under that species).
Cyperus strigosus has widely been confused with
C. odoratus, at least in Bulgaria, Italy, Spain and former
Yugoslavia. At maturity both species are readily distin-
guished. Cyperus odoratus has spikelets that fall apart in
numerous units at maturity, each containing a glume and
part of the indurated and corky rachis that tightly
encloses the nutlet. This is a unique feature in Cyperus;
therefore this species was formerly accommodated in a
separate genus, Torulinium. At the flowering stage their
separation is less straightforward although C. odoratus
has subcylindrical spikelets (versus distinctly flattened
ones in C. strigosus), shorter glumes (c. 2.2–2.8 mm ver-
sus 3.2–4.5 mm) and shorter nutlets (c. 1.2–1.5 mm ver-
sus 1.8–2.4 mm). Also, C. strigosus initially has greenish
yellowish inflorescences (turning golden at maturity)
while these are golden from the beginning in C. odora-
tus. Finally, C. strigosus is a rhizomatous perennial (but
often flowering in the first year) while C. odoratus is
usually an annual although European plants often seem
to behave like short-lived perennials. According to
Raynal (1979)C. strigosus has furthermore been con-
fused in Italy with C. microiria. Both species only share
the typical golden colour of the spikelets (at least at
maturity) but are otherwise very different.
At present C. strigosus is mostly found on exposed
gravelly river shores and in drainage channels. It is still
sometimes found as a weed in rice fields.
Selection of specimens examined
ITALY –Lombardia, Prov. Milano: presso Besate (sul
Ticino), 08.10.1967, F. et E. Ghisotti 9/137 (MSNM);
Piemonte, Prov. Vercelli: Oldenico, rizières de la rive
gauche du Cervo, à l’est de la route de Vercelli, alt.
130 m, bords de fossés, par taches çà et là, 12.09.1978,
J. Raynal 20886 (BR, GENT); Lombardia, Prov. Varese:
Brinzio, 08.1979, Peracchio s.n. (MSNM 10253); Pie-
monte, Prov. Vercelli: Oldenico, bord de rizière,
06.09.1987, M. Desfayes s.n. (GENT); Piemonte, Prov.
Torino: Rivarossa (near Torino), Malone river,
28.07.1990, M. Desfayes 90–07 (GENT); Piemonte,
Prov. Vercelli: Vercelli, left bank of river Sesia, south of
railway bridge, muddy bank of river, scattered speci-
mens, 05.09.2004, F. Verloove 5773 (BR); Piemonte,
Prov. Torino: San Giorgio Canavese, SP 40, ditch,
07.09.2009, F. Verloove 7770 (BR); Piemonte, Prov. Tor-
ino: Chivasso, river Orco, muddy riverbank, 14.09.2009,
F. Verloove 7895 (BR, LG, TO); Piemonte, Prov. Torino:
San Sebastiano da Po, north side of river Po, muddy riv-
erbank, 14.09.2009, F. Verloove 7896 (BR); Piemonte,
Prov. Vercelli: Vercelli, left bank of river Sesia north of
the railway bridge, gravelly river bank, scattered individ-
uals, 07.09.2014, F. Verloove 10988 (BR); Piemonte,
Prov. Alessandria: Valenza, river Po south of the bridge,
sandy river bank, 07.09.2014, F. Verloove 10992 (BR);
Lombardia, Prov. Pavia: Pancarana, river Po, sandy river
bank, a single individual, 10.09.2014, F. Verloove 10999
(BR); Lombardia, Prov. Pavia: Bereguardo, river Ticino
close to the bridge, sandy river bank, 11.09.2014, F. Ver-
loove 11008 (BR); Lombardia, Prov. Pavia: Robbio,
north of the city, rice fields, 12.09.2014, F. Verloove
11003 (BR); Lombardia, Prov. Lodi: San Rocco al Porto,
river Po close to SS 9, muddy river bank, 13.09.2014, F.
Verloove 11012 (BR); Emilia-Romagna, Prov. Piacenza:
Castelvetro Piacentino, località Maginot, river Po, muddy
river bank, 14.09.2014, F. Verloove 11024 (BR).
SPAIN –Prov. Barcelona: Gualba, Montseny, vores hor-
tis, 09.11.1917, Font i Quer (BC 63593; sub C. aurico-
mus); Prov. Girona: Hostalric, Pla d’en Marquès, llocs
humids, 09.1939, S. Llensa de Gelcén (BC 92707; sub
C. auricomus); Prov. Girona: Hostalric, algo más abajo
del puente del Arrabal, a la orilla izquierda del río Tor-
dera, 25.08.1940, S. Llensa de Gelcén (BC 92709; sub
C. auricomus).
Section Spicati
Cyperus rigens J. Presl & C. Presl, Reliquiae Haenke-
anae 1(3): 170. 1828 (Figure 9)
(≡)Mariscus rigens (J. Presl & C. Presl) C.B. Clarke,
Bull. Herb. Boissier 6 (App. 1): 20. 1898.
Native: tropical and subtropical America (Argentina,
Brazil, Paraguay, Uruguay). Naturalized in Europe.
Cyperus rigens was first discovered in Tarnos-Le Boucau
(Lac Lahoun) in France in 1975 (Vivant 1978) (det. J.
Raynal). At present it is well-established there in a small
coastal area immediately north of Bayonne (Vivant 2003).
Field observations in 2013 showed that C. rigens is pres-
ent to the north up to Capbreton and more inland localities
were detected in Orx. It occurs on the margins of exposed
ponds, in damp coastal grassland and marshes, by riverlets
and in other similar wet habitats.
Previous Italian claims of C. rigens (from Sesia river;
see Soldano 1976b,1980) were referable to C. congestus
(Soldano 2000). See also under that species for
distinguishing features. However, a single collection
from the 1950s from the river Ticino in Sesto Calende in
Webbia: Journal of Plant Taxonomy and Geography 203

Figure 9. Cyperus congestus /Cyperus rigens:C. congestus: Habitus (1a, 1b), stem (section) (2), spike (3), spikelet (4a), glume (5),
nutlet + style (6) / C. rigens: Spikelet (4b).
204 F. Verloove

