The significance of Cyperaceae as weeds

Abstract

Weedy Cyperaceae adversely affect natural plant communities and the health of humans and livestock and are major deterrents to agricultural and forest productivity. Most weeds are exogenous and have traits that give them biological and reproductive advantages over other plants. Weeds cost $ billions in agriculture, forestry, and urban areas and threaten diversity in natural communities worldwide. Of an estimated 8,000 species of weeds worldwide, only about 200 species cause approximately 95% of the problems in production of food, feed, fiber, and livestock. About 25% of the world’s weeds are monocots. Of these, sedges are among the most troublesome and difficult to control. The most important cyperaceous weeds in terms of their adverse effect on agriculture include Cyperus rotundus L., C. esculentus L., C. difformis L., C. iria L., and the Fimbristylis miliacea (L.) Vahl/F. dichotoma (L.) Vahl complex, ranking 1st, 16th, 32nd, 33rd, and 40th among the world’s worst weeds, respectively. We provide an overview of cyperaceous weeds, including economic losses, population dynamics, control methods, identification, biology, ecology, dispersal mechanisms, spread, and discussions of major weeds of agriculture, forestry, urban areas, and natural communities.

Full text

The Significance
of Cyperaceae as Weeds
Charles T. Bryson and Richard Carter
15
Chapter 2
ABSTRACT WeedyCyperaceae adversely affect natural plant communities and the health of humans and
livestock and are major deterrents to agricultural and forest productivity. Most weeds are exogenous and have
traits that give them biological and reproductive advantages over other plants. Weeds cost billions of dollars in
agriculture, forestry, and urban areas and threaten diversity in natural communities worldwide. Of an
estimated 8000 species of weeds worldwide, only about 200 species cause approximately 95% of the problems
in production of food,feed, fiber, and livestock. About 25% of the world’s weeds are monocots. Of these, sedges
are among the most troublesome and difficult to control. The most important cyperaceous weeds in terms of their
adverse effect on agriculture include Cyperus rotundus L., C. esculentus L., C. difformis L., C. iria L., and the
Fimbristylis miliacea (L.) Vahl/F. dichotoma (L.) Vahl complex, ranking first, 16th, 32nd, 33rd, and 40th among
the world’s worst weeds, respectively. We provide an overview of cyperaceous weeds, including economic
losses, population dynamics, control methods, identification, biology, ecology, dispersal mechanisms, spread,
and discussions of major weeds of agriculture, forestry, urban areas, and natural communities.
KEY WORDS Abildgaardia,Bolboschoenus,Bulbostylis,Carex,Cladium,Courtoisina,Cyperaceae,Cyperus,
Eleocharis,Fimbristylis,Fuirena,Isolepis,Kyllinga,Lepidosperma,Lepironia,Lipocarpha,Mapania,Oxycaryum,
Rhynchospora,Schoenoplectus,Scirpodendron,Scirpus,Scleria, sedge, weed.

16 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Cyperaceae is a cosmopolitan family with ca.
5000 species and 100 genera (Ball et al.,
2002). Members of Cyperaceae, commonly
called sedges, are monocot flowering plants with
reduced, mostly wind-pollinated (anemophilous)
flowers. The inconspicuous flowers are organized
into spikelets, and the spikelets further arranged into
higher order spicate, paniculate, or umbellate inflo-
rescences. Flowers may be either perfect or imper-
fect, and when imperfect, plants are monoecious (or
rarely dioecious). Fruits are small single-seeded ach-
enes. Sedges are primarily grass-like herbs with lin-
ear leaves and parallel venation. Cyperaceae and
Poaceae have traditionally been treated as related
families (Cronquist, 1981). Recent cladistic analysis
using molecular and morphological data confirms a
closer relationship with Juncaceae, with the “sedge
clade” consisting of Cyperaceae, Juncaceae, and
Thurniaceae (Chase et al., 2000).
Many species of Cyperaceae are heliophytes,
adapted to open, sunny areas with reduced competi-
tion from taller shading trees and shrubs. Such habi-
tats are often dependent upon natural or artificial dis-
turbance. A variety of plants, including many sedges,
have intrinsic characteristics (e.g., high reproductive
output, rapid growth, vegetative proliferation,
extended seed dormancy) that promote population
expansion after disturbance and probably originally
evolved as colonizers of disturbed habitats (Baker,
1965, 1974; McNaughton & Wolf, 1973). In addition
to catastrophic disturbances, more subtle and contin-
ual natural processes provide open areas for colo-
nization by such species, e.g., exposed bars and
banks along streams and coasts (Baker, 1974).
Plants are often called weeds when they oppor-
tunistically colonize and occupy habitats artificially
disrupted and maintained by humans, e.g., agricul-
tural fields, lawns, and gardens (Baker, 1974). The
term “weed” is inherently anthropocentric and,
therefore, is fundamentally problematic when used
in science. Some definitions are entirely subjective
and consequently are of little use in science,e.g., “a
plant growing out of place” (James et al., 1991: 1) or
“a plant growing where it is not desired” (Buchholtz,
1967: 389), and others emphasize only the negative
effects of weeds on natural communities and ecosys-
tems (Zimdahl, 1995; Randall, 1997). Although the
latter are applicable to natural resource management
and basic ecology, they are too restrictive for broad-
er application to agriculture and other applied sci-
ences. Bryson (2003: 1571) defined a weed as “an
undesirable plant that adversely affects humans or
other organisms which humans deem desirable.”
Reducing further the anthropocentric emphasis and
incorporating elements applicable in both pure and
applied sciences, we propose the following defini-
tion: Weeds are plants that alter the structure of nat-
ural communities, interfere with the function of
ecosystems, or have negative effects on humans,
agriculture, or other societal interests.
Cronk and Fuller (1995) clearly distinguish
between invasive plants that invade natural areas and
weeds or ruderals that infest agricultural or other
highly disturbed, artificial habitats, and they provide
a system of ranking weeds and invasive plants. The
same characteristics that enable plants to colonize an
area during ecological succession can make them
invasive pests when they are introduced outside their
natural ranges or habitats. Invasive weeds alter
wildlife habitat by reducing quantity and quality of
food sources, nesting sites, and cover, by increasing
the frequency of fire and soil erosion, and by chang-
ing the natural dynamics of aquatic systems causing
flooding or desiccation. Contrastingly, in agriculture
the most important weeds are those that have the
greatest economic impact through reduction in crop
yield, interference, or reduced efficiency or quality
of harvest.
About 8000 species, or approximately 3% of the
total number of plant species worldwide, have been
documented as weeds (Holm et al., 1977). Of these,
about 200 species, less than 0.1% of the world’s
flora, account for approximately 95% of weed prob-
lems in agriculture (Holm et al., 1977, 1979, 1997).
Invasive weeds possess a variety of characteristics
enabling them to disperse rapidly into new areas and
outcompete crops or native or desirable non-native
vegetation for light, water, nutrients, and space
(Westbrooks, 1998). To varying degrees, many char-
acteristics contribute to the success and competitive-
ness of invasive weeds, and sedges share many of
these traits with other plants (Table 1). The number of
weeds reported in crops and nonagricultural areas is
increasing. Two decades ago the important weeds in
cotton (Gossypium spp.) worldwide slightly exceed-
ed 100 species (Holm et al., 1977; Cronk & Fuller,
1995). Because of changes in production and cultur-
al practices (especially reduced-tillage production

The Significance of Cyperaceae as Weeds 17
systems), chemical control methods, weed shifts,
adaptations of populations, evolution of herbicide-
resistant weeds, and use of transgenic herbicide-
resistant crops, the total number of important weeds
in worldwide cotton production may currently
exceed 200 species, as demonstrated by the total
number recorded in cotton alone within the U.S.A.
(Bryson et al., 1999). Natural barriers and restricted
migration routes have historically prevented many
plants from dispersing over great distances.
However, the current speed and ease of world trans-
portation by humans and cargo have increased the
rate and distance of dispersal of plants. Upon intro-
duction, if a species becomes naturalized, it may
remain near the point of introduction without
becoming a pest. In the case of invasive weeds, the
local population amplifies and disperses, expanding
the range. Unfortunately, newly introduced weeds
often are undetected until after their numbers and
ranges increase greatly. The period of time between
introduction and invasion is the “lag phase”
(Radosivich & Holt, 1984), the duration of which
depends on a number of factors, e.g., size of popula-
tion, dynamics of reproduction, and detection. The
lag phase may vary from a few to many years, and
facilitation of a naturalized population must occur
before it expands, which may be brought about by
new pathways for dispersal, introduction of new pol-
linators or dispersal vectors, environmental change
(e.g., disturbance), and local adaptation through nat-
ural selection (Cronk & Fuller, 1995). Heterosis
resulting from hybridization with related species
may also be a factor in facilitation (Carter, 1990;
Daehler & Strong, 1997).
Table 1. Characteristics of weeds. Adapted from Muenscher (1955), Baker (1965, 1974), Klingman et al. (1982),
Radosevich and Holt (1984), Stuckey and Barkley (1993), Rejmanek (1996), and Westbrooks (1998).
Copious production of small seeds
Early maturation
Extended seed dormancy and discontinuous germination
Germination and survival in a wide range of environments
Long life of propagules in soil or during dispersal
Profuse vegetative reproduction and fragmentation
Rapid growth
Short juvenile period
Self-compatible or if cross-pollinated then by wind or unspecialized floral visitors
Survival and the ability to produce seed under adverse environmental conditions
Seed size similar to associated crops or native plants
Structural modifications (e.g., thorns, prickles, spines, urticating hairs) that cause injury and repel animals
or herbivores
Structural modifications facilitating dispersal
High photosynthetic rate (C4photosynthesis)
Increased water-use efficiency (C4photosynthesis)
Production of toxic secondary compounds that deter herbivores
Production of phytotoxins to prohibit or suppress growth of other plants (allelopathy)
Ability to parasitize other plants
Accumulation of large food reserves in roots, rhizomes, or other plant structures
Alternate host for insect pests and pathogens of crops
Resistance to pathogens
Small inconspicuous flowers
Short- and long-range dispersal mechanisms
Tolerance of environmental and chemical extremes, including fire, herbicides, and soil disturbances

