Bot. Rev.

(2012) 78:184–204
DOI 10.1007/s12229-012-9098-z

Structure of the Cyperaceae Inflorescence

Andrea Reutemann1,2 & Leandro Lucero1,2 &

Nicolás Guarise1 & Abelardo C. Vegetti1,2,3
1
Facultad de Ciencias Agrarias, Universidad Nacional del Litoral, Esperanza, Pcia. Santa Fe, Argentina
2
Instituto de Agrobiotecnología del Litoral, Facultad de Ciencias Agrarias, Universidad Nacional del
Litoral, Kreder 2805, S3080HOF Esperanza, Provincia de Santa Fe, Argentina
3
Author for Correspondence; e-mail: avegetti@fca.unl.edu.ar
Published online: 5 April 2012
# The New York Botanical Garden 2012

Abstract This work presents the basics for interpreting the adult inflorescence
structure in Cyperaceae. It provides an analysis of the variations of the synflorescence
and inflorescence structure in the family. Three types of synflorescence may be
recognized in this family: a synflorescence with a foliate stem, a terminal inflores-
cence and a variable number of lateral inflorescences; a synflorescence with a foliate
stem and only the terminal inflorescence; and a synflorescence with a scape and a
terminal inflorescence. Variations in the structure and form of the inflorescences are
related to variations in inflorescence branching, inflorescence homogenization de-
gree, presence or absence of the distal part of the inflorescence, phyllotaxis, inflo-
rescence position, types of bracts and leaves subtending branches, elongation of
inflorescence internodes and spikelet structure. These variations are correlated with
some of the developmental processes that give origin to the inflorescence.

Keywords Cyperaceae . Synflorescence . Inflorescence . Homogenization process .

Truncation proccess

Introduction

Cyperaceae, the third largest monocotyledonous family, comprise ca. 5.500 species in
ca. 109 genera (Muasya et al., 2009) and constitute a specialized group of plants,
particularly in relation to their reproductive structure (Kukkonen, 1994). In studying
the Cyperaceae family, the characters of the inflorescence have played an important
role in the diversification of many taxa, and have had significant phylogenetic value
(Tucker & Grimes, 1999). Cyperaceae are difficult to classify due to the complex
structure of their inflorescences, which leads to different interpretations and to
establishing uncertain hypotheses of homology (Kukkonen, 1984, 1986, 1994;
Simpson, 1995; Muasya et al., 1998; Vegetti, 2003). The establishment of hypotheses
of structural homologies among different types of inflorescences is one of the main
concerns of cyperologists (Raynal, 1971).
In the formal taxonomy of Cyperaceae, the misapplication of terms with regard to
descriptions of the inflorescence morphology has led to wrong examples in the
Inflorescence of Cyperaceae 185

morphological sense, since uniformity or stability of terminology have been lacking

(Kukkonen, 1994; Browning & Gordon-Gray, 1999). Within this context, homolo-
gous structures are named differently, and different structures are named alike
(Simpson, 1992; Alves, 2000). The main problems in the interpretation of the
inflorescence structure result from studies which did not consider the entire inflores-
cence (Vegetti, 2003); instead, attention was restricted to the position and arrange-
ment of flowers in the final units of the often copiously branched inflorescence
(Kukkonen, 1984).
In order to understand and compare inflorescences within this family, it is neces-
sary to establish a terminology for the different components of the inflorescence; such
terminology should refer to homologous structures and make it possible to determine
variation paths from basic organizational plans (Mora-Osejo, 1987). Several broad
approaches have been presented in the family (Haines, 1966; Guaglianone, 1970;
Raynal, 1971; Meert & Goetghebeur, 1979; Haines & Lye, 1983; Tucker, 1987, 1994;
Goetghebeur, 1998; Vegetti, 2003), however, a global view is lacking.
In view of this, this work aims at: a) stating the basics for interpreting inflores-
cence structures in Cyperaceae and their variations; and b) correlating such informa-
tion with the various developmental processes that give origin to those structures. It
must be noted that this work does not intend to comprise all of the variations
occurring in the inflorescence structure in the family, but it includes some of the
most relevant types.

The Structure of the Plant and its Floriferous Shoots

In Cyperaceae, as in many grass plants, the plant is composed of shoots of consec-

utive branching order. One of the shoots is the main axis of the plant and the others
are axillary shoots (innovations 0 basal branches) generated in the basal zone of short
internodes. Each of these shoots should be considered, in a morphological sense, as a
synflorescence (Troll, 1964; Rua, 1999). The synflorescence is the floriferous shoot
produced from an apical bud of the embryonic axis or from an innovation bud during
the growth period (Troll & Weberling, 1989; Rua, 1999).
The structure and general shape of the synflorescence result from the activity of its
meristems, which determines the number, size and relative disposition of its branches
(Guarise & Vegetti, 2007).
The axillary basal shoots may develop and flower within the same vegetative period
as their parental axis, or their branch development may be substantially delayed relative
to the development of the parental axis (Mora-Osejo, 1960). These are silleptic
innovation shoots or cataleptic innovation shoots, respectively (Rua, 1999; Vegetti,
2003). In most Cyperaceae, basal axillary meristems grow out to produce a highly
tillered plant. As it occurs in other monocotyledonous plants, these buds are still under
hormonal and environmental control (Shimamoto & Kyozuka, 2002; McSteen, 2009).
During the vegetative phase, the apical meristems of the main axis and the axillary
basal shoots generate leaves and shoots and, following floral induction, the terminal
inflorescence (tIn). This inflorescence may be in the terminal portion of a long
internode (scape) (Fig. 1C), or a foliate stem of somewhat developed internodes
(Fig. 1B, C). The axillary buds in the foliate stem may develop (Fig. 1A) or not
186 A. Reutemann et al.

Fig. 1 Structure of synflorescence and inflorescence in Cyperaceae: A, Synflorescence with foliate scape
and terminal (grey block) and lateral (black block) inflorescence; B, Synflorescence with foliate scape and
terminal inflorescence; C, Synflorescence with aphyllous scape and terminal inflorescence; D, Structure of
the inflorescence. References: b, bract; cof coflorescence; EZ, enrichment zone; HZ, inhibition zone; INF
inflorescence; IS, innovation shoot; IZ, innovation zone; lbr, long branch; lbrZ, zone of long branches; lIn,
lateral inflorescence; LIZ, long internode zone; pr, prophyll; shbr, short branch; shbrZ, zone of short
branches; SIZ, short internode zone; tIn, terminal inflorescence; ts, terminal spikelet

(Fig. 1B); thus, in some species, the tIn appears alone in the distal portion of such
foliate stem (Fig. 1C), while in other species the axillary buds develop floriferous
shoots (enrichment shoots, Fig. 1A). Each of these enrichment shoots ends in an
inflorescence (lateral inflorescence, lIn). The subtending leaves where the enrichment
axes originate from bear developed blades and sheaths. The subtendig leaves (bracts)
where the inflorescence branches originate from are modified leaves without sheaths.
Based on the aforementioned, the following synflorescence types may be recog-
nized in the family: (1) a synflorescence with a foliate stem and a tIn and a variable
Inflorescence of Cyperaceae 187