Varese province, initially erroneously identified as
C. glomeratus, belongs to C. rigens. This species should
be looked for along the river Ticino.
All specimens seen have smooth culms and belong
to subsp. rigens (Hefler 2010;Hefler and Longhi-Wagner
2012).
Selection of specimens examined
FRANCE –Dep. Landes: au nord de Bayonne, bords de
l’étang de Lahaun, 19.07.1975, G. Van Buggenhout (BR;
sub C. congestus); Dep. Landes: en aval de Bayonne,
alluvions de l’Adour, 19.07.1975, N. Cnops 75120 (BR;
sub C. congestus); Dep. Landes: Ondres ten N van
Bayonne, oever Lac d’Irieu, 19.07.1979, J.E. De Langhe
422/79 (BR; sub C. eragrostis); Dep. Landes: à 4 km au
Sud de Capbreton, rive gauche du canal du Boudigau,
sables frais, 26.06.1982, J. Vivant s.n. (priv. herb. A.
Soldano); Dep. Landes: Tarnos, N-Bayonne, Lac Lahoun,
marshy area in the dunes, 10.08.1999, F. Verloove 3825
(BR); Dep. Landes: Tarnos, near Métro, les Lacs Lahoun,
marshy area in dunes, 16.06.2004, F. Verloove 5631 (BR);
Dep. Landes: Orx, marais d’Orx (nature reserve), ditch,
common (with C. eragrostis), 17.08.2013, F. Verloove
10431 (BR); Dep. Landes: Capbreton, riverlet Boudigau
south of the village, riverlet margin, locally frequent,
17.08.2013, F. Verloove 10469 (BR, P). ITALY –Lom-
bardia, Prov. Varese: Sesto Calende, lungo Ticino,
14.08.1951, s.coll. (PAV; sub C. glomeratus).
Section Thunbergiana
Cyperus congestus Vahl, Enum. Pl. 2: 358. 1805
(Figure 9)
(≡)Mariscus congestus (Vahl) C.B. Clarke, J. Bot. 35:
72 1897.
Native: South Africa. Naturalized in Europe, Asia
(Japan), Australia.
Cyperus congestus has been known for quite a long time
from various parts of Europe. It used to be, for instance, a
fairly characteristic alien associated with the wool-pro-
cessing industry in several different countries (e.g. Ryves
1976; Kiffe 2004; Verloove 2006a). These occurrences,
however, always proved to be ephemeral. According to
DeFilipps (1980) it was naturalized in Portugal near
Coimbra, where it was known since the end of the nine-
teenth century (Daveau 1891; Almeida 1999). Recent
inquiries, however, demonstrated that it was probably
never genuinely naturalized in Portugal or that it is now
long gone (Castroviejo 2005,2007a). In Italy C. conge-
stus has also been recorded on various occasions (Fiori
1923) but there also probably long remained merely
ephemeral. However, from the 1970s onwards
(e.g. Soldano 1976b,1980; Raynal 1979; in part as
C. rigens) it was increasingly reported but seemed to be
confined to the exposed banks of the river Sesia in the
regions Lombardia and Piemonte (Celesti-Grapow et al.
2010; Banfiand Galasso 2010). It was not confirmed
lately from its single locality in Lombardia (in Palestro)
but it is still firmly established elsewhere beside the river
Sesia in Piemonte, especially in the surroundings of Ver-
celli (pers. obs. author; see also: Desfayes 2005).
In 2013 C. congestus was also repeatedly seen near
Barcelona in Spain, along the river Ripoll between Saba-
dell and Montcada (a stretch of about 15 km) and by the
river Besòs, close to its confluence with the river Ripoll.
In this area it is obviously naturalized but apparently
remained undetected so far. It may be associated with
the former wool-processing industry in the surroundings
of Sabadell (see also under C. exaltatus) although it
might also be an escape from cultivation.
In recent times C. congestus is regularly cultivated as
an ornamental and, like C. eragrostis, easily escapes
from cultivation. In suitable habitats it may well be able
to naturalize.
In the study area C. congestus most closely resembles
the South American C. rigens. Both are rather robust
short-lived perennials with clearly pedunculate, con-
tracted, globose spikes with spikelets arranged along a
rachis. In Italy both species have been confused for some
time (Soldano 1976b,1980,2000; Abbà 1980). They
were both accommodated by Kükenthal (1935–1936)in
section Thunbergiana of subgenus Mariscus and distin-
guished on spikelet form: linear (c. 10× longer than wide)
in C. congestus versus oblong-lanceolate (c. 5× longer
than wide) in C. rigens. Moreover, glumes of the former
always are darker (dark red to maroon).
In Italy as well as Spain C. congestus is naturalized
on exposed gravelly river banks.
Selection of specimens examined
ITALY –Piemonte, Prov. Vercelli: Vercelli, riva sinistra
del Sesia, 04.08.1975, A. Soldano 756b (PAV; sub. C.
rigens); Piemonte, Prov. Vercelli: near Oldenico, Sesia
river, summer bed, 17.09.1990, M. Desfayes 90–06
(GENT); Piemonte, Prov. Vercelli: Vercelli, left bank of
river Sesia, south of railway bridge, stony border of
river, rare, 09.09.2004, F. Verloove 5775 (BR);
Piemonte, Prov. Vercelli: Ghislarengo, right bank of
river Sesia (S of the bridge), gravelly river bank,
locally rather common, 15.09.2008, F. Verloove 7350
(BR); Piemonte, Prov. Vercelli: Arborio, destra idrogra-
fica del fiume Sesia a ovest del centro abitato (UTM
ED50 32T 04529.50381), 179 s.l.m., greto sabbioso,
12.09.2009, G. Galasso s.n. (MSNM 43662, 43663,
43664; sub C. strigosus); Piemonte, Prov. Vercelli:
Arborio, left bank of river Sesia, exposed river bank,
locally, 06.09.2014, F. Verloove 10984 (BR). SPAIN –
Prov. Barcelona: Sabadell, Can Grau, riu Ripoll at
Torre-romeu, north of footbridge, grassy river bank,
dense populations, several tens, 19.08.2013, F. Verloove
10451 (BR, BC, MA); Prov. Barcelona: Barbera del
Vallès, riu Ripoll south of railway bridge, gravelly river
Webbia: Journal of Plant Taxonomy and Geography 205

bank, 3 plants, 21.08.2013, F. Verloove 10416 (BR);
Prov. Barcelona: Cerdanyoles del Vallès, riu Ripoll
north of footbridge, dry gravelly river bed, two plants,
21.08.2013, F. Verloove 10424 (BR); Prov. Barcelona:
El Masrampinyo (Montcada), riu Ripoll, close to its
junction with riu Besòs, river border, c. 10 plants,
22.08.2013, F. Verloove 10433 (BR); Prov. Barcelona:
Montcada, riu Besòs, close to its junction with riu
Ripoll, sandy river bank, several plants, 22.08.2013, F.
Verloove 10445 (BR).
Figure 10. Cyperus lupulinus /Cyperus schweinitzii:C. lupulinus: Habitus (1), spike (2a), glume (4a) / C. schweinitzii: Spike (2b),
Culm (3), Glume (4b).
206 F. Verloove