18 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
ECONOMICS
There is little doubt that weeds cause severe eco-
nomic losses, but placing an exact value on their
impact worldwide is difficult, especially in natural or
nonagricultural areas. In the U.S.A., economic loss
due to invasive species (plants, animals, and
pathogens) was estimated to be more than $138 bil-
lion per year (Westbrooks, 2001). Economic losses
result from interference or competition with crops and
forests and the costs of pest-control chemicals, fuel,
equipment, labor, cultural-control practices, and addi-
tional irrigation and fertilizer (Chandler et al., 1984;
Chandler & Cooke, 1992). Additional costs to human
and animal health (i.e., allergies and toxins) are more
difficult to estimate, but weeds, including sedges,
cause substantial indirect economic losses worldwide.
In the U.S.A., it is estimated that cotton yields
are reduced 8.5% by Cyperus L. weeds (Byrd,
1995a), a loss of about $40.5 million annually. The
two primary Cyperus weeds in cotton and other row
crops are C. esculentus L. (yellow nutsedge) and C.
rotundus L. (purple nutsedge). In Mississippi alone,
31.4% and 23.5% of cotton fields are infested with
C. esculentus and C. rotundus,respectively; howev-
er, population levels of C. rotundus were greater
(75.6 aerial shoots/m2) than those of C. esculentus
(21.8 aerial shoots/m2) (Byrd, 1995b). It is more dif-
ficult to estimate economic impact on nonagricultur-
al areas, especially natural and public-use areas
where losses are measured as reduction in tourism
and recreation. Placing monetary values on native
flora and fauna and wildlife habitat displaced or
degraded by invasive species or the loss of the aes-
thetic value of a natural area is subjective and prob-
lematic. Control of weeds for the preservation of bio-
logical diversity is labor intensive and expensive,
requiring manual labor where chemical methods
may jeopardize natural plant communities (Randall,
1996). Upon control or eradication of invasive
weeds, additional expense is incurred to prevent
recolonization and to reintroduce native or innocu-
ous nonindigenous niche replacements.
The importance of an agricultural weed is not
necessarily correlated with its abundance within a
crop but may depend on herbicide- and cultural-con-
trol regimes, soil type, climatic conditions, number
of viable propagules in the seedbank, or other factors
(McWhorter & Bryson, 1992). Some weeds may be
abundant and conspicuous in crops without interfer-
ing, e.g., winter annuals that germinate, emerge,
flower, and set seeds early enough so growth and
yield of summer crops are unaffected. High popula-
tion levels of Isolepis carinata Hook. & Arn. ex Torr.
often occur in reduced-tillage cotton and soybean
(Glycine max (L.) Merr.) in the southeastern U.S.A.
(Bryson & Hanks, 2001). Because I. carinata com-
pletes its life cycle and dies early in the growing
season, it does not adversely affect crop growth
and yield. In agriculture, weeds that are difficult to
control, compete with crops for light, nutrients,
water, and space (Radosevich & Holt, 1984), inter-
fere with crop harvest efficiency, or reduce quality of
seed and lint (McWhorter & Bryson, 1992; Bryson
et al., 1999) are the most important. Holm et al.
(1977, 1997) list the world’s most important agricul-
tural weeds. Lists of weeds maintained by organiza-
tions include the Weed Science Society of America’s
Composite List of Weeds (WSSA, 1989) and Bayer
AG’s Important Crops of the World and Their Weeds
(Bayer AG, 1992). Bayer AG (1992) is a more com-
prehensive worldwide list and includes more than
5000 scientific names of crops and weeds, while the
WSSA lists about 2000 weeds found exclusively in
the U.S.A. and Canada. Since the second edition of
Bayer AG (1992), rights to the five-digit “Bayer
codes” for weeds have been sold to the European
Plant Protection Organization.
The economic, ethnobotanical, and horticultural
importance of the family Cyperaceae is well docu-
mented (Simpson & Inglis, 2001). Many sedges are
used as foods, food additives, drinks, fibers, animal
poisons, and in the manufacturing of items including
paper, perfumes, medicines, mats, boats, clothing,
shoes, ropes, and roofing (Kükenthal, 1935–1936;
Zeven & Zhukovsky, 1975; Darby et al., 1977; Allan,
1978; Burkill, 1985; Negbi, 1992; Stephens, 1994;
Bryson et al., 1998; Simpson & Inglis, 2001).
Tubers, rhizomes, seed, and foliage of sedges are
important wildlife and domesticated animal feeds or
forage (Hermann, 1970; Miller & Miller, 1999; Abad
et al., 2000). Cyperaceae are also utilized for erosion
control, revegetation after natural disturbances, and
to amend and improve soil fertility (Tachholm &
Drar, 1950; Hermann, 1970; Burkill, 1985; Fagotto,
1987; Simpson & Inglis, 2001). Traits that make
sedge species useful for erosion control and soil sta-
bilization also make them weeds.

The Significance of Cyperaceae as Weeds 19
CONTROL METHODS
Control methods for weedy sedges are diverse.
Cultural methods of hand removal, hoeing, and draft
plowing are still used in much of the world to control
weeds including sedges (Shear, 1985). Mechanical
tillage, flame cultivation, mowing, chemical treat-
ments (herbicides and fumigants), cover crops (e.g.,
sweet potato [Ipomoea batatas (L.) Lam.]), and
shading with a crop or black plastic have proven to
be effective in controlling many sedge weeds of turf,
pasture, and vegetable and row crops (Patterson,
1982; Glaze, 1987; Bryson & Keeley, 1992;
Buchanan, 1992; Peterson & Harrison, 1995).
As shown by Bryson et al. (2003a) with Cyperus
entrerianus Boeckeler, mowing alone will not
effectively control certain perennial sedge weeds,
but it can prevent seed production if mowing
intervals are shorter than the time required to set
fertile achenes. Fumigants are usually applied on
small areas to sterilize the soil for vegetable crop
production. Herbicide treatments may vary depend-
ing on the susceptability of target species, crop
tolerance, and required timing of application
(McWhorter & Bryson, 1992). With each herbicide
developed, research is conducted to determine the
efficacy on weeds and the selectivity on crops (Holt
et al., 1962; Hauser, 1963a, b; Duble et al., 1968,
1970; Hamilton, 1971; Hardcastle & Wilkinson,
1971; Keeley & Thullen, 1971; Keeley et al., 1972;
Wills, 1972; Zandstra et al., 1974; Zandstra &
Nishimoto, 1977; Wills & McWhorter, 1988;
Grichar et al., 1992; Richburg et al., 1993, 1994;
Wilcut et al., 1994; Vencill et al., 1995; Bryson et al.,
2003b).
Effective methods of herbicide application
include pre-emergence broadcast and incorporated
(with tillage) applications to control unwanted
sedges that germinate from seed, rhizomes, and
tubers. Acceptable post-emergence treatments are
dependent on the herbicide selectivity. Nonselective
herbicides are applied in areas where nontarget
species are of little concern, while selective herbi-
cides are applied to control target sedges without
harming crops or other desirable plants. Application
technologies have been developed to spray or wipe
nonselective herbicides on target weeds with special
equipment (e.g., directed sprayers, hooded sprayers,
recirculating sprayers, foam applicators, shielded
sprayers, chemigation, control droplet applicators,
air-assist systems, pneumonic applicators, sensing
devices, electrically charged sprayers, and rope-wick
applicators) to reduce or eliminate damage to crops
(Burr & Warren, 1971, 1972; Wiese, 1986; Bryson
& Wills, 1991; Wills et al., 1991; Barrentine et al.,
1992; Bryson et al., 1992b, 1994a; Bryson & Hanks,
1993; Bryson, 1994, 1997). Directed sprayers and
hooded sprayers are widely used in the U.S.A. for
controlling Cyperus rotundus,C. esculentus,C. iria
L. (rice flatsedge), and other weeds in cotton and
soybean. Additives such as soaps and lightweight
paraffinic oils effectively enhance activity of
some herbicides (McWhorter, 1982; Bryson et al.,
1990; Jordan, 1996). The development of herbicide-
resistant, transgenic crops allows application of
herbicides such as glyphosate over-the-top without
damaging crops (Shaner & Lyon, 1980), while
effectively controlling weeds. However, selection for
herbicide-resistant sedges is a potential problem
with the persistent use of a single herbicide or
herbicide family (Dowler et al., 1974). Resistance to
bensulfuron in C. difformis L. (smallflower umbrella
sedge) populations is well documented in rice
production (Pappas-Fader et al., 1993, 1994; Hill
et al., 1994), and herbicide resistance is also known
in other species (LeBaron, 1991).
Various pesticides also kill herbivores, preda-
tors, or pathogens of weeds, thereby potentially
rendering weeds more competitive. For example,
when nematocides control nematodes harmful to rice
(Oryza sativa L.), they also kill nematodes attacking
weeds of rice (e.g., Echinochloa spp. and Cyperus
haspan L.) (Hollis, 1972).
Although several potential biological control
agents (insects and pathogens) have been evaluated
for controlling Cyperus esculentus,C. rotundus, and
other sedges, none has been effective in reducing
sedge populations outside controlled experiments
(Phatak et al., 1987). It is unlikely that any single
biological agent will provide total control of
nutsedges (Morales-Payan et al., 2005). High para-
sitism and predation by other insects and use of
pesticides that kill biocontrol agents are major
constraints preventing effective biological control
of sedges using insects in row crops (Frick, 1978).
Excessive development, production, and registration
costs, short shelf life,and ineffective delivery
systems are major obstacles to utilizing pathogens