number of lIns (Fig. 1A), as in some species of Carex L., Fuirena Rottb., Rhyncho-
spora Vahl, Scleria P.J. Bergius; (2) a synflorescence with only a tIn and a foliate
stem, as in some species of Hypolytrum Pers. (Fig. 1B); (3) a synflorescence with a
scape and a tIn, like Isolepis R. Br. and Cyperus L., Bulbostylis Kunth, Fimbristylis
Vahl, Eleocharis R. Br., Scirpus L. (Fig. 1C).
Synflorescences with a foliate stem and terminal and lateral inflorescences
(Fig. 1A) are a primitive condition in the family (Guarise & Vegetti, 2007). Accord-
ing to these authors, the inhibition of the development of enrichment shoots from the
axils of the upper leaves makes it possible to obtain a synflorescence with only a tIn
and a foliate stem (Fig. 1B); while the shortening of the internode zone with the
elongation of the internode above the last vegetative leaf leads to the formation of
synflorescences with a scape and a tIn (Fig. 1C).
The change from a synflorescence with a foliate stem and tIn and lIns (Fig. 1A) to
a synflorescence with a scape and only a tIn (Fig. 1C) takes place in the Arthrosty-
lideae/Abildgaardieae clade (Guarise & Vegetti, 2007). According to these authors,
reduction trends, starting from one structure like the plesiomorphic synflorescence
proposed, were used to explain the variation observed in the inflorescences of
Hypolitrum (Alves et al., 2000), Rhynchospora section Dichronema (Michx.) Griseb.
(Thomas, 1984), Carex (Reznicek, 1990) and Kobresia Willd. (Zhang, 2001).
The presence of flowering branches in the axils of bracts with developed blade and
sheath along an axis brings about a discussion on the limits and structure of the
inflorescence in Cyperaceae. Several authors have considered the synflorescence with
the elongated and foliaceous main axis, with or without branches emerging from the
axil of leaves with developed blade and sheath, as a whole inflorescence termed
paniculodium (Mora-Osejo, 1960; Raynal, 1971; Haines & Lye, 1983; Bruhl, 1995;
Goetghebeur, 1998). However, other authors have considered the distal group of
branches as the ultimate inflorescence (Thomas, 1992; Strong, 2001; Alves, 2003) or
the terminal flowering unit (Reznicek, 1990); while the lateral branches produced at
the axil of normal leaves with developed blade and sheath have been named lateral or
partial inflorescence (Guaglianone, 1996; Alves, 2003) or lateral flowering unit
(Reznicek, 1990).
There is some evidence in favor of considering the existence of terminal and lateral
inflorescences in Cyperaceae. According to Mora-Osejo (1987), the enrichment axes
(i.e. the flowering branches arising from the axils of bracts with a well-developed
sheath) are differentiated from the inflorescence branch (i.e. the flowering branching
which emerges from buds originating from the axils of bracts without a developed
sheath) by their different development sequences. In this context, Sell (1980) sug-
gested that the enrichment axes flower basipetally while the inflorescence branches
develop acropetally. This sequence was observed in the reproductive meristem of
Carex disticha Huds. by Mora-Osejo (1987).

Inflorescence Structure in Cyperaceae

As a general model, during inflorescence development, the apical inflorescence

meristem (IM) generates bracts and their axillary meristems, called branch meristems
(BM), which are generally indeterminate and produce the inflorescence branches.
188 A. Reutemann et al.

After several branches are produced, the IM switches and generates axillary meris-
tems that produce one spikelet (spikelet meristem, SM), which is determinate. Next,
the IM may generate or not a varying number of bracts whose axillary meristems do
not develop and, finally, it generates fertile leaves (glumes) with floral meristems
(FM) in their axils, which make up the terminal spikelet.
In a few Cyperaceae, the IM does not produce branches, which means the
inflorescence has a single terminal spikelet (unispicate inflorescence) (Fig. 2E, G,
Goetghebeur, 1998; Vegetti, 2003). However, in most Cyperaceae, the IM produces
lateral meristems that may behave either wholly as SMs and, consequently, do not
produce other branches and end inmediately in a spikelet (Fig. 2D, e.g. species of
Ascolepis Nees ex Steud., Kyllinga Rottb., Lipocarpha R. Br.), or partially as BMs
and partially as SMs, and then the former go on to branch again (Fig. 2A–C, F). As a
consequence, based on the behavior of the meristems, the inflorescence may com-
prise one or several spikelets arranged in a branching system with varying degrees of
complexity (Fig. 2).
In unispicate inflorescences (Fig. 2E, G), the IM does not generate either a BM or
an SM, and if it does, such axillary meristems do not develop—only the FMs show.
Branching in spikes of spikelets (Fig. 2D) is controlled by two types of axillary
meristems: the SM and the FM. Branching in an inflorescence with a branched axis
(Fig. 2A–C, F) is controlled by three types of axillary meristems: BM, SM and FM.
The different inflorescence phenotypes are the result of different developmental
processes (Albert et al., 1998; Doust & Kellogg, 2002). These variations in the
inflorescence structure within the family are determined by variations in: a) inflores-
cence branching, b) inflorescence homogenization degree, c) presence or absence of
the distal part of the inflorescence, d) phyllotaxis, e) inflorescence position, f) types of
bracts and leaves subtending branches, g) elongation of inflorescence internodes, and h)
spikelet structure. All of these characters are controlled by the activiy of a group of
regulatory genes that respond to internal or external signals (Kellogg, 2000).

Inflorescence Branching

The inflorescence results from the activity of its own meristems, which defines the
production or loss of iterative structures, and is finally determined by the number, size
and relative arrangement of their branches (Guarise & Vegetti, 2007). The architec-
ture of the inflorescence depends on its branching pattern (Vegetti, 2003). The
unispicate inflorescence (Fig. 2E, G) lacks branches and consists of the terminal
spikelet only. This reduction may concern an entire genus and characterize its type of
inflorescence (as in Eleocharis), or it may only happen in some species of a genus
(e.g. Bulbostylis, Goetghebeur & Gröger, 1992; Isolepis, Vegetti, 1994) and even
within the same species, as in Abildgaardia ovata (Burm. f.) Kral, Bulbostylis
capillaris (L.) C.B. Clarke var. microstachys (Boeck.) Barros, B. brevifolia Palla
(Reutemann et al., 2009), Schoenoplectus pungens (Vahl) Palla var. longispicatus
(Britton) S.G. Sm. (Vegetti, 1992) and Bolboschoenus maritimus (L.) Palla
(Browning & Gordon Gray, 1999).
However, in most species, the meristem forms lateral branches (primary branches).
In some Cyperaceae, these primary branches are reduced to its terminal spikelet.
When the internode below the spikelet (pedicel 0 hipopodium) is not elongated, the
Inflorescence of Cyperaceae 189

Fig. 2 Different forms of Cyperaceae inflorescences: A Paniculodium; B Anthelodium; C Capituliform

inflorescence or cefalodium; D Stachyodium or spike of spikelets; E, G Inflorescences constituted by only
one spikelet, with involucral bracts (E) and without these bracts (G); F Truncated inflorescence; H Corymb
of spikelets. References: b, bract; coflorescence (grey circle); main florescence (black circle); lbr, long
branch; pr, prophyll; shbr, short branch
190 A. Reutemann et al.

Fig. 3 Branching pattern: A, B Prophyllar branching; C, D Serial branching; E, F Prophyllar and serial
branching; G, H Collateral branching. References: b, bract; br, br´, branches of successive orders; pr,
prophyll

inflorescence is a spike of spikelets (Fig. 2D). The spike of spikelets is observed in

Kyllinga , Lipocarpha and Ascolepis. In Carex, a spike of spikelets is also considered
Inflorescence of Cyperaceae 191

a specialized inflorescence (Reznicek, 1990); it is the result of one advanced degree