Section Laxiglumi
Cyperus lupulinus (Spreng.) Marcks, Trans. Wisconsin
Acad. Sci. 62: 271. 1974 (Figure 10)
Native: USA. Naturalized in Europe.
Cyperus lupulinus was recently discovered in Italy on dry,
gravelly soil by railway tracks in Chivasso (province of
Torino) (Verloove and Soldano 2011). This species grows
in similar circumstances as the related C. schweinitzii and
a future, wider naturalization seems not unlikely.
Both C. lupulinus and C. schweinitzii are perennials
with very distinct, corm-like rhizomes and are very simi-
lar in general appearance. However, the former has
glumes with a shorter mucro (c. 0.05–0.2 mm long),
smaller anthers (c. 0.3–0.6 mm long), culms that are
smooth below the inflorescence, a more compact inflo-
rescence with rays usually absent (rarely 1–4, 10–60 mm
long) and its longest inflorescence bract is horizontal to
reflexed, not erect.
Selection of specimens examined
ITALY –Prov. Torino: Chivasso, west of the city, rail-
way yard, bare, gravelly soil, scattered specimens, along
with Euphorbia davidii, 14.09.2009, F. Verloove 7870
(BR, TO).
Cyperus schweinitzii Torr., Ann. Lyceum Nat. Hist.
New York 3: 276. 1836 (Figure 10)
Native: USA, Mexico. Naturalized in Europe.
Cyperus schweinitzii was probably first reported from
Europe in the 1960s (railway track in Mulhouse, France;
see Litzler 1965). Although relatively numerous, the spe-
cies probably disappeared soon afterwards. In 1990 it
was discovered, also in France, in an abandoned railway
yard in Roanne where it could have been introduced
before 1961 (Bosc and Foirest 1990; Foirest et al. 1996).
In the past decades C. schweinitzii was also found on
several occasions in and around Bordeaux (Le Bouscat,
Lormont, La Bastide and Saint-Louis) (Dupain 2002;
Aniotsbéhère and Dussaussois 2003; J.-C. Aniotsbéhère,
2013 pers. comm.). Also in this area (Bassens:
Appontements) an additional large population was dis-
covered in August 2013 in a more or less disused part of
a railway yard. Recently it was furthermore discovered
in a railway yard in Alès (comm. J.-M. Tison). In all
these localities C. schweinitzii looks firmly established
and a future spread seems very likely.
In 2012 C. schweinitzii was found for the first time
in Italy (Ardenghi 2013). A large, obviously naturalized
population was detected in a railway yard in Stradella.
Elsewhere in Europe (e.g. in the Netherlands; see “Speci-
mens examined”)C. schweinitzii appears to be an
ephemeral introduction.
Cyperus schweinitzii is confined to gravelly, sun-
exposed, dry or only temporarily wet habitats and is
nearly restricted to railway yards. However, in Bordeaux
it has also been recorded on the exposed banks of the
river Garonne.
Section Laxiglumi is an assemblage of closely related,
predominantly American species, several of which with
weedy tendencies. In addition to C. lupulinus and
C. schweinitzii, both naturalized in the study area, other
members may be expected. Cyperus filiculmis Vahl has
been recorded as an alien in the Netherlands (Weevers
et al. 1956) and C. houghtonii Torr. in several different
European countries (e.g. in Poland; Mirek 1995).
Selection of specimens examined
FRANCE –Dep. Loire: Roanne, ancien triage de la gare
SNCF de Roanne, désaffecté depuis 1961, alt. 280 m, sur
sable mélangé à la poussière de charbon sur plusieurs cen-
timètres sur le ballast, 15.09.1995, G. Dutartre s.n. (BR,
LG, MSNM 36999); Dep. Gironde: Bassens (northeast
Bordeaux), railway yard Appontements (port area), by
railway tracks, locally very numerous, 15.08.2013, F. Ver-
loove 10421 (BR, P); Dep. Gard: Alès, railway station,
bare, gravelly area (abandoned railway tracks), common,
since several years, 25.08.2013, F. Verloove 10500 (BR,
BC, P). ITALY –Lombardia, Prov. Pavia: Stradella, stazi-
one FS, scalo merci (WGS84: 45°04’44.81" N,
09°17’53.10" E), detrito ghiaioso consolidato tra la massi-
cciata ferroviaria e il piazzale del deposito, copioso, 74 m,
11.07.2012, N. Ardenghi s.n. (BR); Lombardia,
Prov. Pavia: Stradella, railway yard, gravelly area, sev-
eral tens, 11.09.2014, F. Verloove 11007 (BR). NETHER-
LANDS –Prov. Zuid-Limburg: Simpelveld (IFBL
E8.22.34), stationsemplacement, slechts 1 plantje,
22.09.1984, H. Vannerom 840922.10 (LG).
Section Aristati
Cyperus squarrosus L., Cent. Pl. II 6. 1756 (Figure 11)
(≡)Mariscus squarrosus (L.) C.B. Clarke, Fl. Brit. India
6(19): 623. 1893
(=) Cyperus aristatus Rottb., Descr. Pl. Rar. 22. 1772
(=) C. inflexus Muhl., Descr. Gram. 16. 1817
Native: USA, Mexico, West Indies, Bermuda, Central
America, South America, Asia, Africa, Indian Ocean
Islands, Pacific Islands, Australia. Naturalized in Europe.
Cyperus squarrosus was apparently first discovered in
Italy between Mezzana and Montebolone (province of
Pavia), on the left bank of the river Ticino, in July
1895 (Cavara 1899). It remained rare and local for
quite a long time and has also been widely confused
with C. hamulosus (Raynal 1979). In the 1970s, for
instance, it was only recorded near Trecate, Valenza
and Vercelli (Soldano 1976a; Raynal 1979). At present,
Webbia: Journal of Plant Taxonomy and Geography 207

it is known as a naturalized, non-invasive alien in the
Italian regions Emilia-Romagna, Lombardia and Pie-
monte (e.g. Assini et al. 2005; Desfayes 2005; Banfi
and Galasso 2010; Celesti-Grapow et al. 2010) and
recently also reached the Veneto region (Masin and
Scortegagna 2012). It is also still present in its
original area of introduction (comm. N. Ardenghi
2013).
Figure 11. Cyperus squarrosus: Habitus (1), spike (2), spikelet (3), glume (4), nutlet + style (5).
208 F. Verloove

In 2013 it was discovered for the first time in
Croatia near Motovun, on an exposed sandy river bank
(Rottensteiner 2014). According to Cavara (1899)
C. squarrosus was most likely initially introduced as
an impurity in rice. At present, it is hardly seen as a
rice field weed and is much more frequent now on
temporarily wet, sandy or gravelly exposed river
banks.
Figure 12. Cyperus aggregatus: Habitus (1), spike (2), spikelet (3), glume (4), nutlet + style (5).
Webbia: Journal of Plant Taxonomy and Geography 209