20 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
for biological control of weeds (Boyette, 2000; Duke
& Boyette, 2001).
POPULATION DYNAMICS
Weed species and population levels differ
depending on land use, cropland preparation,
forestation, and disturbance in natural areas. In agri-
cultural systems, weed shifts occur primarily when
management practices or environmental conditions
change (McWhorter & Bryson, 1992; Murray et al.,
1992). A single natural occurrence (e.g., tornado,
hurricane, earthquake, fire, flood) or cultural- and
chemical-control practices in farming operations
may eliminate or reduce populations of one weed,
while enhancing the survival, growth, and reproduc-
tive potential of another. As an example, farmers in
the southeastern U.S.A. claimed that Sida spinosa L.
seed and some sedge weeds such as Cyperus escu-
lentus and C. rotundus arrived in containers of dini-
troanaline (DNA) herbicides. In actuality however,
DNA herbicides controlled annual grasses and small
seed broadleaf weeds and vacated a niche for other
weeds to invade areas previously not infested (Frans,
1969; Dowler et al., 1974). Weed shifts may also
occur when environmental factors are modified
through row spacing, irrigation, and crop rotation in
row crops or when irrigation and fertilization fre-
quency is increased on lawns, turf, and flowerbeds.
In row crops, many sedge weeds thrive on irrigated
soils and occur in higher population levels prior to
crop canopy closure. Canopy closure earlier in the
growing season shades weeds and prevents seed or
tuber germination; thus, it is an effective cultural
practice in controlling many weeds, including sedges
such as C. esculentus and C. rotundus (Bryson et al.,
1990, 2003b).
Weed shifts may also occur as weeds disperse
into new areas. Non-native weeds, such as Cyperus
rotundus,C. iria,C. difformis, and Kyllinga brevifo-
lia Rottb., are excellent examples of weeds that were
introduced into the U.S.A. more than a century ago
and spread (Appendix 1). Within the past 50 years,
sedge weeds such as C. entrerianus,C. sanguinolen-
tus Vahl, and C. eragrostis Lam. have become estab-
lished and spread rapidly in areas previously not
infested in the U.S.A. (Carter, 1990, 2005; Carter &
Bryson, 1996, 2000b; Bryson et al., 1998). Once
introduced into a new area, weeds may take several
years to become established before causing prob-
lems (the lag phase). Duration of the lag phase may
vary depending on factors such as the number of
seeds produced, presence of dispersal vectors, and
environmental conditions (Radosevich & Holt,
1984). Early detection and implementation of con-
trol strategies are important in effectively controlling
non-native invasive weeds soon after introduction or
while still in the lag phase.
Farmers, consultants, and landowners must be
ever observant of new weeds and changes within
populations of weeds. Weed shifts are inevitable
when land use is altered or disturbance occurs. For
instance, weed shifts occur in reduced-tillage pro-
duction systems or where cover crops are utilized
(Bryson & Hanks, 2001). Perennial sedges such as
Cyperus esculentus and C. rotundus and many other
perennial weeds regenerate from greater soil depths
than most annual weeds (Elmore, 1984; Elmore et
al., 1989). Likewise, seeds of many annual sedges
germinate on the soil surface following a rainfall
event without burial, e.g., C. sanguinolentus (Carter
& Bryson, 2000b). No-tillage or conservation crop
production systems tend to favor weeds that germi-
nate from shallow soil depths and perennial weeds.
Unless controlled, perennial weeds are an increased
problem in reduced-tillage production systems.
Difficult-to-control perennial sedges such as C. escu-
lentus,C. rotundus,and perennial Kyllinga Rottb.
species often require repetitive and integrated con-
trol methods (Bryson & Keeley, 1992; McWhorter &
Bryson, 1992; Bryson et al., 1999, 2003b).
In order to assess the impact of a particular
weed species effectively, researchers have devised a
method to determine the competitive potential of
weeds based on field interference studies in agricul-
tural and forest areas (Coble & Byrd, 1992; Reichard
& Hamilton, 1997). Interference is ranked among
weed species to develop a competitive index or rela-
tive competitive abilities table such as the one for
selected weeds in cotton created by Coble and Byrd
(1992). Such an index aids farmers, consultants, and
landowners in determining which species are the
most pernicious and helps establish thresholds for
the number of weeds that can be tolerated in a given
situation. Computerized models (e.g., Soybean Weed
Control [SWC] and Mississippi State University
Herbicide Application Decision Support System
[MSUHADSS]) have been developed to aid farmers

The Significance of Cyperaceae as Weeds 21
and consultants in making recommendations, which
take into account the weed-competitive index, herbi-
cide options and prices, application costs, crop vari-
ety (cultivar), row spacing, crop stage, expected
weed-free yield potential, expected selling price, soil
moisture, and species of weed, population size, and
density (Bryson, 2003). However, little research has
been conducted to evaluate the thresholds of weeds
in natural areas, where populations may far exceed
threshold levels before a problem is perceived.
IDENTIFICATION
Accurate identification is essential in detecting
the presence of weeds and developing the best man-
agement strategies for control (Palm et al., 1968;
Murray et al., 1992). Traditionally, weed scientists
have approached plant identification pragmatically
and have adopted simplified systems to be used
primarily by individuals with minimal training in
systematic botany (e.g., Fischer et al., 1978; Stuckey
et al., 1980; Elmore & Bryson, 1986–2001; DeFelice
& Bryson, 2004). Such weed identification systems,
usually very different from the dichotomous keys
commonly used in taxonomic treatments, group
weeds by similar susceptibility or resistance to
herbicides, effectiveness of cultural-control
practices, time of germination, and other factors
(Bryson, 2003). In the simplest systems, plants are
grouped into general categories (e.g., broadleaved
species, grasses, sedges, annuals, or perennials),
which is usually sufficient for making decisions
about application of broad spectrum and nonselec-
tive herbicides. However, the increasing use of more
selective herbicides and biological control agents
demands greater precision in identification, i.e.,
determination to specific or infraspecific rank.
Among sedges, susceptibility to herbicides is usual-
ly correlated with species; however, infraspecific
biotypes (e.g., Cyperus difformis) do rarely exhibit
differential resistance to herbicides (Pappas-Fader et
al., 1993, 1994; Hill et al., 1994). In the case of her-
bicide-resistant biotypes, visual identification is
impossible, necessitating the use of bioassays
(LeBaron, 1991). Currently, when herbicide resist-
ance is suspected, bioassays are used to determine if
the lack of control is due to herbicide resistance,
herbicide tolerance, environmental conditions, or
misapplication.
Weed scientists and researchers in agriculture
usually work with a relatively small subset of all pos-
sible plant species in their area, and the agricultural
weeds are usually well known. Thus, simplified sys-
tems for identification generally work well for most
common agricultural weeds. However, when new or
unexpected weeds are encountered, more traditional
taxonomic methods must be adopted (e.g., use of
floristic manuals or systematic treatments in primary
literature). Although it may be possible to identify
immature sterile specimens of well-known sedges like
Cyperus esculentus and C. rotundus, reliable identifi-
cation of most sedges to species requires mature fer-
tile specimens and oftentimes the assistance of taxo-
nomic experts. To ensure that the specimen receives
proper attention from a taxonomist, it should be pre-
pared using standard methods and should include
accurate geographical data (Carter, 2003). To avoid
overlooking newly introduced weeds, every effort
should be made to collect unfamiliar sedges and to
identify them accurately. If one is not able to make a
reliable identification, then the specimen with data
should be sent to a competent taxonomist for determi-
nation. Vigilance, prompt action, and cooperation
between plant systematists and weed scientists are
absolutely essential in detecting newly introduced
sedges and dealing effectively with emerging weed
problems. Early detection and rapid response with
effective control methods are essential for eradication
of non-native invasive weeds (Westbrooks, 1998).
FACTORS DETERMINING
COMPETITIVE ADVANTAGE
The general characteristics of weeds summa-
rized in Table 1 are found to varying degrees in many
groups of plants, including sedges. Although no sin-
gle species exhibits all features, it is presumed that
there is usually a direct relationship between the
number of these characteristics and the degree of
invasiveness of a weed (Radosevich & Holt, 1984;
Bryson & Carter,2004). Most sedges reproduce sex-
ually through the production of large numbers of
small achenes. Such small reproductive structures
are well suited to both short- and long-distance dis-
persal. Because of their small size, the achenes of
sedges are difficult to detect and are readily trans-
ported as contaminants of seeds of crop, lawn, and
forage plants. Cyperus difformis and C. iria are

22 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
major agricultural pests, particularly of rice (Holm et
al., 1977). They probably originated as weeds by
invading rice paddies in Asia, where they were sub-
ject to similar selective pressures as rice. Annual
habit, rapid growth, short generation time, high
fecundity, and tolerance of submergence of roots
have enabled C. difformis and C. iria to persist and
disperse as weeds of rice. Cyperus difformis com-
pletes its life cycle in just four to six weeks and can
go through several generations within a single sea-
son (Holm et al., 1977), and an individual plant can
produce as many as 50,000 achenes (Jacometti,
1912). These and other sedges are thought to have
become naturalized throughout rice-producing areas
around the world via dissemination of their achenes
as contaminants of rice seed (Bellue, 1932;
Muenscher, 1955; Kral, 1971).
Obviously, certain characteristics listed in Table
1 are more important as determinants of invasiveness
than others. Given the importance of dispersal during
the phases of introduction and spread of invasive
species (Cronk & Fuller, 1995), characteristics relat-
ing to fecundity and dispersal of seeds would be of
major importance, as would those providing the abil-
ity to spread vegetatively. According to Holm et al.
(1977), Cyperus rotundus is the most pestiferous
plant in the world. It reproduces and disperses pri-
marily from vegetative tubers, with many biotypes
rarely producing viable seeds (Wills, 1987). Cyperus
esculentus, also a major agricultural weed, shares
similar reproductive characteristics. Vegetative struc-
tures such as stolons, rhizomes, and tubers are
important in localized spreading of many perennial
sedges and may even be transported long distances
both naturally and artificially when fragmentation
occurs. Subterranean rhizomes, tubers, and corms
also enable perennation and survival of sedges dur-
ing adverse environmental conditions, e.g., cold tem-
peratures, drought, or fire. Further discussion of dis-
persal in Cyperaceae, including dissemination of
vegetative fragments and structural modifications
facilitating transport of achenes and other structures,
is included below in the Dispersal section.
The highly reduced and inconspicuous flowers
of most sedges generally go undetected until after
they produce seeds, which Muenscher (1955) cited
as characteristic of many weeds. Cyperaceae are
almost exclusively wind-pollinated (anemophilous).
However, entomophily (insect pollination) has been
documented to varying degrees in Hypolytrum Rich.,
Mapania Aubl., Ascolepis Nees ex Steud.,
Rhynchospora Vahl sect. Dichromena (Michx.)
Griseb., Cymophyllus Mack., and even some species
of Bolboschoenus (Asch.) Palla, Carex L., Cyperus,
and Eleocharis R. Br. (Thomas, 1984a, b;
Goetghebeur, 1998). Although there is a paucity of
information, it is suspected that most sedges are
cross-pollinated (allogamous). For example, Cyperus
esculentus is self-incompatible, and therefore an
obligate outcrosser (Brown & Marshall, 1981) with
greater genetic variability within sexually reproduc-
ing populations than C. rotundus, which rarely pro-
duces viable seed (Horak & Holt, 1986; Horak et al.,
1987). Cross-pollination in combination with
anemophily is thought to contribute to the success of
weeds (Baker, 1965, 1974). Some of the most pestif-
erous sedges are very broadly ranging, exhibiting
great infraspecific diversity with many biotypes
adapted to a wide variety of environmental condi-
tions. Cyperus rotundus and C. esculentus are cos-
mopolitan weeds distributed widely throughout the
tropics and throughout much of the temperate zone
(Kükenthal, 1935–1936). In a worldwide treatment,
Kükenthal (1935–1936) recognized numerous infra-
specific taxa within these species, indicating consid-
erable adaptation to local environmental conditions.
C4photosynthesis confers a competitive advan-
tage under conditions of high temperature, high light
intensity, and water stress (Hopkins & Hüner, 2004).
C4plants have a lower transpiration ratio, thus, a
higher water-use efficiency, than C3species, brought
about by a lower CO2compensation point, reduced
photorespiration, and enzymes (ribulose-1,5-bisphos-
phate carboxylase [RUBISCO], phosphoenolpyruvate
carboxylase [PEPcase]) with higher optimal tempera-
tures (Hopkins & Hüner, 2004). In Cyperaceae, C4
photosynthesis is complex, consisting of four differ-
ent anatomical types (chlorocyperoid,rhyncho-
sporoid, fimbristyloid, eleocharoid) and two distinct
carbon assimilation modes (Brown, 1975; Soros
& Bruhl, 2000). In cladistic analyses using
developmental, anatomical, and molecular data,
Soros and Bruhl (2000) concluded that C4photosyn-
thesis arose multiple times (at least four) in the
Cyperaceae. Table 2 shows the occurrence of C4
photosynthesis in the genera of cyperaceous weeds.
In most cases genera are either C3or C4; however,
five genera, Abildgaardia Vahl, Cyperus,Eleocharis,