of reduction and it is one of the most derivative types of inflorescence in the family
(Guarise & Vegetti, 2007).
In other genera, the primary branches go on to branch again. Generally, there is a
gradual transition in the number of spikelets in each inflorescence and in the branch-
ing order of the inflorescence branches (Fig. 2A). The branching degree becomes
distally reduced and the branches become progressively shorter toward the apex
(Fig. 2A). Therefore, the basal and median branches are branched, while the distal
branches are not branched and reduced to their terminal spikelet. In many
inflorescences of Cyperaceae, most of the primary branches have a lower similar
order of branching, except for the very distal ones, which are reduced either to
the minimum branching degree or to the terminal spikelet. These inflorescences
with many branches of similar branching order are homogenized inflorescences
(see next section).
For a proper interpretation of the inflorescence, it is important to know the
inflorescence ramification pattern (Haines, 1966; Meert & Goetghebeur, 1979;
Vegetti & Tivano, 1991) and the branch position in the inflorescences (Guarise &
Vegetti, 2007). In the Cyperaceae inflorescence, three types of branching can be
observed, namely (1) normal branching: the branch is produced by an axillary bud of
a bract (Fig. 2A); (2) prophyllar branching: the prophyllar branch is produced by a
prophyllar bud (Fig. 3A, B; E, F); (c) accessory branch: new branches are observed
between an axillary branch and its bract (Fig. 3C–H). These accessory branches lack
their own subtending bract; because of this, only one bract protects this entire
ramification.
Prophyllar productions may have varying degrees of development in different
species and even within one species (Reutemann et al., 2009); also, they may repeat
themselves several times; thus, the prophyllar branching pattern may or not result in a
whole series of prophyllar branches.
In the accessory branches, each of these accessory axes has its own prophyll,
in whose axils a prophyllar branch can be produced. The accessory pattern is
produced by serial or collateral buds. In the serial branches of Cladium P. Browne
(Mora-Osejo, 1960), variations regarding the extent of development of these buds
may be observed in different positions of the inflorescence: in the lowermost
inflorescence region, the bottom enrichment axis is stronger and has more
ramifications, while the opposite takes place in the distal region, where the top
shoot is the one showing the highest degree of development. This means that
accesory serial buds follow an upward pattern in one case and a downward
pattern in the other.
Normal and prophyllar branching patterns have been described by several
authors (Blaser, 1944; Haines, 1966; Guaglianone, 1970, 1980, 1981, 1982; Meert
& Goetghebeur, 1979; Bruhl, 1995; Goetghebeur, 1998; Vegetti & Guaglianone,
2005). Branches arising from accessory serial buds are observed in Hypolytrum
(Alves et al., 2000), Cladium mariscus (L.) Pohl (Mora-Osejo, 1960) and Cyperus
(Guarise & Vegetti, unpublished), while supernumerary collateral buds are observed
in Schoenus ferrugineus L. (Mora-Osejo, 1960) and Coleochloa setifera (Ridl.) Gilly
(Kukkonen, 1986). Prophyllar and accessory-axillary productions can be either
branched or reduced to their terminal spikelet.
192 A. Reutemann et al.

Although normal branching takes place along the inflorescence, the presence and
position of prophyllar and accessory-axillary branches in the inflorescences may vary.
In this context, whereas prophyllar branching is an important feature of the main
branching system of the inflorescence, it is relatively rare in the spikelet clusters
(Haines, 1966), except in the spikelet clusters of some species of Cyperus section
Luzuloidei (Guarise & Vegetti, 2007), in Bulbostylis (Reutemann et al., 2009) and
Rhynchospora (Haines, 1966). In different species, prophyllar and accessory-axillary
axes may or may not be arranged in different ways along the main inflorescence axis
and branches. In Bulbostylis, the development of prophyllar productions does not
occur in a given region of the inflorescence, whereas in Cyperus incomtus Kunth var.
incomtus and C. ochraceus Vahl, these branches are located in the distal region of the
main axis. In some species of Cyperus section Luzuloidei Kunth, accessory-axillary
axes of the 1st order can either develop from the proximal region upwards throughout
the main axis (e.g. C. eragrostis Lam. var. compactus (E. Desv.) Kük., C. fraternus
Kunth, C. reflexus Vahl and C. surinamensis Rottb.), in the median-distal region (C.
eragrostis var. eragrostis, C. intricatus Schrad. Ex Schult., C. luzulae (L.) Retz., C.
virens Michx. var. drummondii (Torr. & Hook.) Kük. and C. virens var. virens) or just
in the distal region of the main axis (C. entrerianus Boeck., C. hieronymi Boeck., C.
virens var. montanus (Boeck.) Denton). The presence or lack of prophyllar and
accesory branches, as well as their position, have proved to be useful as taxonomic
characters in some species of Cyperus section Luzuloidei (Guarise & Vegetti, 2007);
whereas the presence or lack of prophyllar productions, as well as their degree
of development, have turned out to be both taxonomically and phylogenetically
useful in some species of Bulbostylis (Guaglianone, 1970; López & Reutemann,
unpublished).

Inflorescence Homogenization Degree

Homogenization is the process by which some or all branches positioned in consec-

utive nodes along one axis have the same ramification degree and become similar to
one another (Rua, 1999). Homogenization may be total or partial. In a fully homog-
enized inflorescence (homogenized inflorescence), the degree of ramification is the
same for all branches. In a partially homogenized inflorescence, only the distal and
median parts have been affected by the homogenization process, while the basal
branches are more extensively branched with respect to the distal and median
branches (Rua, 1999). In homogenized inflorescences, two types of branches may
be recognized: long branches (lBr 0 long paraclade) and short branches (shBr, the
distal one reduced to the terminal spikelet).
In homogenized inflorescences (Fig. 2B), the axillary meristems made up by the
BMs only form SMs, which, in turn, generate branches reduced to one spikelet. After
generating a certain number of primary branches with secondary branches reduced to
one spikelet, the IM of the homogenized inflorescence switches abruptly to produce a
certain number of branches reduced to their terminal spikelet. The BMs are indeter-
minate meristems and the SMs are determinate meristems. Indeterminate meristems
make an indefinite number of organs, whereas determinate meristems are consumed
after making a specific number of organs (Vollbrecht et al., 2005; Bortiri & Hake,
2007). The homogenization process of the inflorescence is related to the meristem
Inflorescence of Cyperaceae 193

determinancy (Perreta et al., 2009a). In this sense, the meristem determinancy would
guide the degree of homogenization (Salariato et al., 2010).
The homogenization process is usually associated with the truncation process
(Perreta et al., 2009a); however, this association is rare in Cyperaceae (see next section).

Presence or Absence of the Distal Part of the Inflorescence

After producing a certain number of branches, the IM generally ends in a terminal

spikelet (Fig. 2A–C). In a few species, the IM forms a variable number of
branches and then stops and does not develop the distal floral structures
(Fig. 2F). The process characterized by the lack of formation of the distal floral
structures has been called “truncation” (Troll, 1964; Weberling, 1989). Truncation
may affect the development of the terminal spikelet only, or it may also affect
the development of the distal region with short branches and some long branches
(Perreta et al., 2009a).
Although truncation is not frequent in this family, a few species show truncated
inflorescences. Truncation in Cyperaceae may affect only the terminal spikelet on the
main axis (Cariceae, Reznicek, 1990; Vegetti, 2002), or it may also affect the primary
shBrs (Fig. 2F) (Vegetti, 2002) and the primary lBrs (e.g. Cyperus papyrus L., C.
prolifer Lam., C. giganteus Vahl (Fig. 2F, Mora-Osejo, 1960; Raynal, 1971; Haines
& Lye, 1983; Perreta & Vegetti, 2002; Vegetti, 2003). On the lBrs, truncation may
affect the terminal spikelet and the shBrs (Perreta & Vegetti, 2002; Vegetti, 2003).
In some families, such as grasses, truncation is a very common process which is
frequently associated with homogenization, and both processes account for much of
the diversity among grass inflorescences (Reinheimer & Vegetti, 2008; Perreta et al.,
2009b). However, while most inflorescences show varying degrees of homogeniza-
tion in Cyperaceae, truncation is not common in this family. In those few species of
Cyperaceae where truncation does occur, inflorescences are homogenized (Fig. 2F).

Phyllotaxis and Branch Disposition

Phyllotaxis varies along the synflorescence (main shoot and branches); the disposi-
tion of the leaves in the vegetative zone is tristichous (phyllotaxis 1/3), whereas in the
inflorescence branch zone the bracts, and their branches, have a spiral arrangement.
The inflorescence primary branches emerge in an alternate arrangement, one per
node; however, some inflorescences show branches with a subopposite or pseudo-
verticillate arrangement (primary branches disposed in nodes very close around the
main axis). In the inflorescence of some species of Cyperus (Guarise & Vegetti,
2007), the formation of more than three orthostichies can be observed. Each of these
orthostichies is made up by branches from the basal and distal regions of the
inflorescences; whereas the branches of the median region are not positioned over
any orthostichies. If the number of primary branches on the main axis is not nine or
more than nine, there are no orthostichies (e.g. some C. incomtus var. incomtus
inflorescences). In the axillary and prophyllar branching, the disposition of branches
can follow a right-hand or left-hand spiral; showing an antidromic arrangement. This
antidromic arrangement is not observed in the accessory-axillary ramification
(Guarise & Vegetti, 2007).
194 A. Reutemann et al.