In Euro+Med Plantbase (Jiménez-Mejías and Luceño
2011)C. squarrosus is not upheld. This binomial is con-
sidered to be a misapplied name (for C. hamulosus).
However, in Italy both species grow sympatrically, at
least in Lombardia (Celesti-Grapow et al. 2010),
C. hamulosus being rare and ephemeral. Both species are
closely related, morphologically similar and often con-
fused. However, on closer examination they are readily
distinguished (see Key).
Selection of specimens examined
ITALY –Lombardia, Prov. Pavia: Mezzana, presso
Pavia, 1895, F. Cavara s.n. (PAV); Lombardia, Prov.
Pavia: Mezzinella, rive del Ticino, 1895, F. Cavara s.n.
(PAV); Lombardia, Prov. Pavia: Pavia, Ticino morto, con
Lindernia dubia, 08.1953, J. Viola s.n. (MSNM 232,
6756, 6757); Lombardia, Prov. Pavia: presso Pavia
(Vul), spiaggia del Ticino, 04.09.1953, A. Pirola s.n.
(PAV); Emilia-Romagna, Prov. Piacenza: Piacenza, lungo
il Po, lama sabbiosa inondata, 10.09.1986, C. Lazioli s.n.
(MSNM 26812); Lombardia, Prov. Pavia: Bereguardo
(northwest Pavia), lit du Tessin, 05.09.1987, M. Desfayes
s.n. (GENT); Piemonte, Prov. Novara: Trecate (San Mar-
tino), right bank of river Ticino, sandy bank of river,
locally, 06.09.2004, F. Verloove 5780 (BR); Lombardia,
Prov. Cremona: Cremona, Colonie Padana, muddy river
bank of river Po, common, 08.09.2008, F. Verloove 7405
(BR); Lombardia, Prov. Cremona: Cremona, Colonie
Padane, sterrato lungo il Po, c. 38 m s.l.m., 08.09.2008,
G. Galasso s.n. (BR, MSNM 41654, 43872); Emilia-Ro-
magna, Prov. Parma: Zibello (southeast of Cremona),
right bank of river Po, muddy river bank, locally (along
with Cyperus michelianus), 09.09.2008, F. Verloove
7425 (BR); Lombardia, Prov. Pavia: Viale Lungo Ticino
Sforza, left bank of river Ticino (WGS84: 45°10’49" N,
09°09’22" E), sandy bank, 64 m, 25.09.2013, N. Arden-
ghi et P. Cauzzi s.n. (BR) [the site where this specimen
was collected is only about 3.5 km west from the site
where C. squarrosus was for the first time collected
in Italy and Europe by F. Cavara; comm. N. Ardenghi];
Lombardia, Prov. Pavia: Viale Lungo Ticino Sforza,
left bank of river Ticino (WGS84: 45°10’51" N,
09°09’17–18" E), abandoned flowerbed (previously
flooded), on wet sandy soil, 66 m, 25.09.2013, N. Arden-
ghi et P. Cauzzi s.n. (BR); Piemonte, Prov. Alessandria:
Valenza, river Po north of the bridge, muddy, shady
track, 07.09.2014, F. Verloove 10993 (BR); Lombardia,
Prov. Pavia: Bereguardo, river Ticino close to the bridge,
sandy river bank, 11.09.2014, F. Verloove 11009 (BR);
Lombardia, Prov. Lodi: San Rocco al Porto, c. 500 m
west of bridge SS 9, exposed sandy river bank,
13.09.2014, F. Verloove 11025 (BR); Emilia-Romagna,
Prov. Reggio Emilia: Guastalla, canale Crostolo close to
river Po, muddy bank, 15.09.2014, F. Verloove 11034
(BR).
Section Umbellati
Cyperus aggregatus (Willd.) Endl., Cat. Horti Vindob.
1: 93. 1842 (Figure 12)
(≡)Mariscus aggregatus Willd., Enum. Pl. 1: 70. 1809
Native: Central and South America, Mexico, southern
USA. Naturalized in Australia and Europe.
Cyperus aggregatus was relatively recently reported
from Labenne in the Landes department in France where
it was discovered on the verge of a pine wood and in a
sandy roadside in 1993 (Chaffin and Vivant 1996). How-
ever, a herbarium revision demonstrated its presence in
this area since at least the 1950s (see “Specimens exam-
ined”). At present this species is locally firmly estab-
lished and expanding in southwestern France, especially
in the surroundings of Labenne. In 2013 C. aggregatus
was also recorded in abundance in Ychoux, about
100 km to the north.
In Spain (province of Huelva), C. aggregatus has
been recorded on several occasions, the first time appar-
ently in 2004 (Castroviejo 2006; Verloove and Sánchez
Gullón 2010). Records are available from Huelva and
Mazagón but, at least for the time being, C. aggregatus
remains an ephemeral alien there (comm. E. Sánchez
Gullón, May 2013). In France C. aggregatus grows in
nitrophilous ruderal habitats, usually in sandy and sun-
exposed, dry localities (mainly roadsides).
Selection of specimens examined
FRANCE –Dep. Landes: Dax, berges de l’Adour,
07.1953, L. Delvosalle s.n. (BR); Dep. Landes: Dax, rive
de l’Adour, 08.1973, A. Sotiaux s.n. (BR, LG, TAIF,
WAG); Dep. Landes: Labenne, aire de Labenne (motor-
way A63), sandy road verge, 08.09.2006, F. Verloove
6476 (BR); Dep. Landes: Ychoux, close to railway station,
roadside, very common, in dense nearly monospecific
stands, 16.08.2013, F. Verloove 10432 (BR, P). SPAIN –
Prov. Huelva: Huelva, parking Carrefour, ruderal aceras,
09.06.2008, E. Sánchez Gullón 127 (BR, MGC); Prov.
Huelva: Huelva, Punta del Sebo, ruderal cuneta carretera,
16.09.2011, E. Sánchez Gullón s.n. (BR).
Cyperus croceus Vahl, Enum. Pl. 2: 357. 1805
(Figure 13)
Native: southeastern USA, Mexico, West Indies, Central
and South America (Venezuela).
Cyperus croceus is known since 2004 from irrigated
lawns in two different localities in Mazagón (province of
Huelva, Spain) where it was probably unintentionally
introduced as a contaminant in grass seed (Verloove and
Sánchez Gullón 2010). It grows among other turf weeds
[Axonopus fissifolius (Raddi) Kuhlm., C. retrorsus,
210 F. Verloove

Figure 13. Cyperus croceus /Cyperus retrorsus:C. retrorsus: Habitus (1a, 1b), spike (2a), spikelet (3), glume (4), nutlet + style (5)
/C. croceus: Spike (2b).
Webbia: Journal of Plant Taxonomy and Geography 211