The Significance of Cyperaceae as Weeds 23
Mapanioideae (13/140)
Hypolytreae (9/130)
Mapania (70) C3
Scirpodendron (2) C3
Chrysitricheae (4/13)
Lepironia (1) C3
Cyperoideae (71/2380)
Scirpeae (6/60)
Scirpus (20) C3
Fuireneae (5/90)
Fuirena (30) C3
Bolboschoenus (11) C3
Schoenoplectus (50) C3
Actinoscirpus (1) C3
Eleocharideae (3/200)
Eleocharis (200) C3[C4]
Abildgaardieae (6/420)
Abildgaardia (10) C4[C3]
Fimbristylis (300) C4[C3]
Bulbostylis (100) C4
Cypereae (19/900)
Cyperus (incl. Anosporum,Juncellus,
Mariscus,Torulinium) (550) C4[C3]
Kyllinga (60) C4
Queenslandiella (1) C4
Pycreus (100) C4
Lipocarpha (35) C4
Oxycaryum (1) C3
Isolepis (60) C3
Courtoisina (2) C3
Schoeneae (29/700)
Rhynchospora (250) C3[C4]
Cladium (4) C3
Lepidosperma (55) C3
Sclerioideae (15/340)
Sclerieae (1/250)
Scleria (250) C3
Caricoideae (5/2150)
Cariceae (5/2150)
Carex (2000) C3
1Data on photosynthetic pathway from Bruhl (1993,1995) and Soros and Bruhl (2000); C3[C4] = mostly C3,C4[C3] = mostly C4.
2Subfamily and tribal classification and numbers of genera and species in parentheses are from Goetghebeur (1998).
3Authority names for genera in Table 2 not discussed elsewhere in this paper are as follows: Actinoscirpus (Ohwi)
R. W. Haines & Lye; Cyperus sect. Anosporum (Nees) Pax.
Table 2. The occurrence of C3and C4photosynthesis by genus of cyperaceous weeds.1, 2, 3
Fimbristylis Vahl, and Rhynchospora have both C3
and C4species. Of these, the mostly aquatic to sub-
aquatic Eleocharis is almost entirely C3, and all of the
subgenera of Cyperus are C4except Pycnostachys C.
B. Clarke [= Protocyperus]. Although many weeds are
not, some of the most competitive are characterized by
C4photosynthesis (Black et al., 1969; Elmore & Paul,
1983). Holm et al. (1977) rank C. rotundus,C. escu-
lentus,C. difformis, and C. iria among the world’s
worst weeds. Cyperus rotundus,C. esculentus,and
C. iria are C4plants; C. difformis is C3(Hesla et al.,
1982). Because C4photosynthesis is only one of
many factors contributing to the competitiveness of
weeds (Baskin & Baskin, 1978), it is not surprising
that other characteristics enable certain C3
Cyperaceae to be highly competitive weeds. C4pho-
tosynthesis is normally most advantageous in the ter-
restrial environment under conditions of drought,
high light, and high temperatures (Hopkins & Hüner,
2004). Cyperus difformis is almost exclusively a pest
of rice and is well adapted to aquatic environments,
where excessive water in the environment amelio-
rates high temperatures, and water stress is normally
not a factor. Thus, it is not surprising that C. difformis
has C3photosynthesis. Similarly, the C3species C.
haspan is a major weed of rice agriculture.
Although data on photosynthetic pathways for
most species of cyperaceous weeds are lacking,
generic-level data for species listed in Appendix 2
indicate a predominance of weeds in genera that are
exclusively or primarily C4(Fig. 1). Thus, it appears
that C4photosynthesis has been a major factor in the
success of genera such as Cyperus,Fimbristylis,
Kyllinga, and Bulbostylis DC. as weeds.
Certain plants, including weeds, achieve a com-
petitive advantage through allelopathy, the production
of chemical compounds that suppresses seed germina-
tion and growth in competing plants. Allelopathy is

24 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
well known in Cyperus rotundus and C. esculentus
and has been cited as a factor in its competition with
cotton and other crops (Friedman & Horowitz, 1971;
Mallik & Tesfai, 1988; Martinez-Diaz, 1997).
Although it has not been investigated, the nearly
monotypic nature of invasive populations of C.
entrerianus, observed in southern Louisiana and
eastern Texas, U.S.A. (Carter, 1990; Carter &
Bryson, 1996), suggests an allelopathic effect.
Weeds may also harbor insects and pathogens
that adversely affect agricultural crops and native
plants (USDA, 1960; Tietz, 1972). Cyperus dives
Delile, the natural host for a moth (Eldana sacchari-
na Walker) whose larvae cause losses to the sugar
industry, is of some concern as a weed in southern
Africa, where it is native and where an increase in its
frequency is associated with clearing of natural veg-
etation for the cultivation of sugarcane (Gordon-
Gray, 1995). Cyperus papyrus L. is also thought to
harbor this same moth (Gordon-Gray, 1995).
Noctuiid moth larvae of Spodoptera frugiperda (J. E.
Smith) [= Laphygma frugierda (Abbott & Smith)]
reportedly feed on C. rotundus,Carex spp., barley
(Hordeum vulgare L.), cotton, milo (Sorghum bicol-
or (L.) Moench.), potato (Solanum tuberosum L.),
rice, soybean, sweet potato, and other crops and
native plant species (Tietz, 1972). Colletotrichum
graminicola (Ces.) G. W. Wils., a fungal pathogen,
infects Carex spp., other Cyperaceae, and grass
crops (USDA, 1960). Cyperus esculentus,C. rotun-
dus, chili peppers (Capsicum annuum L.), and other
crops are hosts to the southern root-knot nematode
(Meloidogyne incognita (Kofoid & White) Chitwood)
(Schroeder et al., 1993).
DISPERSAL
Dispersal is fundamentally important in deter-
mining distributional patterns of plant species.
Dispersal may be complex and dynamic involving
both sexual and asexual systems, multiple vectors,
and shifts in vectors. When released from competi-
tion, predation, and disease, many species, upon
introduction outside their natural ranges, have poten-
tial to become weeds. Dispersal is crucial at two
points during invasion by plants: first, during the ini-
tial introduction of the species and later, after natu-
ralization, as the invasive species spreads, expanding
its range (Cronk & Fuller, 1995). Consequently,
basic knowledge about attributes of reproduction and
natural dispersal can provide insight into which
species are likely to become invasive weeds and how
they might be dispersed.
Vegetative growth from rhizomes, stolons,
runners, tubers, and corms is common in many
perennial sedges and is undoubtedly very important
in local expansion. Some species, e.g., Eleocharis
melanocarpa Torr., E. microcarpa Torr., and E. rostel-
lata Torr., have arching aerial stems that take root
apically upon contact with the ground, and others,
like E. vivipara Link, proliferate vegetatively from
spikelets. Cyperus pectinatus Vahl forms plantlets
vegetatively from its inflorescence (Haines & Lye,
1983). Vegetative growth when coupled with frag-
mentation and transport of asexual propagules can
also result in more distant dispersal. This is perhaps
most effective in the dispersal of fragments broken
from rafts (sudds) of floating or submerged natant
aquatic sedges by water currents or wind. Such dis-
persal has been noted in C. cephalotes Vahl, C.
colymbetes Kotschy & Peyr., C. mundtii Kunth, C.
papyrus,C. pectinatus, and Oxycaryum cubense
(Poepp. & Kunth) Palla (Kern, 1974; Haines & Lye,
1983; Gordon-Gray, 1995). We have noted this phe-
nomenon in C. alopecuroides Rottb., C. prolifer
Lam., Eleocharis baldwinii (Torr.) Chapm., E. vivip-
ara, and O. cubense and suspect that it occurs in
other species similar in habit and habitat, e.g., C. ela-
tus L. and Websteria confervoides (Poir.) S. S.
Hooper (Kern, 1974).
Sedges exhibit a variety of modifications
exploiting various agents of dispersal, most of which
directly involve fruits or inflorescences. A number of
mechanisms involving dispersal of achenes by wind
(anemochory) are known in Cyperaceae. In
Afrotrilepis (Gilly) J. Raynal, Carpha Banks & Sol.
ex R. Br., Costularia C. B. Clarke, Eriophorum L.,
and Scirpus L., a persistent perianth adnate to the
achene is modified into long, silky bristles or hairs
that facilitate transport by wind (Kern, 1974; Pijl,
1982; Haines & Lye, 1983; Goetghebeur, 1998), and
in Androtrichum Brongn. and Machaerina Vahl, per-
sistent elongated filaments have the same function
(Goetghebeur, 1998). Also, the flattened wing-like
floral scales of Anosporum spp. and the flattened
winged spikelets of certain Kyllingaspp. (Haines &
Lye,1983) promote wind dispersal of the achenes
retained within. Such dispersal of spikelets has been

The Significance of Cyperaceae as Weeds 25
observed over short distances during collection of
specimens of the introduced weed K. squamulata
Thonn. ex Vahl (Carter, pers. obs.).
Dispersal by water (hydrochory) is well docu-
mented in Cyperaceae. The fruits or spikelets of
most terrestrial sedges are disseminated to some
extent by rain; however, such dispersal is usually
quite local (Ridley, 1930). The achenes of the aquat-
ic and wetland sedges Cyperus (Anosporum) colym-
betes,C. pectinatus,C. platystylis R. Br., Oxycaryum
cubense,several Scirpus spp., and certain wetland
Carexspp. have a spongy suberized pericarp that
facilitates flotation and dispersal by moving water
(Chermezon, 1924; Ridley, 1930; Kern, 1974; Lye,
1981; Haines & Lye, 1983). Achenes of Cladium P.
Browne were observed to float in the laboratory for
up to 15 months (Ridley, 1930). Similarly in Cyperus
odoratus L. and Remirea maritima Aubl., the achene
remains enclasped in a buoyant corky rachilla and is
thereby dispersed by moving water (Kern, 1974;
Haines & Lye, 1983). Floods undoubtedly transport
even unmodified, nonbuoyant achenes, vegetative
fragments of plants (e.g., rhizomes, tubers), and
whole plants (Kern, 1974) and deposit them far from
the main channel along basins of major rivers.
Cyperus fuscus L., a potential rice weed in the
U.S.A., has apparently been dispersed by floodwa-
ters along the Missouri River in the central U.S.A.
(McKenzie et al., 1998).
Dispersal of achenes by animals (zoochory),
especially birds, is important in Cyperaceae.
Zoochory may involve the internal (endozoic) trans-
port of achenes within the digestive system or exter-
nal (epizoic) transport. The achenes of Carex,
Cladium,Cyperus,Fimbristylis,Rhynchospora, and
Scirpus have been identified in the alimentary sys-
tems of waterfowl (Ridley, 1930). Waterfowl and
other birds consume large quantities of achenes,
Figure 1. Photosynthetic pathways among genera of Cyperaceae with weeds; data on photosynthetic pathways from
Bruhl (1993, 1995) and Soros and Bruhl (2000).