In the last branching order (spikelet), the glume phyllotaxis can be either spiral
(e.g. Schoenoplectus (Rchb.) Palla) or distichous (e.g. Cyperus). In Cyperus sect.
Luzuloidei (Guarise & Vegetti, 2007), all the glumes have a distichous arrangement
(1/2 phyllotaxis), except for the prophyll. The distichous disposition of the glumes
would indicate a change in the inflorescence phyllotaxis, from a spiral arrangement
(3/8 phyllotaxis) to a distichous phyllotaxis (1/2 phyllotaxis), and this switch may be
mediated by the prophyll of the spikelets in a transitional position, because it forms a
90 ° angle with respect to the glumes (Guarise & Vegetti, 2007).
In species of Scleria, the glumes are disposed distichously in the pistillate spike-
lets, while the subandrogynous spikelet, staminate spikelet, and androgynous spikelet
have one proximal portion with glumes disposed distichously and one distal portion
with glumes disposed spirally (Camelbeke, 2002; Ahumada & Vegetti, 2009); this
arrangement of the glumes is also characteristic of the bisexual spikelets of
Abildgaardia ovata. In species of Scleria with the latter type of spikelet—considering
each axis separately (the main axis and the branched paracladia)—, the phyllotaxy
pattern changes first from spiral (in the vegetative part) to distichous and then
again from distichous to spiral. The transition from the tristichous disposition,
being characteristic of the leaves of the vegetative region, to the spiral arrange-
ment in the inflorescence is a relevant feature that should be noted (Guarise &
Vegetti, 2008). However, a double and bidirectional change in the phyllotaxy
pattern along the same morphological axis is very unusual, as it occurs in the
inflorescences of Scleria with subandrogynous, staminate, or androgynous spikelets.
Extensive spikelet and floral ontogenetic research is needed to clarify these phyllo-
taxy changes.
The transition from the tristichous disposition, being characteristic of the leaves of
the trophotagma region, to the spiral arrangement in the paracladial zone is a relevant
feature that should be noted (Vegetti & Anton, 1995; Cámara-Hernández, 2001a, b;
Reinheimer & Vegetti, 2004; Reinheimer et al., 2005; Kern et al., 2008).

Inflorescence Position

The inflorescence position as related to the culm can be terminal or pseudolateral.

When the inflorescence takes a pseudolateral position, the lower bracts and proximal
inflorescence branches point into the same direction as the stem, turning the main axis
aside. In this type of inflorescence, the first bract may be culmlike (e.g. Schoeno-
plectus: Vegetti, 2002) or foliaceous (e.g. Cyperus surinamensis, unpubl.). In Cyper-
aceae, pseudolateral inflorescences evolve independently more than once and can
occur in isolation, as in the Cyperus genera (Heinzen & Vegetti, 1994; Guaglianone,
1996), or be a feature for all the species belonging to a genera, as in Schoenoplectus
(Vegetti & Tivano, 1991; Vegetti, 1992) and Isolepis (Vegetti, 1994).
In terminal inflorescences, the main axis goes on the same direction as the stem. In
some cases, in species with terminal inflorescences, the bracts and inflorescence
branches are erect, as in the pseudolateral inflorescences (e.g. Cyperus intricatus).
This feature, which has been observed in the family (Bruhl, 1995), could be consid-
ered as an intermediate position between terminal and pseudolateral. Some authors
only refer to the pseudolateral arrangement when the bract goes on the same direction
as the stem (Haines & Lye, 1983; Goetghebeur, 1998) and make no reference
Inflorescence of Cyperaceae 195

whatsoever to the inflorescence main axis. This type of consideration has led to
considering one true terminal inflorescence as pseudolateral.

Bracts and Leaves Subtending Branches

The inflorescence main axis bears bracts that decrease in length up the axis;
each bract subtends an axillary bud that produces a primary lateral inflorescence
branch. Each primary branch has a prophyll and a variable number of bracts.
Buds in the axils of the bracts and, in some species, in the axils of the
prophylls expand and repeat the branching pattern in the successive branches
(Richards, 2002; Vegetti, 2003).
Different types of bracts may be recognize in the inflorescence: (1) foliaceous,
with a well-developed blade and a scarcely or not developed sheath; (2) setiform,
without a sheath, the leaf blade enlarges at the base but the mid and upper portions are
narrow and stretched (setalike), sometimes acuminate; and (3) glumaceous, similar to
a glume, but with a short setiform appendix.
The length reduction of the inflorescence internodes of different order and the
reduction of the branching degree of the inflorescence branches are generally corre-
lated with the reduction in the shape of the bracts (Guarise & Vegetti, 2007). Because
of this, the main axis may show, in acropetal succession, all three types of bracts
(foliaceous-setiform-glumaceous), or two types (foliaceous-glumaceous, setiform-
glumaceous) or only the glumaceous type. Given the clear acropetal development
of its componentes, every inflorescence has glumaceous bracts, both in the distal
portions on the main axis and on the inflorescence branches.
The primary bracts are often described as involucral bracts, the lowermost bracts
are usually foliaceous (sheathing or not) or setiform, and the uppermost bracts are
very small and glumaceous; however, not rarely, primary bracts can be all foliaceous
(Cyperus imbricatus Retz.) or all glumaceous (Cyperus papyrus). The bracts of the
ultimate branchlets (spikelet bracts) are often barely distinct from glumes (Goetghebeur,
1998). In some species, bracts can be patent or reflex; also, in some species, the lower
bracts can be upright, continuing the culm and culmlike (see Inflorescence Position).
Lower bracts may be longer or shorter than the inflorescence.
The prophylls can be well developed, but they are often reduced or even absent.
They show an acropetal variation in size and form: tubular (cladoprophyll), laminar
and glumaceous. All of them are two-keeled, a hardly observable character in the
glumaceous ones. The length of the lowermost prophylls varies among species,
whereas the length of the distal ones (glumaceous) is smaller in all species studied.
The occurrence of the different types of bracts and prophylls varies according to
the region of the inflorescence (Guarise & Vegetti, 2007). Some authors (Haines,
1966; Tucker, 1983) consider the length and shape of the bracts to bear taxonomic
significance.

Elongation of the Inflorescence Internodes

The length variation in the internodes both on the main axis and on the branching
axes of different order determines important variations in the external appearance of
the Cyperaceae inflorescence. Internodes lengthening is related to the time of
196 A. Reutemann et al.

development of each species (Tucker & Grimes, 1999); it is genetically determined

(Kellogg, 2000).
In Cyperaceae, the inflorescence main axis may show long or short internodes,
determining the so-called panicle (Fig. 2A) and anthela (Fig. 2B) respectively or, in
the sense of Troll’s terminology, the paniculodium and the anthelodium (Rua, 1999;
Guarise & Vegetti, 2008). Raynal (1971) and Goetghebeur (1998) consider the
panicle of spikelets as the basic cyperaceous inflorescence which can be modified
by the contraction of the internodes and various reductional trends.
In the inflorescence branches, the internode length is variable; with the epipodium
being the internode showing the main variations. The variations in the anthela of
spikelets depend on the length of the epipodium of their branches (see Inflorescence
Form). A capitate inflorescence may derive from a paniculodium, an anthelodium, or
from a spike of spikelets because of the reduction of the internode length of the main
axis and the inflorescence branches. There is no reason to suppose that the reverse
pathway might happen to produce a type of inflorescence with an epipodium devel-
oped from any more congested form (Guarise & Vegetti, 2007).
The internode growth from the inflorescence branch axes (except the epipodium)
affects the inflorescence shape, especially in the manner in which branches group in
the distal portion of an inflorescence branch with a developed epipodium; these
groups can be simply described as a contracted head or glomerulous (e.g. Cyperus
sect. Luzuloidei), digitate inflorescences (C. laxus Lam.), lax (C. esculentus L.) or
congested spikes (C. aggregatus (Willd.) Endl).