C. sesquiflorus and Digitaria violascens Link] and seems
firmly established although it does not seem to spread
beyond the initial area of introduction. In Mazagón
C. croceus and C. retrorsus grow in close proximity and
were both initially erroneously ascribed to C. cyperoides
(L.) Kuntze (Castroviejo 2006,2007a).
Cyperus croceus was known for quite a long time as
C. globulosus Aubl. However, this name is a synonym
of the very different C. luzulae (Carter and Kral 1990).
Selection of specimens examined
SPAIN –Prov. Huelva: Mazagón, Parador Nacional de
Turismo (UTM 29SPB9809), 09.2004, E. Sánchez Gul-
lón 16 (BR); Prov. Huelva: Mazagón, Faro, 24.07.2008,
E. Sánchez Gullón 128 (BR); Prov. Huelva: Mazagón,
Parador Nacional, lawn weed, 08.07.2009, E. Sánchez
Gullón 217 (BR); Prov. Huelva: Mazagón, Parador Nac-
ional, arvense en césped, 15.09.2009, E. Sánchez Gullón
232 (BR).
Cyperus retrorsus Chapm., Bot. Gaz. 3(3): 17. 1878
(Figure 13)
(≡)Mariscus retrorsus (Chapm.) C.B. Clarke, Bull.
Misc. Inform., Addit. Ser. 8: 15. 1908
(=) Cyperus ovularis auct. eur. non (Michx.) Torr., Ann.
Lyceum Nat. Hist. New York 3: 278. 1836
(=) Cyperus echinatus auct. eur. non (L.) Alph.Wood,
Class-book Bot. (ed. 1861): 734. 1861
Native: southeastern USA, Mexico. Naturalized in
Europe.
In the Azores C. retrorsus was discovered in 1929 in
two localities (Lajes and São Mateus) on the southern
slopes of the Pico Island (Tutin and Warburg 1932;
Palhinha 1966; sub C. ovularis). At present the species
is also known from Faial Island (Schäfer 2003). In Pico
it is least rare. In both islands it is confined to the low-
lands in the south, from close to sea level up to c. 300 m
(Schäfer 2003). In the Azores C. retrorsus was initially
found growing on lava on top of dry cliffs (Tutin and
Warburg 1932). At present it exclusively occupies dis-
turbed habitats (roadsides, waste places and coastal
rocks; Schäfer 2003).
In Spain, C. retrorsus was discovered in an irrigated
lawn at the lighthouse in Mazagón (province of Huelva)
in 2004. Soon afterwards it was found in identical cir-
cumstances at the Parador Nacional, also in Mazagón. It
was initially erroneously ascribed to C. cyperoides
(Castroviejo 2006,2007a) and subsequently corrected to
C. retrorsus (Verloove and Sánchez Gullón 2010). At
present it is fully established in both localities but shows
no tendencies (yet) to spread. In Mazagón it grows along
with the very similar C. croceus. Both these species are
also frequent in poorly kept lawns in coastal areas in the
southeastern United States (Carter and Bryson 1991;
Bryson et al. 1998).
There has been a lot of confusion with C. retrorsus in
the Atlantic Islands, both as regards its nomenclature and
distribution. The plants from the Azores were initially
ascribed to C. ovularis (Tutin and Warburg 1932) and this
binomial [or its synonym in the genus Mariscus,M. ovu-
laris (Michx.) Vahl] was used by all subsequent authors
(e.g. Palhinha 1966; Eriksson et al. 1979; DeFilipps
1980; Schäfer 2003). Carter and Kral (1990) advocated
that C. ovularis is in fact conspecific with C. echinatus,
the latter binomial having priority. Soon afterwards, in
the fourth edition of the Flora of Macaronesia, Hansen
and Sunding (1993) included for the Azores (Pico) both
C. retrorsus and “Mariscus ovularis”(= C. echinatus).
These data were also incorporated in the recent Euro+-
Med Plantbase (Jiménez-Mejías and Luceño 2011). How-
ever, after having examined several Azorean specimens
(see below) it is evident that only one of them occurs in
the Azores, C. retrorsus. This species has smaller spik-
elets, glumes, anthers and nutlets than C. echinatus
(Tucker et al. 2002). Moreover, its spikes are longer than
wide (not spherical) and much less tight.
In the Euro+Med Plantbase (Jiménez-Mejías and
Luceño 2011)“C. echinatus”is also acknowledged from
Madeira (Porto Santo), based on Hansen and Sunding
(1993). This appears to be erroneous as well.
Cyperus cyperoides, naturally occurring from Africa
to the Pacific, is an exceedingly variable species and
some forms or depauperate plants are much reminiscent
of C. retrorsus (both belong to section Umbellati in
Kükenthal 1935–1936). In Spain both have been inter-
mixed (see above). Cyperus cyperoides usually has
longer (20–40 mm) cylindrical spikes that are more than
twice as long as wide, longer glumes (up to 4 mm long)
and much narrower, longer nutlets (linear-oblong).
Future occurrences of C. cyperoides in the warmer parts
of the Flora Europaea area (Mediterranean area, Atlantic
Islands) are not unlikely, especially because it is also
increasingly cultivated as an ornamental, often under its
horticultural name “Cyperus zumula”(see also Simpson
and Inglis 2001).
Selection of specimens examined
AZORES –Pico: São Mattheus, cliff top, on lava,
07.1929, E.F. Warburg & T.G. Tutin s.n. (CGE; sub C.
ovularis); Pico: Lages [Lajes], cliff top, 07.1929, E.F.
Warburg & T.G. Tutin s.n. (CGE; sub C. ovularis); Pico:
Lajes, in salt marsh, 04.07.1987, A. Hansen 359 (C; sub
C. ovularis). - SPAIN: Prov. Huelva: Mazagón, close to
the lighthouse, lawn, very abundant, with Kyllinga
odorata, 10.10.2007, F. Verloove 6969 (BR); Prov.
Huelva: Mazagón, Parque del Faro, lawn weed, 06.2009,
E. Sánchez Gullón 177 (BR, LG); Prov. Huelva:
Mazagón, Parque del Faro, arvense en césped,
15.09.2009, E. Sánchez Gullón 233 (BR); Prov. Huelva:
212 F. Verloove

Figure 14. Cyperus brevifolius /Cyperus brevifolioides:C. brevifolius: Habitus (1), spikelet (2a), nutlet + style (3) /
C. brevifolioides: Spikelet (2b).
Webbia: Journal of Plant Taxonomy and Geography 213

Mazagón (UTM 29SPB9312), en césped ornamental,
20.09.2010, E. Sánchez Gullón 271 (BR).
[Kyllinga]
Section Kyllinga
Cyperus brevifolioides Thieret & Delahouss., Sida 3:
129. 1967 (Figure 14)
(=) Kyllinga gracillima Miq., Ann. Mus. Bot. Lugduno-
Batavi 2: 142. 1865
(=) K. brevifolia var. leiolepis (Franchet & Savatier) H.
Hara, J. Jap. Bot. 14: 339. 1938
Native: temperate East Asia. Naturalized in southeastern
USA and Europe.
Cyperus brevifolioides was discovered in Italy in Vernate
(province of Milano) in 2002. Soon afterwards it was
recorded in an additional locality in the same area
between Milano and Pavia. It is now considered a natu-
ralized and invasive alien (Galasso et al. 2006). Previous
inaccurate records of this species between Milano and
Pavia (in the 1950s; see below) may indicate an earlier
and wider naturalization (Galasso et al. 2006). This spe-
cies was not yet included in the Euro+Med Plantbase
(Jiménez-Mejías and Luceño 2011).
In Italy C. brevifolioides grows alongside small irri-
gation canals and ditches in the rice-growing area.
The taxonomic position of C. brevifolioides is
debated. It was accepted as a distinct species in the Flora
of North America (Tucker 2002 as K. gracillima), while
in the Flora of China (Dai et al. 2010b) it was only
given varietal rank. An upcoming monographic study of
(sub-)genus Kyllinga (Huygh, in prep.) applies a broad
species concept without recognition of any infraspecific
taxa. Cyperus brevifolioides is distinguished by having
glumes with smooth keels, two or three anthers and
longer styles.
Selection of specimens examined
ITALY –Lombardia, Prov. Milano: (…) di Milano,
09.1950, (…) in herb. C. Stucchi s.n. (MSNM 255);
Lombardia, Prov. Milano: Milano, strada pavese,
10.1954, Viola in herb. C. Stucchi s.n. (MSNM 256);
Lombardia, Prov. Milano: Vernate, località Pasturago,
risaie poco a nord dell’Oasi di Pasturago, lungo un
canale, c.100 m s.l.m., sponda di un canale, 25.10.2005,
G. Galasso s.n. (MSNM 40188, 40189, 40190, 40191,
40192, 40193, 40194); Lombardia, Prov. Milano:
Lacchiarella (Mettone), canale presso il Cavo Socio, rag-
giungibile dallo sterrato che si stacca dalla strada tra
Mettone e Casirate, 93 m s.l.m., sponda di un canale,
25.10.2005, G. Galasso s.n. (BR, MSNM 40195, 40196,
40197, 40198, 40199, 40200, 40201); Lombardia, Prov.
Milano: Pasturago, north of Oasi di Pasturago, ditch,
11.09.2014, F. Verloove 11006 (BR).
Cyperus brevifolius (Rottb.) Endl. ex Hassk., Cat. Hort.
Bot. Bogor. 24. 1844 (Figure 14)
(≡)Kyllinga brevifolia Rottb., Descr. Icon. Rar. Pl. 13,
pl. 4, f. 3. 1773
(=) Kyllinga monocephala auct. lusit., non Rottb., Descr.
Icon. Rar. Pl. 13, pl. 4, f. 4. 1773
Native: pantropics but probably not originally native in
Africa (Huygh, in prep.). Naturalized in Europe.
In the Azores C. brevifolius was already documented by
Cederkreutz (1941). Palhinha (1966) reported it from the
islands Flores, São Miguel and Terceira and now it is
also known from Corvo, Faial, Graciosa, Pico, Santa
Maria and São Jorge (Jiménez-Mejías and Luceño 2011).
In Madeira it was first reported by Hansen (1969)
from Ribeira Brava. It later turned out to have been
known before: it was already recorded in abundance in
1963 and 1965 in Caniçal and Estádio dos Barreiros
(Vieira 2002). Press and Short (1994) further cite São
Jorge in the north and Água de Pena in the east. It is
also known from Porto Santo (Vieira 2002). It has
become an aggressive weed of agricultural fields, espe-
cially (but not exclusively) in southern coastal areas.
In the Canary Islands C. brevifolius is a recent intro-
duction. It was discovered as a weed on a golf course
(“green”) in Buenavista del Norte in Tenerife in 2009
(Siverio Núñez et al. 2013) and in 2014 also near La
Orotava (see “Specimens examined”). In Gran Canaria it
was found for the first time in 2011 in irrigated lawns in
Arguineguín (Verloove 2012).
In Portugal C. brevifolius is known since the 1880s
from the estuary of the river Minho in Caminha (Daveau
1891) and at present it is well-established in several dif-
ferent parts of the country (Douro Litoral, Estremadura,
Minho, Trás-os-Montes; see Rainha 1959; Almeida
1999; Castroviejo 2007b).
In Spain C. brevifolius was first discovered in Los
Barrios (province of Cádiz) in damp meadows in 2001
(Luceño and Marín 2002). An earlier Spanish claim
(Sánchez Gullón 1999) turned out to be referable to
C. sesquiflorus (Sánchez Gullón and Verloove 2013). In
2012 it was also found in irrigated lawns in Palos de la
Frontera (Monasterio de La Rábida, province of Huelva)
(Sánchez Gullón and Verloove 2013).
In the study area C. brevifolius is found as a weed in
a wide range of (mostly) disturbed, at least temporarily
damp habitats: irrigated lawns, agricultural fields, road-
sides, etc.
Selection of specimens examined
AZORES –Terceira: Ribeira dos Moinhos, 05.1937,
R.T. Palhinha & L.G. Sobrinho s.n. (C); Santa Maria
(W): in the area between the airport and the
214 F. Verloove