26 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
especially of Cyperus spp. and Eleocharis spp., and
their endozoic transport plays an important role in
dispersal of sedges over long and short distances
(Ridley, 1930; Kern, 1974; Haines & Lye, 1983).
Vlaming and Proctor (1968) experimentally deter-
mined that sedge achenes remained viable after
retention in avian digestive systems for periods up to
120 hours: Cyperus ochraceus Vahl, max. 37 hr.;
Eleocharis albida Torr., max. 38 hr.; E. macrostachya
Britton, max. 77 hr.; E. parvula (Roem. & Schult.)
Link ex Bluff, Nees & Schauer max. 30 hr.; and E.
quadrangulata (Michx.) Roem. & Schult., max. 120
hr. Brightly colored fruits in the tropical genus
Gahnia J. R. Forst. & G. Forst. are consumed and
dispersed by birds (Benl, 1937; Pijl, 1982; Lye,
2000), and, according to Sauer (1988), seeds of Carex
nigra (L.) Reichard were brought to Iceland by snow
buntings from Great Britain. Short-distance endozoic
dispersal by cattle (Carex,Scirpus) and water buffalo
(Fimbristylis globulosa (Retz.) Kunth, F. littoralis
Gaudich.) has been reported by Kern (1974).
Similarly, the epizoic transport of achenes in
mud adhering to the feet of migratory waterfowl is
implicated in long-distance dispersal in Cyperus,
Eleocharis,Rhynchospora, and Scirpus (Ridley,
1930; Kern, 1974). Such mechanisms could account
in part for the wide distributions of C. drummondii
Torr. & Hook., C. odoratus,C. virens Michx., and
Oxycaryum cubense. A number of epizoic mecha-
nisms involving various structural modifications are
known in Cyperaceae. The achenes of many species
of Eleocharis,Fuirena Rottb., Rhynchospora,
Schoenoplectus (Rchb.) Palla, and Websteria S. H.
Wright are subtended by persistent, hypogynous
bristles beset with retrorse barbs that readily attach
to feathers or hair of animals (Kern, 1974; Haines &
Lye, 1983), and the North American sedge, C.
plukenetii Fernald, exhibits a number of modifica-
tions that facilitate dispersal of intact spikelets by
attachment to animal hair (Carter, 1993). Uncinia
Pers., widely distributed in the Southern Hemisphere,
including many islands of the Pacific, is characterized
by a hooked inflorescence axis that extends beyond
the utricle, attaching readily to feathers and enabling
transport by birds (Pijl, 1982; Mabberley, 1997).
Carex pauciflora Lightf. has a springing mechanism
that disperses its perigynia over relatively short dis-
tances when touched by animals (Hutton, 1976), and
the perigynia of certain other Carex spp. produce oil-
rich appendages and are dispersed by ants (Handel,
1976, 1978; Gaddy, 1986). Similarly, a fleshy peri-
anth in Lepidosperma Labill. reportedly facilitates
dispersal by ants (Goetghebeur, 1998).
High fecundity and small fruits (achenes) make
sedges especially susceptible to unintentional dissem-
ination directly by humans or through their activities.
A variety of human activities are known or suspected
to disperse sedges, and most of these involve move-
ment of their small, inconspicuous achenes. Sedge
achenes are readily dispersed as contaminants of com-
mercial seed supplies (Koyama, 1985; Bryson &
Carter, 1992; Sell & Murrell, 1996), and achenes or
even live plants may contaminate ornamental nursery
stock, potted plants, or mulch. A number of sedges
associated with rice agriculture around the world (cf.
Bolboschoenus,Cyperus,Eleocharis,Fimbristylis,
Schoenoplectus in Appendix 1) are thought to have
dispersed via achenes as contaminants of rice seed
(Bellue,1932; Muenscher,1955; Kral, 1971).
Shipments of shorn wool may contain achenes of
sedges, which when dispersed result in the introduc-
tion of so-called “wool aliens” (Sell & Murrell, 1996).
Other kinds of cargo, including live animals, trans-
ported by land, sea, or air may harbor achenes result-
ing in the unintentional introduction of sedges (Carter
& Mears, 2000). Dumping of ballast contaminated
with achenes or vegetative propagules (e.g., rhizomes,
stolons, tubers) has long been associated with disper-
sal of sedges and other plants (e.g., Smith, 1867;
Brown, 1880; Britton, 1886; Mohr, 1901). The inad-
vertent transport of achenes or vegetative propagules
embedded in mud or lubricants adhering to wheels or
other parts of freight cars, trucks, automobiles, and
airplanes undoubtedly disperses sedges, and migra-
tion of plants, including sedges, along railroads (fer-
roviatic migration) is well documented (e.g.,
Mühlenbach, 1979, 1983). It also seems likely that
tiny achenes of sedges, drawn by jet airplane engines,
could lodge in the housing of the engine or other parts
and be carried great distances. The transport of turf-
grass sod, mulch, soil, hay, and fodder has been asso-
ciated with dispersal of sedges, e.g., Cyperus esculen-
tus,C. rotundus,Kyllinga brevifolia, and K. gracilli-
ma Miq. (Bryson et al., 1992b, 1996, 1997; Sell &
Murrell, 1996), and movement of achenes and vegeta-
tive propagules occurs during construction and main-
tenance of roads, e.g., Cyperus entrerianus,C. san-
guinolentus,Carex oklahomensis Mack., and

The Significance of Cyperaceae as Weeds 27
C. praegracilis W. Boott (Kern, 1974; Reznicek
& Catling, 1987; Carter, 1990; Carter & Bryson,
1996, 2000b).
Because sedges are generally inconspicuous,
and other than as weeds are of minimal economic
importance, they escape all but casual notice and
interest of most humans; consequently, it is pre-
sumed that the intentional dispersal of sedges by
humans is infrequent. However, as shown in
Appendix 1 and in Figures 2 and 3, there is an
increased interest in using sedges as ornamentals,
and a surprising number of species are subject to
deliberate transfer by humans. Some of these have
become naturalized weeds from cultivation, and any
could potentially become pests. Carex comans
Berggr., C. morrowii Boott, C. pendula Huds.,
C. riparia Curtis, Cymophyllus fraserianus (Ker
Gawl.) Kartesz & Gandhi, Cyperus compressus L.,
C. eragrostis,C. longus L., C. owanii Boeckeler, and
C. strigosus L. are used in gardens, and Carex bac-
cans Nees, Cyperus albostriatus Schrad., C. fertilis
Boeckeler, and Isolepis cernua (Vahl) Roem. &
Schult. are sometimes used as potted plants or in
hanging baskets (Bailey, 1935, 1949; Bailey &
Bailey, 1976; Everett, 1980–1982; Brickell & Zuk,
1997). Of these, Carex riparia,Cyperus compressus,
C. eragrostis, and C. longus are listed as weeds (cf.
Appendix 2), and the South African C. owanii is
naturalized,but apparently not invasive, in
California, U.S.A. (Tucker et al., 2002).
Cyperus alternifolius L. subsp. flabelliformis
Kük. (umbrella plant, umbrella sedge) has been used
as an ornamental in water gardens and as a potted
plant for more than 200 years (Bailey & Bailey,
1976). It is widely naturalized from cultivation
throughout the tropics and subtropics (Kern, 1974;
Koyama, 1985) and is frequently cited as a weed
(cf. Appendix 2). Other sedges cultivated in water
gardens include C. papyrus (papyrus), C. prolifer
(dwarf papyrus, miniature papyrus), C. sexangularis
Nees, C. textilis Thunb., and various bulrushes,
Bolboschoenus robustus (Pursh) Soják, Schoeno-
plectus acutus (Muhl. ex J. M. Bigelow) Á. Löve &
D. Löve, S. heterochaetus (Chase) Soják, S. taber-
naemontani (C. C. Gmel.) Palla, S. lacustris (L.)
Palla, and Scirpus cyperinus (L.) Kunth (Bailey,
1935, 1949; Everett, 1980–1982; Gordon-Gray,
1995). Cyperus papyrus is naturalized in Australia
(Wilson, 1993) and in Florida, U.S.A. (Wunderlin,
1998), and C. prolifer is naturalized in Florida
(Carter et al., 1996).
Appendix 1 is a list of sedges known or suspect-
ed to be transported by human activities. The fre-
quencies of various modes of anthropogenic disper-
sal in Cyperaceae are shown in Figure 2: ornamen-
tals (53%); wool aliens (13%); ballast (7%); rice
agriculture (7%); revegetation, reclamation, erosion
control, and soil improvement (7%); and highways
and railroads (3%). The burgeoning human popula-
tion and the current ease and frequency of rapid
long-distance transportation of humans and cargo
make it inevitable that such dispersal of sedges, both
unintentional and deliberate, will continue into the
foreseeable future.
INVASION BY CYPERUS
ENTRERIANUS:A CASE STUDY
The following case study of Cyperus entreri-
anus (deeprooted sedge), based upon Carter (1990)
and subsequent investigation (Carter, unpubl. data),
shows how basic research in the field and the herbar-
ium allows for the detection of invasive weeds and
illustrates the need for accurate and timely identifi-
cation in order to take appropriate action against
them. In 1987, Carter found a species of Cyperus in
Ware County, Georgia, that did not fit any descrip-
tions of species known from the southeastern U.S.A.
During 1988 and 1989, intensive searching in the
field resulted in discovery of numerous additional
populations of this perplexing sedge in Florida,
Georgia, Alabama, Louisiana, and eastern Texas.
During this same period, an examination of herbari-
um specimens at FSU, IBE, and VDB revealed addi-
tional ones, variously misidentified, that were col-
lected from northern Florida in the 1970s and 1980s,
southern Louisiana in 1975, and eastern Texas in
1981. In early 1989, Carter correctly determined that
the enigmatic sedge was C. entrerianus.
Based upon data gleaned from herbarium speci-
mens and intensive field research and Rosen et al.
(2006), the following hypothetical scenario for the
introduction, naturalization, and dispersal of Cyperus
entrerianus in the U.S.A. is proposed (Fig. 4). Cyperus
entrerianus was introduced into the U.S.A. before
1941, and the suspected points of introduction are
Cameron County, Texas, and Pensacola, Florida
(Brinker 413,US). The species was not found again in