Structure of the Spikelet

In Cyperaceae, the spikelet is the morphological unit of the inflorescence (Eiten,

1976). Studies of inflorescences in Cyperaceae focus mainly on the spikelet, whose
reduced size and complexity have led to varying interpretations. In this context, there
is much controversy over the spikelet in the family and until now its interpretation has
remained unclear (Vrijdaghs, 2006). This author considers that this confusion par-
tially results from old discussions influenced by the euanthial and pseudanthial
hypotheses and related monopodial or sympodial interpretation of spikelets. There
is plenty of literature on the structure and development of flowers and spikelets in the
family (Payer, 1857; Pax, 1886; Celakovsky, 1887; Schönland, 1922; Mattfeld, 1938;
Blaser, 1941, 1944; Holltum, 1948; Mora-Osejo, 1960, 1987; Kern, 1962; Raynal,
1971; Meeuse, 1975a, b; Eiten, 1976; Haines & Lye, 1976; Kukkonen, 1986; Meert
& Goetghebeur, 1979; Goetghebeur, 1986, 1998; Bruhl, 1991, 1995; Muasya, 1998;
Timonen, 1998; Vrijdaghs et al., 2003, 2004, 2005a, b, c, 2007, 2009, 2010; Zhang et
al., 2004a, b; Richards et al., 2006), that is why this subject is not covered in this
review. Based on findings from many ontogenetic studies (Vrijdaghs et al., 2003,
2004, 2005a, b, c, 2007, 2009; Richards et al., 2006; Reutemann et al., 2009), we
consider spikelets to be monopodial ramifications.
Many variations which are valuable from a taxonomic or systematic point of view
have been observed in spikelets (see Eiten, 1976; Bruhl, 1995). The most important
variations include: a) flower sexuality: spikelets with male and female flowers may be
gynandrous (with the female flower distal), androgynous (with the male flower distal)
and mesogynous (with female flowers proximal and distal to the male flowers); b)
Inflorescence of Cyperaceae 197

shape (shortly ovate, elliptical, lanceolate, linear, ovate, etc); c) compression (laterally
or dorsiventrally compressed or more o less terete); d) rachilla internodes more o less
straight, zigzag or flexuose; e) rachilla prolongation exceeding (e.g. Uncinia Pers.) or
not the flowers; f) rachilla persistent or deciduous at maturity; g) type of rachilla
disarticulation at maturity (not disarticulating, disarticulating above the prophyll,
disarticulating below the prophyll, shattering with points of abscission at the base
of each fruit or not shattering); h) rachilla with wings or wingless, i) wings deciduous
or persistent; j) glumes spirally or distichously disposed; and k) glumes persistent or
deciduous (completely, incompletely, individually or collectively deciduous).

Hypothesized Evolutionary Processes

Reduction is the most frequent way of modification in the inflorescences of Cyper-

aceae (Thomas, 1984; Zhang, 2001), and it is considered the main trend of synflor-
escence evolution among and within the tribes (Guarise & Vegetti, 2008). The
reductive processes do not always affect equivalent areas. Indeed, reductive processes
can occur in branches, resulting in less complex inflorescences, as in supradecom-
pound and simple anthelas in Cyperus; they can affect the proximal region of the
inflorescence and complex branches, resulting in a spike of spikelets (with this
inflorescence being the most derived stage in Cyperaceae, Guarise & Vegetti,
2008); or they may affect the entire inflorescence, which is then reduced to its
terminal spikelet. In other cases, it is the distal part of the inflorescence that is
reduced in a sequence that can affect either the terminal spikelet alone , the short
branches or some of the long branches (Fig. 2F), as occurs in truncated inflorescences
of Cariceae (Vegetti, 2002) and Cyperus giganteus (Perreta & Vegetti, 2002).
Truncation is not frequent in this family.
The broad diversity shown by inflorescences in the family can be explained by the
analysis of some processes that operate in different ways, either combined or inde-
pendently (Guarise & Vegetti, 2007). These authors characterize 25 processes that
have contributed to variations in the Cyperaceae inflorescence. These processes may
be grouped into the following classes: (1) reduction/elongation of the internodes; (2)
branch development; (3) reduction/development of foliar structures; (4) homogeni-
zation; and (5) truncation.
Several trends and processes create morphological differences and similarities
between and within the tribes, and even in the same genus. This may cause parallel-
ism and reversions in the evolutionary course. Examples of parallelism can be
observed in the synflorescence of different genera such as Chrysitrix L. and Eleo-
charis R. Br., among certain synflorescences of Hypolytrum (Alves, 2003) and
Eriophorum L., or among Scleria, Fuirena and Rhynchospora (Guarise & Vegetti,
2008). Ficinia Schrad. and Bolboschoenus (Asch.) Palla might be good examples of
reversion because these genera show synflorescences with a foliate stem (considered
an ancestral state) but belong to clades where the synflorescence has all basal leaves
and the inflorescence is supported by a scape (considered a derived state) (Guarise &
Vegetti, 2008).
Beyond parallelism and reversion within the inflorescence architecture, it is
possible to identify groups of taxa in which particular types of inflorescences have
198 A. Reutemann et al.

evolved (Stebbins, 1974; Tucker & Grimes, 1999). This allows to identify basic
structures for a given group and to understand the way in which the processes have
changed the synflorescence architecture.

Inflorescence Form

The following inflorescence forms have been described in the Cyperaceae family:

– Panicula of spikelets (or paniculodium): conical indeterminate inflorescence;

the terminal spikelet and the distal branches emerge distinctly higher than the
proximal branches (Fig. 2A) (e.g. the panicle-like inflorescences of Hypolytrum,
Alves et al., 2000; Alves, 2003). In the panicula of spikelets, the branches may
have their internodes more or less lengthened or completely shortened, looking
like lateral capitula or heads (densely paniculate inflorescence; Scirpodendron
Zipp. ex Kurz, Goetghebeur, 1998).
The panicula of spikelets, as an inflorescence, evolved early in basal groups in the
phylogeny of the family, as in some species of Hypolytrum (Hypolytreae I), Scleria
and Rhynchospora. Raynal (1971) and Goetghebeur (1998) consider the panicle as
the basic cyperaceous inflorescence which can be modified through the elongation or
contraction of the internodes in various reduction trends. This type of inflorescence is
also observed in Juncaceae (e.g. Oxychlöe Phil. and Luzula DC.; Pedersen, 1968;
Barros, 1969) and it can be considered plesiomorphic based on the outgroup com-
parison criterion (Guarise & Vegetti, 2008).

– Anthela of spikelets (or anthelodium): crateriform indeterminate inflorescence,

with the terminal spikelet and the short and distal branches hidden among the
long and proximal ones, which overtop them. This results from the inhibited
lengthening of the main axis internodes and the distal branch epipodium, with an
important development of the basal branch epipodium (Fig. 2B). The anthela of
spikelets may be simple, compound, decompound and supradecompound
depending on the branching order with expanded epipodium (either first, second,
third, or fourth and above, respectively). This variation in the form of the
anthela of spikelets has been described in species of Cyperus (Wilson, 1991;
Guaglianone, 1996; Guarise & Vegetti, 2007, 2008).
– Corymb of spikelets (or corymbodium): indeterminate inflorescence with the
primary branch spikelets arranged like the flowers of a corymb (Rua, 1999). In
this inflorescence, the variable epipodium length determines the spikelets to be
disposed all at the same level (Fig. 2H). Haines & Lye (1983) describe terminal
and lateral corymbs in Rhynchospora corymbosa (L.) Britton.
– Umbel of spikelets (or sciadodium or umbelliform inflorescence): indetermi-
nate inflorescence, with the spikelets disposed like the flowers of an umbel (Rua,
1999). Haines & Lye (1983) only recognize as true umbels the truncated inflor-
escences of Cyperus papyrus and C. prolifer. We consider that true umbelliform
inflorescences do not exist in Cyperaceae. In the umbel, the stalks arise from a
same point (Weberling, 1989), while in umbelliform Cyperaceae inflorescences,
the branches follow a one-per-node disposition (alternately arranged) and are
Inflorescence of Cyperaceae 199

separated by very short internodes. In this context, the inflorescence of C.