Figure 15. Cyperus sesquiflorus: Habitus (1a, 1b), spike (2), nutlet + style (3). spikelet (4).
Webbia: Journal of Plant Taxonomy and Geography 215

Figure 16. C. odoratus: Habitus (1a, 1b), Spike (2), Spikelet (3), Nutlet + style (4), Glume + nutlet + rachis (5).
216 F. Verloove

“Hotel-City”, ditch, 28.08.1970, A. Hansen s.n. (C); São
Miguel: between Feteiras and Sete Cidades, roadside,
31.08.1970, A. Hansen s.n. (C); São Miguel: c.6km
south of Furnas, roadside on wet ground, 01.09.1970, A.
Hansen s.n. (C); São Miguel: Furnas, roadside (wet) near
the “boilers”, 29.06.1971, A. Hansen 44 (C); São
Miguel: Sete Cidades, at the shore of the Lagoa Azul,
30.06.1971, A. Hansen 122 (C); São Miguel: Ponta
Delgado, among pavement in a street, 30.06.1972, A.
Hansen 111 (C); Faial: above Flamengos, on roadside,
05.07.1973, A. Hansen 211 (C); Faial: near Horta, in bar-
ranco de Flamingos, 08.07.1973, A. Hansen 246 (C);
Santa Maria: near the airport, swamp, 11.06.1974, A.
Hansen 44 (C); São Miguel: at lake Furnas, 27.06.1979,
A. Hansen 31 (C); São Miguel: Sete Cidades, in a
swamp, 27.06.1987, A. Hansen 1224 (C); Pico: Cais do
Pico, in fields above the town, 05.07.1987, A. Hansen
1415 (C); São Miguel: Sete Cidades, 450 m, talus,
16.07.2000, J. Bouharmont 30481 (BR). CANARY
ISLANDS –Gran Canaria: Arguineguín towards Patala-
vaca, barranco, lawn weed, very common, 10.11.2011, F.
Verloove 9169 (BR); Gran Canaria: Arguineguín towards
Patalavaca, GC500 at Sun Wing Resort, lawn weed,
19.11.2011, F. Verloove 9175 (BR); Gran Canaria:
Arguineguín, centro ciudad, irrigated lawn, 18.09.2013,
F. Verloove 10586 (BR, LPA); Tenerife: La Orotava, TF
217 (En la Urbanizacion), irrigated lawn, very common
but only locally, 27.06.2014, F. Verloove 10,852 (BR,
ORT). MADEIRA –Machico, bord de fossé, zone de
suintements aux abords des cultures, au-dessus du
complexe touristique Matur, 20.07.1975, J. Duvigneaud
75M537 (BR, LG); Caniçal, suintements en bordure de
la mer, un peu à l’est du village, 21.07.1975, J. Duvig-
neaud 75M550 (BR, LG); R101 west of São Vicente,
30.04.2008, R. Barendse s.n. (BR). - SPAIN: Prov.
Huelva: La Rábida, Palos de la Frontera (UTM
29SPB8420), césped ornamental acompañado por
Paspalum dilatatum, Axonopus affinis, etc., 15.09.2012,
E. Sánchez Gullón 383 (BR).
Cyperus sesquiflorus (Torr.) Mattf. & Kük., Pflanzenr.
IV. 20 (Heft 101): 591. 1936
subsp. cylindricus (Nees) T. Koyama, Bot. Mag. 83:
187. 1970 (Figure 15)
(≡)Kyllinga cylindrica Nees, Contr. Bot. India 91. 1834
(≡)Kyllinga odorata Vahl, Enum. Pl. 2: 382. 1805
subsp. cylindrica (Nees) T. Koyama, Gard. Bull.
Singapore 30: 161. 1977
(≡)Kyllinga odorata var. cylindrica (Nees) Kük., J.
Straits Branch Roy. Asiat. Soc. 75: 80. 1917
(≡)Kyllinga sesquiflora Torr., Ann. Lyceum Nat. Hist.
New York 3: 287. 1836 subsp. cylindrica (Nees) T.
Koyama, Fl. Taiwan 5: 288. 1978
(≡)Cyperus sesquiflorus var. cylindricus (Nees) Kük.,
Pflanzenr. IV.20 (Heft 101): 593. 1936
(=) Cyperus kernianus Ohwi & T. Koyama, J. Jap. Bot.
30: 126. 1955
(=) Cyperus sesquiflorus var. subtriceps (Nees) T.
Koyama, Quart. J. Taiwan Mus. 14: 191. 1961.
Native: pantropics. Naturalized in Europe.
Cyperus sesquiflorus was first discovered in Spain in
1998 in irrigated lawns at the Parador Nacional in
Mazagón (province of Huelva) (Sánchez Gullón 1999
sub K. brevifolia). Soon afterwards it was recorded, in
identical circumstances, near the lighthouse in Mazagón
(Castroviejo 2006). At present it is well-established in
these localities but it shows no tendency to spread (yet).
Plants naturalized in Europe (Spain) are characterized
by small spikelets (c. 2.5 mm long), narrow leaves (less
than 3 mm wide) and glumes with a smooth keel (not
spinulose). Such plants belong to subsp. cylindricus.
Some, mainly Asian authors, warrant specific status to
this taxon (e.g. Koyama 1955; Dai et al. 2010b), others
only varietal status (e.g. Kükenthal 1935–1936; Kern
1974; Hoenselaar et al. 2010). No infraspecific variabil-
ity is upheld in South Africa (Getliffe 1983) and an
upcoming monographic study of (sub-) genus Kyllinga
(Huygh, in prep.) also applies a broad species concept
without recognition of any infraspecific taxa.
Selection of specimens examined
SPAIN –Prov. Huelva: Mazagón, Parador Nacional,
lawn, 10.10.2007, F. Verloove 6939 (BR); Prov. Huelva:
Mazagón, close to the lighthouse, lawn, abundant,
10.10.2007, F. Verloove 6940 (BR); Prov. Huelva:
Mazagón, Parque del Faro, lawn weed, 06.11.2008, E.
Sánchez Gullón s.n. (BR); Prov. Huelva: Mazagón, Para-
dor Nacional, lawn weed, 05.07.2009, E. Sánchez Gullón
220 (BR).
[Torulinium]
Section Diclidium
Cyperus odoratus L., Sp. Pl.: 46. 1753 (Figure 16)
(≡)Torulinium odoratum (L.) S·S. Hooper, Kew Bull.
26: 579. 1972
(=) Cyperus ferax Rich., Actes Soc. Hist. Nat. Paris 1:
106. 1792
(=) Torulinium ferax (Rich.) Urban, Symb. Antill. 2(1):
165. 1900
Native: pantropics. Naturalized in Europe.
The naturalization history of C. odoratus in the study
area is very complicated as it has widely been confused
with C. strigosus and other species. The first confirmed
European occurrences (at least of stable populations)
apparently are from Italy where C. odoratus now is
Webbia: Journal of Plant Taxonomy and Geography 217