28 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
the U.S.A. until 1974, when it was collected again in
Pensacola (Godfrey 73755, FSU). It was collected in
southern Louisiana in 1975 (Allen 6674, VDB), addi-
tional collections were made in Escambia and Gulf
counties, Florida, during the late 1970s and 1980s, and
it was found in eastern Texas in 1981 (Carter, 1990).
All of the collections of C. entrerianus made by others
before it was reported new to the U.S.A. by Carter
(1990) were variously misidentified as C. pseudovege-
tus Steud., C. robustus Kunth, C. virens,and C. virens
var. drummondii (Torr. & Hook.) Kük. The paucity of
herbarium records before the mid-1980s suggests that
C. entrerianus was in its lag phase until then. Although
the apparent rapid expansion of range in the late 1980s
and 1990s is undoubtedly in part an artifact of inten-
sive searching for C. entrerianus by Carter and others
(Carter, 1990; Carter & Jones, 1991; Bryson & Carter,
1994; Carter & Bryson, 1996), its collection at a num-
ber of sites in Louisiana and Florida during the later
1970s by researchers who had no knowledge of its cor-
rect identity indicates that its lag phase had ended
some years earlier.
It is suspected that Cyperus entrerianus was
introduced independently in southern Texas and at
Pensacola from temperate South America or Mexico
(Carter, 1990; Rosen et al., 2006). There are other
cases of introduced Cyperus weeds that were proba-
bly imported into Pensacola via ballast: C. aggrega-
tus (Willd.) Endl., C. difformis,C. pilosus Vahl, and
C. reflexus Vahl (Burkhalter, 1985; Wunderlin,
1998); thus, introduction of C. entrerianus via bal-
last is plausible. Distribution and habitat indicate
that C. entrerianus has spread from its point of intro-
duction at Pensacola via dispersal from road con-
struction and maintenance activities, primarily along
highway Interstate 10 and secondarily along inter-
secting highways (Carter,1990; Carter & Bryson,
1996). It is probably also now dispersed endozoical-
ly by birds or other animals that consume its ach-
enes. Certain populations of C. entrerianus in the
southeastern U.S.A. show evidence of introgression
with C. surinamensis Rottb., which could account in
part for the robust habit (heterosis) observed in
plants there (Carter, 1990). Vigorous growth and
robust form have probably facilitated the rapid
expansion of C. entrerianus in the southeastern
Figure 2. Percentages of various kinds of anthropogenic dispersal of cyperaceous weeds listed in Appendix 1.

The Significance of Cyperaceae as Weeds 29
U.S.A. from Florida and southern Georgia west into
eastern Texas, and it has begun to invade natural
areas in eastern Texas (Rosen et al., 2006).
SURVEY OF GENERA
AND SELECTED SPECIES
There is no comprehensive, contemporary, cos-
mopolitan enumeration and description of species of
Cyperaceae, and such comprehensive accounts of
most cyperaceous genera do not exist. Furthermore,
there is still considerable disagreement about taxo-
nomic limits and circumscriptions of many genera.
Consequently, estimates of numbers of taxa (gen-
era/species) vary considerably: ca. 70/ca. 4000
(Cronquist, 1981); 98/4350 (Mabberley, 1997); 104
genera/5000+ (Goetghebeur, 1998); ca. 100/ca. 5000
(Ball et al., 2002). For example, there is little con-
sensus about the circumscription of Cyperus,i.e.,
whether it should be defined broadly to include
Diclidium Schrad. ex Nees, Juncellus C. B. Clarke,
Kyllinga,Mariscus Vahl, Pycreus P. Beauv., and
Queenslandiella Domin with infrageneric rank, or
whether it should be defined narrowly with the seg-
regates treated as genera. This problem has major
implications with respect to nomenclature in
Cyperus,the most important genus of weeds in the
family (Carter & Bryson, 2000a). Use of molecular
techniques (e.g., Muasya et al., 2000a, b) should help
to stabilize nomenclature by resolving the taxonom-
ic status and rank. However, until such basic prob-
lems are resolved through additional research and
alignment of nomenclature, we think a conservative
approach is warranted. Herein where possible,
nomenclature at the generic level follows the recent-
ly published Flora of North America, volume 23.
However, in the absence of a synonym under
Cyperus, one species, Pycreus decumbens T.
Koyama, reported as a weed in Brazil by Kissmann
(1997), was not listed in Appendix 2. Cyperus
decumbens Govind., the name for a different species
from India published in 1973, prevents legitimate
transfer of the name under Cyperus.
Based upon a survey of more than 60 publica-
tions, Appendix 2 is a worldwide list that includes
449 species of Cyperaceae that have been cited as
weeds. Additionally, we have included other sedges
indigenous to the southeastern U.S.A. that we have
observed to be weeds. Table 3 summarizes numbers
of weedy species by genus. Cyperus is by far the
Figure 3. Cumulative numbers of ornamental and cultivated species of Cyperaceae listed in selected horticultural
references from 1935 to 2001.

30 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Rhynchospora,Kyllinga,Bulbostylis,Fuirena,
Scirpus, and Bolboschoenus had fewer than 5% each,
and the remaining 10 genera had fewer than 1% each.
Cypereae, which includes Cyperus, is the largest
tribe of weeds (Fig. 5), and subfamily Cyperoideae,
which includes Cypereae, has the overwhelming
majority of weedy sedges (Fig. 6).
The previous lists of Holm et al. (1977, 1979,
1997) and WSSA (1989) show a substantially larger
proportion of weeds in Cyperus (ca. 42%); ca. 43% in
Eleocharis,Fimbristylis,Scirpus (incl. Bolboschoenus,
Isolepis R. Br., Schoenoplectus); and the remaining
15% in Carex,Cladium,Fuirena,Kyllinga,Rhyn-
chospora (incl. Dichromena,Psilocarya Torr.), and
Scleria. Our survey (Appendix 2) shows a much small-
er proportion in Cyperus and substantial increases in
Carex and other genera. Bayer AG (1992) was not used
in compiling Appendix 2 because it does not
separate weeds from crops and because it is based upon
key sources cited in Appendix 2.
ABILDGAARDIA
Abildgaardia is a genus of ca. 15 species distrib-
uted mostly in the pantropics and subtropics in both
the Eastern and Western hemispheres (Kral, 2002d).
Although Abildgaardia spp. have been placed in
Bulbostylis and Fimbristylis, embryological and
anatomical data support segregation as a separate
genus (Lye, 1973). The results of our survey
(Appendix 2) show only one species, A. ovata (Burm.
f.) Kral, cited as a weed, which is reported to be a
weed in Asia, North America, and the Pacific Islands
(Holm et al., 1979). In southern Florida, U.S.A., it is
occasionally a weed of gravelly soils in waste areas,
along highways, and in lawns (Carter, pers. obs.).
BOLBOSCHOENUS
Bolboschoenus is a genus of 6 to 15 species
(Smith, 2002a), five of which are listed as weeds in
Appendix 2. Bolboschoenus maritimus (L.) Palla, con-
sidered among the world’s worst weeds, is a pest in
agricultural lands and waterways in Africa, Asia,
Australia, Europe, and North and South America
(Holm et al., 1997; Kissmann, 1997). It is a trouble-
some rice weed in paddy fields (Holm et al., 1977,
1997), and in the southern Korean peninsula B. mar-
itimus infests more than 80% of rice fields reducing
yields by as much as 50% when adequate control
Figure 4. The dispersal of Cyperus entrerianus
Boeckeler in the U.S.A. —A. 1941–1979. —B.
1941–1989. —C. 1941–1999. —D. 1941–2003.
largest genus with more than 147 species or 33% of
the total, followed by Carex with 82 species and
18%, Eleocharis with 53 species and 12%,
Fimbristylis with 46 species and 10%, and Scleria P.
J. Bergius with 24 species and 5%. Schoenoplectus,

The Significance of Cyperaceae as Weeds 31
measures are not taken (Ryang et al., 1978). Integrated
weed management schemes, including rotation of
crops, water regimes, and chemical and cultural meth-
ods, effectively control B. maritimus in rice-producing
areas of Asia (De Datta & Jereza, 1976; Verga et al.,
1977). Bolboschoenus maritimus is less a problem in
the equatorial zone than in semitropical and temperate
regions of the world (Holm et al., 1997). The achenes
Table 3. Numbers and percentages of cyperaceous
weeds by genus (data extracted from Appendix 2).
Species Percent
Genus (infrasp.) of Total
Cyperus1147 (2) 33
Carex 82 18
Eleocharis 53 12
Fimbristylis 46 10
Scleria 24 5
Rhynchospora 20 4
Schoenoplectus 20 4
Kyllinga 13 3
Bulbostylis 92
Fuirena 8 2
Scirpus 82
Bolboschoenus 51
Lipocarpha 4< 1
Cladium 2< 1
Abildgaardia 1< 1
Courtoisina 1< 1
Isolepis 1< 1
Lepidosperma 1< 1
Lepironia 1< 1
Mapania 1< 1
Oxycaryum 1< 1
Scirpodendron 1< 1
Total 449 (2) 100
1Includes Diclidium, Juncellus, Mariscus, Pycreus, and
Queenslandiella.
of B. maritimus are readily dispersed by birds (Holm
et al., 1997) and by water (Guppy, 1893). Bolbo-
schoenus fluviatilis (Torr.) Soják is reported as an
aquatic weed in Asia, Australia, and North America
(Holm et al., 1979; WSSA, 1989).
BULBOSTYLIS
Bulbostylis is a genus of ca. 100 species, occur-
ring mostly in dry or periodically dry, sunny, sandy
uplands and savannas in warm temperate and tropical
regions worldwide (Kral, 2002c). Nine species are
listed as weeds in Appendix 2; however, none is a
major weed. Bulbostylis barbata (Rottb.) C. B.
Clarke and B. capillaris (L.) C. B. Clarke are occa-
sionally weeds of sandy soil in flowerbeds and poor-
ly managed turf in the southeastern U.S.A. In late
summer and fall in the Coastal Plain of the southeast-
ern U.S.A., B. barbata can be a conspicuous feature
of the landscape when en masse its reddish brown
inflorescences appear in sandy cultivated fields
(Kral, 1971). Bulbostylis capillaris and B. ciliatifolia
(Elliott) Fernald are common weeds of sandy fallow
fields, roadsides, and on gravel and cinders of rail-
road right-of-ways (Kral, 1971; Godfrey & Wooten,
1979). All three species often grow in sandy soil in
flowerbeds and lawns or through cracks in sidewalks
and parking lots. Bulbostylis barbata is reported as a
weed of cultivated lands in Taiwan (Lin, 1968), and
B. capillaris is reported as a weed in Brazil
(Kissmann, 1997).
CAREX
Of the more than 2000 species worldwide (Ball
& Reznicek, 2002), only a small proportion of Carex
spp. are major weeds when compared to other sedge
groups (e.g., Cyperus,Kyllinga). Although not
among the most troublesome weeds of rice, Carex
diandra Schrank and C. pycnostachya Kar. & Kir.
are reported from rice field habitats in Pakistan
(Kukkonen, 2001). Very few Carex spp. are invasive,
and none is a principal agricultural weed (Holm et
al., 1977), which may be due to several factors
including more restrictive habitat requirements,
fewer or larger seeds, shorter period of sexual repro-
duction, fewer vectors for dispersal, lack of tolerance
to mowing or tillage, and greater susceptibility to
herbicides. In Appendix 2, 82 species of Carex are
listed as weeds.