papyrus and C. prolifer depicted as a true umbelliform inflorescence is a
truncated anthela of spikelets as it has been described for C. giganteus
(Perreta & Vegetti, 2002).
– Capituliform inflorescence (or capitate inflorescence or cephalodium): inde-
terminate inflorescence, similar to a capitulum or head, due to a pronounced
shortening of the internodes on the main axis and branches of different order. In
some cases, there is also a reduction of the branching degree (Fig. 2C). Many of
the above described inflorescences may be reduced to a capitate inflorescence; in
many cases, probably due to unfavorable environment conditions (Raynal, 1971;
Browning & Gordon-Gray, 1999; Rua, 1999).
– Spike of spikelets (or stachyodium): indeterminate inflorescence with spikelets
disposed sessile on the main axis (Fig. 2D). In some cases, the terminal
spikelet is absent (truncation) and only branches reduced to final spikelets
remains (Cariceae, Vegetti, 2002). The spike of spikelets can be composed of
spikelets with perfect flowers (e.g. Schoenoplectus, Isolepis and Cyperus) or
unisexual flowers (e.g. Cariceae). In some spikes of spikelets, the unisexual
flowers may form bisexual spikelets (male and female flowers on a same rachilla)
or unisexual spikelets (male and female flowers on different rachillae) (Meert &
Goetghebeur, 1979; Timonen, 1998). The spikes with unisexual and bisexual
spikelets (in given cases) may be gynandrous (i.e. with the female-only spikelet
distal) or androgynous (i.e. with the male-only spikelet distal) (Bruhl, 1995).

In extremely complex inflorescences, the distal region of the main axis, as well as
the distal region of the many branches (especially the basal branches with long
epipodium), can also show a branch arrangement similar to one of the above
described forms. This leads to a set of combinations including: panicula of congested
spikes (e.g. Carex uruguensis Boeck.; Carex trachycystis Griseb.; Scirpodendron),
anthela of spikes of spikelets (e.g. Cyperus digitatus Roxb.), anthela of fascicles of
spikelets, anthela of capitula (e.g. Bulbostylis consanguinea (Kunth) Clarke, B.
junciformis (H.B.K.) Clarke, Cyperus entrerrianus); anthela with monostachius axis
(e.g. Fimbristylis robusta K. Lye; F. dipsaceae (Rottb.) C. B. CL).
Within the various forms that have been characterized, it is important to describe,
in turn, subtypes of inflorescences according to: (1) the truncation process (truncated
or non-truncated inflorescences, Perreta & Vegetti, 2002); (2) the homogenization
proccess (fully homogenized inflorescences, partially homogenized inflorescences,
non-homogenized inflorescences, Guarise & Vegetti, 2007; Lucero et al., 2009;
Reutemann et al., 2009).

Conclusions

The structural diversity of the Cyperaceae inflorescences may be analized at the

spikelet level or at the inflorescence branch level. In this review, we analyze the
variations at the inflorescence and synflorescence branch level. The most significant
variations that affect the inflorescence branches determine that inflorescences may be
composed of numerous spikelets, arranged in a more or less complex branching
200 A. Reutemann et al.

system, or consist of a few spikelets or even one spikelet. Variations at the level of the
inflorescence branches are related to:
& number of primary inflorescence branches;
& number of the primary branches in each node: alternate, subopposite or pseudo-
verticillate branches;
& development degree of the primary branches:
– reduced to the terminal spikelet,
– formed by the terminal spikelet and a variable number of branches of
consecutive order (only secondary branches or branches of order n);
& different elongation of the internodes of : (1) the main axis; (2) the epipodioum;
and (3) the rest of the internodes of the inflorescence branches;
& disposition of the branches of consecutive order;
& development of the bracts and prophylls;
& development of the prophyllary buds;
& development of the accessory buds;
& homogenization degree;
& truncation degree.
Variations at the synflorescence level are related to:
& presence of scape or of foliate stem;
& absence or presence of enrichment shoots and its axillary inflorescences in the
foliate stem.
In this work, we analize the structure of the mature inflorescence. The extensive
variation in adult inflorescences of Cyperaceae is the result of the diverse inflores-
cence development patterns, as it has been shown in Grasses (Doust & Kellogg,
2002; Reinheimer, 2007; Perreta et al., 2009a). Research on the evolution of morpho-
logical diversity of the Cyperaceae inflorescences in large clades and the combination of
developmental and mature structure studies are essential; these will contribute to a more
detailed interpretation of the inflorescences in the family’s different genera as well as to a
description of a larger morphological variability. Such studies will help to establish new
characters that may be useful in future taxonomic and phylogenetic studies.
It is important to characterize the inflorescence and the synflorescence properly.
For comparative purposes, only homologous structures should be compared,
that is, inflorescences should only be compared with inflorescences but not
with synflorescences.

Literature Cited

Ahumada, O. & A. C. Vegetti. 2009. Inflorescence structure in species of Scleria subgenus Hypoporum
and subgenus Scleria (Sclerieae-Cyperaceae). Plant Systematics and Evolution 281: 115–135.
Albert, V. A., Gustafsson M. H. & Di Laurenzio L. 1998. Ontogenetic systematics, molecular develop-
mental genetics, and the Angiosperm petal. Pp. 349–374. In: D. Soltis, P. Soltis & J. Doyle (eds.),
Molecular Systematics of Plants II, Chapter 12. Kluwer Academic Press, Boston.
Alves, M. V. 2000. Abordagem Tipológica das Inflorescencias em Monocotiledóneas. Tópicos Atuais em
Botánica. 51º Congreso Nacional de Botánica. Brasilia, DF. pp 17–22.
Inflorescence of Cyperaceae 201

——— 2003. Hypolytrum Rich. (Cyperaceae) nos Neotrópicos, Tese de Doutorado, Universidade de São
Paulo, São Paulo
———, M. Wanderley & W. W. Thomas. 2000. Typology of the inflorescence in species of Hypolytrum
Rich., (Cyperaceae) from Brazil. Beitrage zur Biologie der Pflanzen 72: 59–73.
Barros, M. 1969. Cyperaceae. In: M. Correa (ed.), Flora Patagónica T. VIII, Parte II. Colección Científica
del INTA.
Blaser, H. W. 1941. Studies in the morphology of the Cyperaceae I. Morphology of flowers. A. Scirpoid
genera. American Journal of Botany 28: 542–551.
——— 1944. Studies in the morphology of the Cyperaceae. II. The prophyll. American Journal of Botany
31: 53–64.
Bortiri, E. & S. Hake. 2007. Flowering and determinacy in maize. Journal of Experimental Botany 58:
909–916.
Browning, J. & K. D. Gordon-Gray. 1999. The inflorescence in southern African species of Bolboschoenus
(Cyperaceae). Annales Botanici Fennici 36: 81–97.
Bruhl, J. J. 1991. Comparative development of some taxonomically critical floral/inflorescence features in
Cyperaceae. Australian Journal of Botany 39: 119–127.
——— 1995. Sedge genera of the world: relationships and a new classification of the Cyperaceae.
Australian Systematic Botany 8: 125–305.
Cámara Hernández, J. 2001a. La polistiquia en las inflorescencias de las Poáceaes. Boletín de la Sociedad
Argentina de Botánica 36: 9–10
——— 2001b. Morfología de las inflorescencias de Diplachne y Leptochloa (Poaceae Boletín de la
Sociedad Argentina de Botánica 36: 267–278
Camelbeke, K. 2002. Morphology and taxonomy of genus Scleria (Cyperaceae) in Tropical South
America. Tesis Doctoral. Gent. Belgium.
Celakovský, L. 1887. Über die ährchenartigen Partialinfloreszenzen der Rhyncosporeen. Berichte der
Deutschen Botanischen Gesellschaft 5: 148–152.
Doust, A. N. & E. A. Kellogg. 2002. Integrating phylogeny, developmental morphology and genetics: a
case study of inflorescence evolution in the “bristle grass” clade (Panicoideae: Poaceae). Pp 298–314.
In: Cronk Q. C. B., R. M. Bateman & J. A. Hawkins (eds), Developmental Genetics and Plant
Evolution. Ed. Taylor & Francis, London.
Eiten, L. T. 1976. Inflorescence units in the Cyperaceae. Annals of the Missouri Botanical Garden
63: 113–119.
Goetghebeur, P. 1986. Genera Cyperacearum. Dr. Sci. Thesis, State University, Ghent.
——— 1998. Cyperaceae. In: Kubitzki, K., H. Huber, P. J. Rusall, P. S. Stevens & T. Stüzel (eds), The
families and genera of plants, 4: 141–190. Springer Verlag, Berlin.
——— & A. Gröger. 1992. Studies in Cyperaceae II. The unispiculate species of Bulbostylis in the
Americas. Dodonaea 60: 64–75.
Guaglianone, E. R. 1970. Un nuevo carácter, útil en la distinción genérica entre Fimbrystilis Vahl y
Bulbostylis Kunth (Cyperaceae). Darwiniana 16: 40–48.
——— 1980. Contribución al estudio del género Rhynchospora Vahl (Cyperaceae) II. Darwiniana 22:
449–509.
——— 1981. Contribución al estudio del género Rhynchospora Vahl (Cyperaceae) III. Darwiniana 23:
489–506.
——— 1982. Contribución al estudio del género Rhynchospora Vahl (Cyperaceae). IV. Rhynchospora
iberae nueva especie de América Austral. Darwiniana 24: 469–473.
——— 1996. Cyperaceae. In: Lahitte, H. B. J. & A. Hurrell (eds), Las plantas hidrófilas de la isla Martín
García Buenos Aires. Ser Informes 52: 70–96.
Guarise, N. J. & A. C. Vegetti. 2007. The Inflorescences structure of Cyperus L. section Luzuloidei Kunth
(Cyperaceae). Plant Systematic and Evolution 271: 41–63.
——— & ——— 2008. Processes responsible of the structural diversity of the Cyperaceae inflorescence:
hypothetical evolutionary trends. Flora 203: 640–647.
Haines, R. W. 1966. Prophylls and branching in Cyperaceae. Journal of the East Africa Natural History
Society XXVI (113): 51–78.
——— & K. A. Lye. 1976. Studies in African Cyperaceae XlV. The genus Hellmuthia Steud. Botaniska
Notiser 129: 61–68.
——— & ——— 1983. The sedges and rushes of East Africa. East African Natural History Society,
Nairobi. 404 pp.
Heinzen, F. A. & A. C. Vegetti. 1994. Typology of the Inflorescence in Cyperus corymbosus var.
subnodosus and C. rotundus (Cyperaceae). Beiträge zur Biologie der Pflanzen 68: 263–273.
202 A. Reutemann et al.