widely naturalized in the valley of the river Po as seen
in September 2014. It is here reported for the first time
but evidence from herbarium specimens demonstrates its
presence there as early as 1958 in Casalmaggiore (see
“Specimens examined”). A more thorough herbarium
revision will probably shed new light on the species’
invasion history in Italy (Verloove, in prep.).
In Romania it was first discovered in Sulina (Roman
1992), soon afterwards also in Letea and Sfântu
Gheorghe (Ciocârlan 1992,1994), all located in the
Danube Delta Biosphere Reserve (Negrean 2011; Sîrbu
and Oprea 2011). In the same area, Anastasiu (2011)
recently added several new localities. Cyperus odoratus
obviously is in expansion but not (yet) considered an
invasive species in Romania (Anastasiu et al. 2008).
The first Bulgarian records of C. odoratus date back
to 2000 and 2001 when it was recorded in the Danube
plain in the surroundings of Belene and Novgrad respec-
tively (Tzonev et al. 2003). These and further Bulgarian
records (see “Specimens examined”) were initially erro-
neously ascribed to C. strigosus and here correctly
assigned. It is now fully established along the river
Danube and several of its tributaries.
Similarly, C. odoratus was recently reported for the
first time from Serbia and Montenegro, again under the
erroneous binomial C. strigosus (Stevanovićet al. 2004,
2005). It was first seen in Mali Lap marsh near the village
of Dubovac (km 1086) and subsequently, during the
investigation of macrophytic vegetation in the Danubian
river plain, in numerous additional localities, mainly in
small populations. Cyperus odoratus now is a widespread
alien along the Danube river from Kovin (km 1110) to
Mihajlovac very near to Djerdap hydropower dam II (km
872). It is obviously a fast-spreading species although it
could not be considered as invasive as, for instance, Pasp-
alum distichum L. (comm. V. StevanovićApril 2013).
A third area of naturalization in Europe is located in
northeastern Spain. Along the river Segre C. odoratus was
already very common in 2006, at least between Balaguer
and its confluence with the river Ebro near Mequinenza, a
stretch of c. 75 km long (comm. J. Conesa, March 2006).
It was probably introduced long before but remained
undetected. Castroviejo (2005,2006) was the first to refer
to these plants. They were initially erroneously ascribed to
C. strigosus but soon afterwards correctly assigned to
C. odoratus. A recent claim of “C. digitatus subsp. au-
ricomus”(Cardero Aguilera et al. 2004) from Tortosa
(province of Taragona) also belongs here (see “Specimens
examined”). At present C. odoratus seems mostly con-
fined to the basins of the rivers Ebre and Segre (and their
tributaries), although it has also been recorded as a weed
in orchards in the same area.
Claims from France (e.g. Castroviejo 2007a) are
probably the result of confusion with C. strigosus and
require confirmation. Recently, C. odoratus was also
reported for the first time from Israel (Danin and
Dufour-Dror 2008). A future, wider naturalization in
suitable habitats in the Mediterranean area is to be
expected.
In Europe C. odoratus has widely been confused
with C. strigosus although both are only superficially
similar. Apart from the very different mode of spikelet
disarticulation (respectively spikelets falling apart in
units containing a glume and part of the rachis that
tightly encloses the nutlet versus spikelets falling as a
whole) both differ, among others, in spikelet shape (ter-
ete versus flattened), colour (golden versus yellowish
green at flowering). Ramachandran and Soosairaj (2014)
also emphasized the close resemblance between these
two species. See also under C. strigosus. In Italy, C. odo-
ratus has also been confused with the rather different
C. glomeratus. The latter has more densely clustered
spikes, some of which are obviously longer than wide
(versus well-separated, spherical spikes), much narrower
glumes, etc.
Cyperus odoratus is an exceedingly variable species
and numerous segregates have been described. These
have been treated as distinct species, varieties or syn-
onyms. A morphometric analysis by Tucker (1984) dem-
onstrated that these segregates should be accepted as
mere synonyms of C. odoratus.
In the study area C. odoratus is mostly found by riv-
ers: estuaries, mud flats, gravelly or sandy exposed
banks, etc. However, it has also been recorded by tracks
in riparian woodland, in ditches and drainage channels
and even as a weed in orchards. It seems to spread in a
very fast way and locally behaves as an invasive envi-
ronmental weed.
Selection of specimens examined
BULGARIA –Prov. Pleven: Belene, Danube riverside,
6–7 km west of town, 03.10.2000, R. Tzonev s.n. (SO
101736, 101737; sub C. strigosus); Prov. Ruse: near
Novgrad, in the old estuary of Yantra river, 30.08.2001,
R. Tzonev s.n. (SO 101635; sub C. strigosus); Prov. Ple-
ven: Danube plain region, Pisarevo village, near to Iskar
river, 23.09.2006, R. Tzonev s.n. (SO 104291; sub
C. strigosus). ITALY –Lombardia, Prov. Cremona: Ca-
salmaggiore, rives du Pô, 30 m, 10.09.1958, R. Barbezat
s.n. (BR; sub C. glomeratus); Lombardia, Prov. Pavia:
Arena Po, 1 km E dal ponte sul Po (SP 199), destra id-
rografica del fiume Po, ZPS « Po da Albaredo Arnaboldi
ad Arena Po », sponda su suelo sabbioso-limoso, veget-
azione pioniera rada, 50 m, 19.09.2009, N. Ardenghi s.n.
(priv. herb. N. Ardenghi; sub C. strigosus); Emilia-
Romagna, Prov. Piacenza: Piacenza, river Po close to
bridge SS 9, muddy river bank, scattered individuals,
13.09.2014, F. Verloove 11011 (BR); Lombardia, Prov.
Lodi: San Rocco al Porto, river Po close to SS 9, muddy
river bank, 13.09.2014, F. Verloove 11014 (BR);
Emilia-Romagna, Prov. Piacenza: Castelvetro Piacentino,
località Maginot, river Po, muddy river bank,
14.09.2014, F. Verloove 11022 (BR); Lombardia, Prov.
218 F. Verloove