32 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Carex blanda Dewey and C. leavenworthii
Dewey are occasional weeds of poorly kept lawns,
especially under shade of deciduous trees in the south-
eastern U.S.A. (Bryson, 1985a). Carex blanda is often
locally abundant and capable of being weedy in
diverse environmental and edaphic conditions (Bryson
& Naczi, 2002). It is highly likely that C. blanda and
other weedy Carex species are dispersed as contami-
nates of grass seed, sod, or clippings for turf establish-
ment (Jones et al., 1993). In lawns and on golf cours-
es, C. blanda and C. leavenworthii are controlled by
frequent mowing and herbicide treatments (Bryson,
1985a). Listing of C. cephalophora Muhl. ex Willd. as
a weed (Callahan et al., 1995) may be due to taxonom-
ic confusion with C. leavenworthii and literature that
considered the two taxa conspecific. Another occa-
sional lawn weed, C. cherokeensis Schwein., is a weed
of pastures (Burns & Buchanan, 1967; Burns et al.,
1969; Bryson, 1985a). In the Black Prairie and Lower
Coastal Plains regions of Alabama, C. cherokeensis is
reported to displace desirable forage species in poor
quality pastures (Burns & Buchanan, 1967; Burns et
al., 1969). Carex cherokeensis persists and spreads in
the early spring or late fall by extensive rhizomes
when many pasture grasses are dormant. It is more
prevalent in poorly managed pastures lacking herbi-
cide applications, and mowing alone is not effective in
C. cherokeensis control.
Carex longii Mack. is weedy along roadsides
and in lawns and flowerbeds (Bryson, 1985a). Unlike
most Carex,C. longii flowers and fruits throughout
the frost-free months. Frequently in the southeastern
U.S.A., establishment of this species occurs follow-
ing dispersal of pine bark mulch around shrubs and
in flowerbeds suggesting contamination by C. longii
seeds. From flowerbeds, C. longii can invade sur-
rounding areas; however, it is not as aggressive as
several invasive Cyperus and Kyllinga spp. in lawns,
turf,gardens, and row crops (Bryson, pers. obs.).
Listing of C. albolutescens Schwein. as a weed
(WSSA, 1989) may be due to taxonomic confusion
recently clarified by Rothrock (1991). Although C.
albolutescens may be locally common, it is not
weedy along roadways and in lawns, pastures, and
flowerbeds like C. longii.
Non-native Carex species have become invasive
weeds in natural areas through accidental introduc-
tion or escape from cultivation as ornamentals. On
sandy beaches and dunes, C. kobomugi Ohwi, native
to Japan, has become an invasive weed along Atlantic
coasts (Standley, 1983). It was first collected in the
U.S.A. in 1929 (Fernald, 1930), but at that time, it
was misidentified as the closely related species C.
macrocephala Willd. ex Spreng. Since 1929, C.
kobomugi has spread on sandy beaches from Rhode
Island southward to North Carolina and displaced
native vegetation and altered the structure of beaches
(Small, 1954; Svenson, 1979; Stalter, 1980; Standley,
1983). Its range is likely to expand (Mastrogiuseppe,
2002) despite current eradication efforts in several
states. Following introduction as an ornamental, C.
pendula has recently escaped into natural areas and is
beginning to appear on roadsides and stream banks;
however, its potential as an invasive weed is
unknown (Reznicek, 2002).
Some Carex species native to one region of a
continent have become weedy in other regions of the
same continent. Northern and eastern records of C.
oklahomensis are most recent, and this sedge may be
increasing its range (Standley, 2002). Because C.
oklahomensis has been frequently collected from
recently completed construction sites, it is probably
dispersed in hay mulch used for erosion control
along roadsides, lakesides, and ditch banks (Bryson
et al., 1992a, 1996). Carex opaca (F. J. Herm.) P.
Rothr. & Reznicek appears to be similarly dispersed
(Bryson et al., 1994b).
Seeds and rhizomes of Carex praegracilis are
dispersed along highways by traffic and by construc-
tion and maintenance equipment, and it is sometimes
called “tollway sedge” or “freeway sedge” (Swink &
Wilhelm, 1979; Bruton & Catling, 1982). Carex
praegracilis is adapted to extreme environmental
conditions (e.g., salty or dry roadsides) and is spread-
ing rapidly eastward and southward from its native
range, especially along roadsides where salt is
applied for deicing (Reznicek et al., 1976; Bruton &
Catling,1982; Cusick, 1984; Reznicek & Catling,
1987, 2002).
Carex nebrascensis Dewey is listed as a weed
(WSSA, 1989; Callahan et al., 1995); it was appar-
ently introduced into Missouri and Illinois, U.S.A.,
and has become weedy along roadsides (Standley et
al., 2002). Heavy infestations of C. lanuginosa
Michx. were effectively controlled by herbicides, and
tillage provided better control of this sedge in light
(sandy) soils than in heavier (silt or clay) soils in
New Mexico, U.S.A. (Hollingsworth, 1969).

The Significance of Cyperaceae as Weeds 33
Carex aquatilis Wahlenb., C. atherodes Spreng.,
C. glaucescens Elliott, C. frankii Kunth, C. lacustris
Willd., C. lasiocarpa Ehrh., C. louisianica L. H.
Bailey, C. pallescens L., C. rostrata Stokes in With.,
C. senta Boott, and C. verrucosa Muhl. are listed as
weeds by the WSSA (WSSA, 1989), while C. lupuli-
na Muhl. ex Willd. is listed as a weed by WSSA
(1989) and Callahan et al. (1995). Carex comosa
Boott is considered weedy by Callahan et al. (1995).
The U.S. Fish and Wildlife Service (USFWS) (1988)
lists C. comosa as an obligate wetland species.
According to Bernard and Seischab (1994), C.
comosa invades gaps in wetlands and persists for up to
a decade while producing seeds that are dispersed into
new gaps. Treated in Flora of North America as dis-
tinct from C. frankii,C. aureolensis Steud. (Ford &
Reznicek, 2002) is weedy in the southeastern U.S.A.
in pastures and along wet roadsides and agricultural
field borders. Carex heterostachyaBunge and C.
rigescens (Franch.) V. Krecz. are reported as weeds in
China along roadsides and field borders or in orchards
and nursery gardens (Zhirong et al., 1990).
CLADIUM
There are four species of Cladium worldwide
with three in North America (Tucker, 2002a), of
which two, C. jamaicense Crantz and C. mariscoides
(Muhl.) Torr., are cited as weeds (Holm et al., 1979;
WSSA, 1989). Both of these wetland species occur
in the U.S.A. Cladium jamaicense (saw grass) inhab-
its marshes near the coast and is the predominant
species of the Everglades marshes of southern
Florida, U.S.A. (Steward & Ornes, 1975; Godfrey &
Wooten, 1979). Much of this formerly vast marsh-
land has been drained for flood control and convert-
ed into agricultural fields for the cultivation of sug-
arcane and other crops (Godfrey & Wooten, 1979).
In such an altered and unnatural landscape, C.
jamaicense is viewed as an impediment to drainage
and navigation and a hindrance to agriculture.
Figure 5. Number of cyperaceous weeds by tribe; classification follows Goetghebeur (1998).

34 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
However, a massive venture is currently underway
(U.S. Army Corps of Engineers, 1999) to reverse the
damage done by drainage projects of the past and to
reclaim portions of the Everglades ecosystem, which
if successful will also restore the natural habitat of C.
jamaicense, taking it from weed to its former status
as the predominant plant of its natural community.
Weediness is oftentimes an artifact of human percep-
tion and folly.
COURTOISINA
Courtoisina Soják is a small genus of two
species found in Africa, Madagascar, India, and
southeastern Asia (Haines & Lye, 1983; Gordon-
Gray,1995; Vorster, 1996; Mabberley, 1997).
Courtoisina cyperoides (Roxb.) Soják was cited as a
weed in rice fields (Simpson & Koyama, 1998;
Simpson & Inglis, 2001) and has also been reported
from wet mud of freshwater pans, seasonally wet
grasslands, and temporary pools (Haines & Lye,
1983; Gordon-Gray, 1995).
CYPERUS
There are about 600 species of Cyperus world-
wide (Tucker et al., 2002). In terms of their signifi-
cance as weeds, Cyperus species are by far the most
important in Cyperaceae. Appendix 2 lists 147
species of Cyperus that have been cited as weeds.
The adverse economic impact of Cyperus is great.
According to Holm et al. (1977), it contains the
world’s worst weed and three additional species
listed among the 33 worst agricultural weeds in
the world. The most recent comprehensive,
universal treatment of Cyperus was by Kükenthal
(1935–1936), who defined the genus broadly as con-
sisting of six subgenera: Cyperus,Mariscus (Vahl)
C. B. Clarke, Torulinium (Desv.) Kük., Juncellus (C.
B. Clarke) C. B. Clarke, Pycreus (P. Beauv.) A. Gray,
and Kyllinga. Cyperus is taxonomically complex,
and the status of its subgenera is widely disputed
even among contemporary workers (cf. Kern, 1974;
Haines & Lye, 1983; Koyama, 1985; Lye, 1992;
Wilson, 1993; Adams, 1994; Gordon-Gray, 1995;
Figure 6. Number of cyperaceous weeds by subfamily; classification follows Goetghebeur (1998).