Holttum, R. E. 1948. The spikelet in Cyperaceae. Botanical Review 14: 525–541.

Kellogg, E. A. 2000. Molecular and morphological evolution in Andropogoneae. Pp. 149–158. In: W. L.
Jacobs & J. E. Everett (eds.), Grasses: Systematics and Evolution (S.). CSIRO, Melbourne, Australia.
Kern, J. H. 1962. A new look at same Cyperaceae mainly from the tropical standpoint. Advancement of
Science 19: 141–148.
Kern, V., N. Guarise & A. Vegetti. 2008. Typology of the inflorescence in Spartina (Poaceae:
Eragrostoideae). Plant Systematic and Evolution 273: 51–61.
Kukkonen, I. 1984. On the inflorescence structure in the family Cyperaceae. Annales Botanici Fennici 21:
257–264.
——— 1986. Special features of the inflorescence structure in the family Cyperaceae. Annales Botanici
Fennici 23: 107–119.
——— 1994. Definition of descriptive terms for the Cyperaceae. Annales Botanici Fennici 31: 37–43.
Lucero, L. E., O. Ahumada, A. G. Reutemann & A. C. Vegetti. 2009. Filogenia morfológica de las
especies sudamericanas de Scleria Bergius (Cyperaceae). XXXII Jornadas Argentinas de Botánica.
Huerta Grande (Córdoba). Pp. 203.
Mattfeld, J. 1938. Das morphologische Wesen und die phylogenetische Bedeutung der Blumenblätter.
Berichte der Deutschen Botanischen Gesellschaft 56: 86–116.
McSteen, P. 2009. Hormonal Regulation of Branching in Grasses. Plant Physiology 149: 46–55.
Meert M. & P. Goetghebeur. 1979. Comparative floral morphology of Bisboeckelereae and Cariceae
(Cyperaceae) on the basis of the anthoid concept. Bulletin de la Societe Royale Botanique de Belgique
112: 128–143.
Meeuse, A. D. J. 1975a. Changing floral concepts: anthocorms, flowers, and anthoids. Acta Botanica
Neerlandica 24: 23–36.
——— 1975b. Interpretative floral morphology of the Cyperaceae on the basis of the anthoid concept. Acta
Botanica Neerlandica 24: 23–36.
Mora-Osejo, J. E. 1960. Beitträge zur Entwicklungsgeschichte und vergleichenden Morphologie der
Cyperaceen. Beiträge zur Biologie der Pflanzen 35: 293–341.
——— 1987. Estudios morfológicos, autoecológicos y sistemáticos en Angiospermas. Academia
Colombiana de Ciencias Exactas, Físicas y Naturales, Ser. J. Alvarez Lleras 1, Bogotá.
Muasya, A. M. 1998. A synopsis of Fuirena (Cyperaceae) for the flora of tropical East Africa. Kew
Bulletin 53: 187–202.
———, D. A. Simpson, M. W. Chase & A. Culham. 1998. An assessment of suprageneric phylogeny in
Cyperaceae using rcbL DNA sequences. Plant Systematics and Evolution 211: 257–271.
———, ———, G. A. Verboom, P. Goetghebeur, R. F. Naczi, M. W. Chase & E. Smets. 2009.
Phylogeny of Cyperaceae Based on DNA Sequence Data: Current Progress and Future Prospects.
Botanical Review 75: 2–21.
Pax, E. 1886. Beiträge zur Morphologie und Systematik der Cyperaceen. Botanische Jahrbücher für
Systematik, Pflanzengeschichte und Pflanzengeographie 7: 287–318.
Payer, J. B. 1857. Traité d’organogénie comparée de la fleur. Masson: Paris.
Pedersen, T. M. 1968. Cyperaceae. Pp. 315–421. In: Cabrera, A. L. (ed.), Flora de la Provincia de Buenos
Aires. T. IV (I). Colección Científica del I.N.T.A. Buenos Aires.
Perreta, M. & A. C. Vegetti. 2002. The inflorescence of Cyperus giganteus Vahl (Cyperaceae). Feddes
Repertorium 113: 256–260.
———, J. Ramos & A. C. Vegetti. 2009a. Development and Structure of the Grass Inflorescence.
Botanical Review 75: 377–396.
———, ———, J. C. Tivano & A. Vegetti. 2009b. Descriptive characters of growth form in Poaceae – an
overview. Flora 206, in press.
Raynal, A. 1971. Quelques notes morphologiques sur les Cypéracées. Mitteilungen der Botanischen
Staatssammlung München 10: 589–603.
Reinheimer, R. 2007. Desarrollo y tipología de la inflorescencia de especies sudamericanas de Brachiaria
y Urochloa y sus implicancias sistemáticas y taxonómicas (Poaceae: Panicoideae: Paniceae). Tesis
doctoral. Facultad de Bioquímica y Ciencias Biológicas, Universidad Nacional del Litoral
——— & A. C. Vegetti. 2004. The inflorescence of South American species of Panicum subg.
Phanopyrum sect. Laxa (Poaceae: Panicoideae: Paniceae). Beiträge zur Biologie der Pflanzen 73:
185–199.
——— & ——— 2008. Inflorescence diversity and evolution in the pck clade (Poaceae: Panicoideae:
Paniceae). Plant Systematics and Evolution 275: 133–167.
———, R. Pozner & A. C. Vegetti. 2005. Inflorescence, spikelet, and floral development in Panicum
maximum and Urochloa plantaginea (Poaceae). American Journal of Botany 92, 565–575.
Inflorescence of Cyperaceae 203