Cremona: Martignana di Po, river Po, damp track along-
side the river, 14.09.2014, F. Verloove 11027 (BR);
Lombardia, Prov. Cremona: Casalmaggiore, river Po
west of railway bridge, known at least since 1958 from
this locality, 14.09.2014, F. Verloove 11029 (BR); Emili-
a-Romagna, Prov. Reggio Emilia: Boretto, river Po east
of the bridge, muddy river bank, 15.09.2014, F. Verloove
11031 (BR); Emilia-Romagna, Prov. Reggio Emilia:
Guastalla, Isola Internati, river Po, muddy river bank,
15.09.2014, F. Verloove 11032 (BR); Emilia-Romagna,
Prov. Reggio Emilia: Guastalla, canale Crostolo close to
river Po, muddy bank, frequent, 15.09.2014, F. Verloove
11033 (BR); Emilia-Romagna, Prov. Reggio Emilia:
Guastalla, north of bridge SP 57bis, exposed bank of
sand pit, very common, 15.09.2014, F. Verloove 11035
(BR); Emilia-Romagna, Prov. Ferrara: Ferrara,
Pontelagoscuro, river Po, muddy river bank, 16.09.2014,
F. Verloove 11039 (BR); Lombardia, Prov. Mantova:
Quatrelle, river Po, sandy river bank, 16.09.2014, F. Ver-
loove 11042 (BR). ROMANIA –Sfântu Gheorghe, Delta
Dunării, marginea canalelor, 12.09.1991, V. Cioĉarlan
s.n. (BUAG, BR); Vatafu-Imputita (Delta Dunării, jud.
Tulcea), loc mlăştinos,10.09.2013, M. Doroftei s.n.
(IASI, BR). SERBIA –Banat: Dubovac, Mali Lap, na
obodu trske u dnu nasipa, 26.07.2003, V. Stevanovic
22336 (BEOU; sub C. strigosus); Banat: village Dubo-
vac, Mali Lap marsh, crevices of cement bricks on lake-
ward slope of dam, alt. c.70–80 m a.s.l., 08.09.2004, P.
Uotila 45577a (H 1,730,595; sub C. strigosus); Banat:
village Dubovac, Mali Lap marsh, wet, dense Glyceria
maxima etc. grass and sedge belt at foot of dam, alt.
c.70–80 m a.s.l., 08.09.2004, P. Uotila 45577b (H
1,730,586; sub C. strigosus). SPAIN –Prov. Tarragona:
Tortosa (Baix Ebre), Illa d’Audí (UTM 31TBF92), 12 m
s.m., Bidention, enmig de poblaments de Leersia,
16.09.2003, Hodgson, Palmer, Torres & Royo 5304
(BCN 21521; sub C. digitatus subsp. auricomus); Prov.
Lérida: Balaguer (N-Lérida), near old bridge over riu
Segre, muddy riverbank, locally common, 04.09.2005, F.
Verloove 6142 (BR, GENT, LG); Prov. Lérida: Balaguer,
riu Segre (close to new bridge), riverbank, common in
this area, 09.09.2007, F. Verloove 6905 (BR); Prov. Lér-
ida: Torres de Segre, north side of Pantana d’Utxesa
(Mas de Ribes), ditch along orchard, very locally (usu-
ally confined to riu Segre), 12.09.2007, F. Verloove 7025
(BR); Prov. Tarragona: Tortosa (Baix Ebre), pr. pont del
Mil·lenari (UTM 31TBF904202), 10 m, herbazales junto
al río Ebro, suelo arenoso, 08.09.2010, F. Royo, R.
Curto, M.J. Tohá & V.J. Arán 136–03-06a (MA
851240).
New combinations
Cyperus sect. Fasciculati (Boeckeler) F. Verloove, comb.
nov.
(≡)Cyperus [unranked] Fasciculati Boeckeler, Linnaea
36: 342. 1870.
Cyperus sect. Stoloniferi (Nakai) F. Verloove, comb.
nov.
(≡)Juncellus sect. Stoloniferi Nakai, Bot. Mag. (Tokyo)
26: 199. 1912.
(=) Cyperus sect. Serotini Kük., nom. illeg.
Cyperus sect. Umbellati (C.B. Clarke) F. Verloove,
comb. nov.
(≡)Mariscus sect. Umbellati C.B. Clarke, Fl. Brit. India
6: 620. 1893.
Cyperus alternifolius subsp. textilis (Thunb.) F. Verloove,
comb. nov.
(≡)Cyperus textilis Thunb., Prodr. Pl. Cap.: 18. 1794.
Acknowledgements
I am very thankful to the curators of the herbaria BCN, BEOU,
BR, C, CGE, GENT, H, LG, MA, MSNM, PAV and SO for
offering relevant collections (including scans) for examination.
Numerous botanists kindly provided various information (speci-
mens, localities, literature, etc.): A. Alessandrini and colleagues
(Italy), J. Amigo (Spain), J.-C. Aniotsbéhère (France), N.
Ardenghi (Italy), J. Conesa (Spain), J. Dexheimer (France), M.
Doroftei (Romania), G. Faggi (Italy), G. Galasso (Italy), P.
Goetghebeur (Belgium), C. Grieco (Italy), Q. Groom
(Belgium), D. Iamonico (Italy), H. Michaud (France), R. Otto
(Germany), D. Saiani and colleagues (Italy), E. Sánchez Gullón
(Spain), H. Schäfer (Germany), M. Sequeira (Madeira), C.
Sîrbu (Romania), A. Soldano (Italy), V. Stevanović(Serbia),
M. Tabacchi (Italy), J.-M. Tison (France), N. Torres (Spain), R.
Tzonev (Bulgaria) and P. Uotila (Finland). M. Reynders is
thanked for sharing his ideas and knowledge and for interesting
discussions. Sven Bellanger and Liliane Tytens are acknowl-
edged for preparing the line drawings. Finally, field work in
Italy in September 2014 was granted by COST (Cooperation in
Science and Technology), Action TD1209.
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