Reutemann A. G., N. J. Guarise, M. G. López & A. C. Vegetti. 2009. Structure of inflorescences of

selected South American species of Abildgaardia Vahl, Bulbostylis Kunth and Fimbristylis Vahl
(Abildgaardieae-Cyperoideae-Cyperaceae). Plant Systematic and Evolution 283: 93–110.
Reznicek, A. A. 1990 Evolution in sedges (Carex, Cyperaceae). Canadian Journal of Botany 68:
1409–1432
Richards, J. 2002. Flower and spikelet morphology in sawgrass, Cladium jamaicense Crantz (Cyperaceae).
Annals of Botany 90: 361–367.
———, J. J. Bruhl & K. L. Wilson. 2006. Flower or spikelet? Understanding the morphology and
development of reproductive structures in Exocarya (Cyperaceae, Mapanioideae, Chrysitricheae).
American Journal of Botany 93: 1241–1250.
Rua, G. 1999. Inflorescencias. Bases teóricas para su análisis. Sociedad Argentina de Botánica. Buenos
Aires. 100 pp.
Salariato, D., F. Zuloaga, L. Giussani & O. Morrone. 2010. Molecular phylogeny of the subtribe
Melinidinae (Poaceae: Panicoideae: Paniceae) and evolutionary trends in the homogenization of
inflorescences. Molecular Phylogenetics and Evolution 56: 355–369.
Schönland, S. 1922. South African Cyperaceae. Memoirs of the Botanical Survey of South Africa 3,
Pretoria, S. Africa.
Sell, Y. 1980. Physiological and phylogenetic significance of the direction of flowering in inflorescence
complexes, Flora 169: 282–294.
Shimamoto, K. & J. Kyozuka. 2002. Rice as a model for comparative genomics of plants. Annual Review
of Plant Biology 53: 399–419.
Simpson, D. A. 1992. A revision of the genus Mapania (Cyperaceae) . Royal Botanic Gardens, Kew, UK.
——— 1995. Relationships within Cyperales. Pp. 497– 509. In: Rudall P. J., P. J. Cribb, D. F. Cutler &
C. J. Humphries (eds). Monocotyledons: systematics and evolution. Royal Botanic Gardens
Kew, UK.
Stebbins, G. L. 1974. Flowering Plants: Evolution above the species level. Belknap Press, Cambridge,
M.A. 399 p.
Strong, M. T. 2001. Novelties in Rhynchospora (Cyperaceae) from the Guianas. Novon 11:261–273.
Thomas, W. W. 1984. The systematics of Rhynchospora sect. Dichromena. Memoirs of the New York
Botanical Garden 37: 1–116.
——— 1992. A synopsis of Rhynchospora (Cyperaceae) in Mesoamerica. Brittonia 44: 14–44.
Timonen, T. 1998. Inflorescence structure in the sedge tribe Cariceae (Cyperaceae). Publications in Botany
from the University of Helsinki 26.
Troll, W. 1964. Die Infloreszenzen, Typologie und Stellung im Aufbau des Vegetationskörpers 1. Gustav
Fischer. Jena.
——— & F. Weberling. 1989 . Infloreszenzuntersuchungen an monotelen Familien. G. Fischer, Stuttgart.
Tucker, G. C. 1983. The taxonomy of Cyperus (Cyperaceae) in Costa Rica and Panama. Systematic
Botany Monographs 2: 1–85.
——— 1987. The genera of Cyperaceae in the SE United States. Journal of the Arnold Arboretum
68: 361–445.
——— 1994. Revision of the Mexican Species of Cyperus (Cyperaceae). Systematic Botany Monographs
43: 213.
Tucker, C. S. & J. Grimes. 1999. The inflorescence: introduction. Botanical Review 65, 303–316.
Vegetti, A. 1992. Typology of the inflorescence in species Schoenoplectus (Cyperaceae) in Austral
America. Beiträge zur Biologie der Pflanzen 67: 241–249.
——— 1994. Typology of the Inflorescence in Species of Isolepis. Beiträge zur Biologie der Pflanzen 68:
21–26.
——— 2002. Typological reinterpretation of the inflorescences in Cariceae (Cyperaceae). Phyton 42:
159–168.
——— 2003. Sinflorescence typology in Cyperaceae. Annales Botanici Fennici 40: 35–46.
——— & J. C. Tivano. 1991. Sinflorescence in Schizachyrium microstachyum (Poaceae). Beiträge zur
Biologie der Pflanzen 66: 165–178.
——— & A. Anton. 1995. Some Trends of Evolution in the Inflorescence of Poaceae. Flora 190: 225–228.
——— & E. R. Guaglianone. 2005. Estructura de las Inflorescencias en especies de Rhynchospora
(Cyperaceae) de América Austral. XXX Jornadas Argentinas de Botánica. Pp. 48.
Vollbrecht, E., P. S. Springer, L. Goh, E. S. Buckler & R. Martienssen. 2005. Architecture of floral
branch systems in maize and related grasses. Nature 436: 1119–1126.
Vrijdaghs, A. 2006. A floral ontogenetic approach to homology questions in non-mapanioid Cyperaceae.
PhD thesis, Laboratory of Plant Systematics, K.U. Leuven, Belgium. 472 pp.
204 A. Reutemann et al.

———, P. Goetghebeur, E. Smets & P. Caris. 2003. The Schoenus L. spikelet a rhipidium? A floral
ontogenetic approach. In: Bayer, C., S. Dressler, J. Schneider & G. Zizka (eds.) 16th International
Symposium Biodiversity and Evolutionary Biology, 17th International Senckenberg Conference.
Abstracts: 24. Palmarum Hortus Francofurtensis 7, Frankfurt am Main.
———, ———, A. M. Muasya, E. Smets & P. Caris. 2004. The nature of the perianth in Fuirena
(Cyperaceae). South African Journal of Botany 70: 587–559.
———, P. Caris, P. Goetghebeur & E. Smets. 2005a. Floral Ontogeny in Scirpus, Eriophorum and
Dulichium (Cyperaceae), with special reference to the perianth. Annals of Botany 95: 1199–1209.
———, P. Goetghebeur, A. M. Muasya, P. Caris & E. Smets. 2005b. Floral ontogeny in Ficinia Schrad.
and Isolepis R. Br. (Cyperaceae), with focus on the nature and origin of the gynophore. Annals of
Botany 96: 1247–1264.
———, ——— & E. Smets. 2005c. Homology problems in cyperoid flowers: a floral ontogenetic
approach. In: XVII International Botanical Congress, Abstracts. Vienna, Austria, 219.
———, ———, ——— & P. Caris 2007. The Schoenus spikelet a rhipidium? A floral ontogenetic
answer. Aliso 23: 204–209.
———, A. M. Muasya, P. Goetghebeur, P. Caris, A. Nagels & E. Smets. 2009. A floral ontogenetic
approach questions of homology within the Cyperoideae (Cyperaceae). Botanical Review 75: 30–51.
———, M. Reynders, I. Larridon, A. M. Muasya, E. Smets & P. Goetghebeur. 2010. Spikelet structure
and development in Cyperoideae (Cyperaceae): a monopodial general model based on ontogenetic
evidence. Annals of Botany 105: 555–571.
Weberling, F. 1989. Structure and Evolutionary tendencies of Inflorescences in the Leguminosae. In: C. H.
Stirton & J. L. Zarucchi (eds.). Advances in Legume Biology. Monographs in Systematic Botany from
the Missouri Botanical Garden 29: 35–58.
Wilson, K. L. 1991 Systematic studies in Cyperus section Pinnati (Cyperaceae). Telopea 4: 361–496.
Zhang, S. 2001. A preliminary revision of the supraespecific classification of Kobresia Willd.
(Cyperaceae). Botanical Journal of the Linnean Society 135: 289–294.
Zhang, X., A. Marchant, K. L. Wilson, & J. J. Bruhl. 2004a. Phylogenetic relationships of Carpha and
its relatives (Schoeneae: Cyperaceae) inferred from chloroplast trnL intron and trnL-trnF intergenic
spacer sequences. Molecular Phylogenetics and Evolution 31: 647–657.
———, K. L. Wilson & J. J. Bruhl. 2004b. Sympodial structure of spikelets in the tribe Schoeneae
(Cyperaceae). American Journal of Botany 91: 24–36.