The Significance of Cyperaceae as Weeds 35
Muasya et al., 2000a, b, 2002; Tucker et al., 2002).
Although we question the apparent inconsistency in
segregating Kyllinga and not Pycreus and Juncellus,
herein we pragmatically adopt the generic taxonomy
in Flora of North America, Vol. 23 (Tucker et al.,
2002). Recent molecular evidence seems to support
a broad circumscription of Cyperus to include
Kyllinga,Pycreus, and other segregate genera
(Muasya, 2002).
To varying degrees, the following characteristics
undoubtedly contribute to the aggressive, invasive
tendencies of Cyperus spp. and other sedges: large
numbers of small, readily dispersed achenes; vegeta-
tive reproduction; longevity of tubers, rhizomes, or
other subterranean structures; production of allelo-
pathic compounds; paucity of pathogens; short life
reproductive cycle, especially in annual species; tol-
erance of broad ranges of environmental conditions;
C4photosynthesis; and resistance to control with her-
bicides and cultural methods, including tillage.
Cyperus rotundus is considered the world’s
worst weed because of its ability to survive, spread,
and compete, especially in agricultural areas (Holm
et al., 1977; Terry, 2001). It was reported in 52 crops
and 92 countries (Holm et al., 1977, 1979, 1997). In
the U.S.A., Elliott (1821) described C. rotundus (C.
hydra Michx.) as a “scourge” of plantations in
Georgia and South Carolina and recommended daily
tilling of the soil for control. The infraspecific taxon-
omy of this cosmopolitan weed is extremely com-
plex and in need of revision (cf. Kükenthal,
1935–1936). In addition to threatening agriculture,
C. rotundus is a troublesome weed in urban areas
and natural communities after disturbance. Although
it rarely sets viable seeds (Holm et al., 1977; Thullen
& Keeley, 1979), C. rotundus produces numerous
rhizomes that reportedly can penetrate and grow
through fleshy subterranean organs of root crops and
even asphalt pavement (Hauser, 1962a, b; Thullen &
Keeley, 1979). These rhizomes form tubers that give
rise to new aerial plants or produce other rhizomes or
they may remain dormant during periods of adverse
environmental conditions including heat, cold,
drought, flooding,or inadequate aeration (Ranade &
Burns, 1925; Williams, 1978; Bendixen &
Nandihalli, 1987; Wills, 1987; Miles et al., 1996).
The tubers of C. rotundus are bitter, rough, and are
often connected serially by rhizomes with or without
giving rise to new plants (Plowman, 1906; Ranade &
Burns, 1925; Hauser, 1962a; Wills & Briscoe, 1970;
Holm et al., 1977; Wills, 1987). Dormant tubers
make C. rotundus difficult to control in turf, and only
a few selective herbicides that effectively control
sedges are approved for use in turf or in row crops
(Aleixo & Valio, 1976; Keeley, 1987; Pereira et al.,
1987; Holt & Orcutt, 1996). Tubers and rhizomes of
C. rotundus produce allelopathic compounds that
reduce growth in crops such as cotton (Martinez-
Diaz, 1997).
Diagnostic features of Cyperus rotundus
include abruptly tapering leaves, inflorescence
bracts equaling or longer than the inflorescence, and
purplish floral scales (Wills & Briscoe, 1970;
Horowitz, 1972; Wills, 1987). In a comparative
study of C. rotundus morphology based on collec-
tions from 13 states in the U.S.A. and 21 locations
from around the world, Wills (1998) detected differ-
ences in numbers of shoots produced by single
tubers, numbers of leaves per shoot, lengths and
widths of leaves, lengths of culms, flowering times
during the year, numbers and lengths of rachises,
lengths of rachillae and spikelets, and numbers,
lengths, and widths of involucral bracts. Infraspecific
variation in C. rotundus is also documented in
Ceylon (Koyama, 1985), East Africa (Haines & Lye,
1983), and Natal Province (now KwaZulu-Natal
Province), South Africa (Gordon-Gray, 1995).
Although these differences occurred within some
traits on a worldwide basis, the basic characteristics
distinguishing C. rotundus from other taxa were con-
sistent (Wills, 1998) and differed from closely relat-
ed taxa such as C. bifax C. B. Clarke. Worldwide, C.
rotundus is a troublesome weed in corn (Zea mays
L.), cotton, peanut (Arachis hypogaea L.), rice,
sorghum (Sorghum vulgare Pers.), soybean, sugar-
cane (Saccharum officinarum L.), turf grass species,
and many other vegetable, nursery, row, rotation, and
plantation crops (Long et al., 1962; Bryson, 1985b;
Bendixen & Nandihalli, 1987; Holt & Orcutt, 1991;
Derr & Wilcut, 1993; Grichar et al., 1992; Wills,
1998; Bryson et al., 2002, 2003b).
Cyperus esculentus is ranked as the world’s
16th worst weed (Holm et al., 1977). Highly variable
and widely distributed in tropical, subtropical, and
temperate regions around the world, its infraspecific
taxonomy was revised by Schippers et al. (1995).
Schippers et al. (1993) attribute invasiveness in C.
esculentus to an increase in the rate of population

36 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
growth brought about by tillage. Cyperus esculentus
has rhizomes and tubers (Thumbleson &
Kommedahl, 1961; Jansen, 1971; Stoller et al.,
1972); however, its rhizomes are fleshy and termi-
nate in a sweet-tasting tuber (Garg et al., 1967).
Additional diagnostic characters include gradually
tapering, acute leaves, yellow to yellowish orange
floral scales, and bracts longer than the inflores-
cence. Cyperus esculentus is pernicious and difficult
to control in agricultural and urban areas. Although
it produces seeds more frequently than C. rotundus
(Wills, 1987), C. esculentus reproduces primarily
from tubers (Thumbleson & Kommedahl, 1961).
Cyperus esculentus tubers remain dormant for pro-
longed periods during adverse environmental condi-
tions and only produce tubers from rhizomes of the
parent plant (Wills, 1987). Cyperus esculentus is
able to survive colder winter conditions than C.
rotundus and thus is more widespread worldwide
(Stoller & Sweet, 1987; Wills, 1987). The tubers of
C. esculentus are called chufas, tiger nuts, or rush
nuts (Abad et al., 2000; DeFelice, 2002). Cyperus
esculentus is often planted for its tubers that provide
food for deer, turkey, wild hogs, and other animals
(Miller & Miller, 1999; Abad et al., 2000; DeFelice,
2002). Humans also use the tubers as food for
domesticated animals (e.g., chickens, swine) and
directly consume them as food, use them as a spice,
and use them to prepare a drink called “horchata de
chufas” (Zeven & Zhukovsky, 1975; Darby et al.,
1977; Allan, 1978; DeFilipps, 1980c; Negbi, 1992;
Stephens, 1994; Bryson et al., 1998). Unfortunately,
the tubers used by humans contribute to the invasive
character of C. esculentus and to its dispersal. In
addition to the crops mentioned above for C. rotun-
dus,C. esculentus is also a principal weed of potato
(Solanum tuberosum), sugarbeet (Beta vulgaris L.),
and many cool-season crops (Bendixen &
Nandihalli, 1987).
Cyperus difformis and C. iria are ranked 32nd
and 33rd among the world’s worst weeds, respective-
ly (Holm et al., 1977). Both are caespitose annuals
and often produce clumps of many culms and have
become established in tropical and temperate areas
of the world. In the southeastern U.S.A., C. difformis
and C. iria are primarily weeds of drainage ditches,
rice fields, and poorly drained sites in other agricul-
tural fields or disturbed areas. Cyperus difformis and
C. iria produce multiple generations per year under
optimal growing conditions and in the tropics flower
and produce seeds year-round (Holm et al., 1977).
Cyperus difformis can complete its life cycle every
four to six weeks throughout the growing season
(Holm et al., 1977). A single plant of C. iria may
produce more than 5000 viable seeds, while an indi-
vidual of C. difformis can produce 50,000 seeds with
a germination rate of 60% or more (Jacometti, 1912).
Short generation times and high seed production
favor rapid dispersal (Vaillant, 1967), large seed
reservoirs in the soil, high population levels (Holm et
al., 1977; Bryson, 1984), and an increased potential
for the development of herbicide resistance. In rice-
production areas of California, multiple C. difformis
generations per year and large seed production may
be primary factors in the rapid development of herbi-
cide resistance to bensulfuron (Pappas-Fader et al.,
1993, 1994; Hill et al., 1994). Despite similarities in
habitat and growth and reproductive patterns, C. dif-
formis is C3and C. iria is C4(Hesla et al., 1982).
Cyperus rotundus,C. esculentus,C. difformis,
and C. iria are all suspected to have originated in
Asia. Other Cyperus spp. of probable Asian origin
include C. compressus,C. haspan,C. pilosus, and C.
sanguinolentus (Holm et al., 1979). All are natural-
ized weeds in other regions of the world (Bryson &
Carter, 1995; Carter & Bryson, 2000b).
Cyperus haspan is among the world’s worst
weeds (Holm et al., 1997). It has been reported as a
weed in 12 crops and 39 countries throughout tropi-
cal and semitropical areas of Africa, Asia, Australia,
South America, and North America (Lin, 1968;
Holm et al., 1977, 1979; Kissmann, 1997). Cyperus
haspan is a recent introduction into Hawaii with the
first collection made in 1957 (Wagner et al., 1990).
An individual plant can produce more than 50,000
achenes per year (Datta & Banerjee, 1976), and
although plants produce achenes during the first sea-
son of growth, they do not form rhizomes until the
second year (Tadulingam & Venkatanaryana, 1955).
Cyperus haspan,a C
3plant, commonly occurs in
shallow standing water and germinates and grows
well in wet, sandy, acidic soils (Bertels, 1957; Eyles
& Robertson, 1963; Dirven, 1970). Cyperus haspan
is sometimes broken into two subspecies; C. haspan
subsp. juncoides (Lam.) Kük. is a taller plant with
conspicuous rhizomes (Kükenthal, 1935–1936;
Kern, 1974; Koyama, 1985). Cyperus haspan is
sometimes confused with closely related C. tenuispi-

The Significance of Cyperaceae as Weeds 37
ca Steud., a species with more widely spaced floral
scales, and both species are cited as frequent weeds
in rice fields in Asia (Kern, 1974; Koyama, 1985).
Cyperus entrerianus is native to temperate
regions of South America; it is also known from the
Caribbean, Mexico, and the Coastal Plain of the
southern U.S.A. (Kükenthal, 1935–1936; Barros,
1960; Carter, 1990; Tucker, 1994). In his compre-
hensive revision of Cyperus, Kükenthal (1935–1936)
accorded specific rank to C. entrerianus; however,
Barros (1960) reduced it to varietal status under C.
luzulae (L.) Rottb. ex Retz., and Denton (1978) gave
it no rank, treating it as a synonym of C. luzulae.
Carter (1990) and subsequent authors (Tucker, 1994;
Tucker et al., 2002) treated this taxon at the rank of
species. In the southeastern U.S.A., flooding, con-
struction equipment, mowing, and soil-moving
activities, especially along highways, disperse the
small achenes of C. entrerianus, resulting in infesta-
tions in a variety of disturbed habitats (Carter, 1990;
Carter & Bryson, 1996). Cyperus entrerianus dis-
places native vegetation even in undisturbed habitats,
and, without widespread control, it will likely con-
tinue to spread rapidly, infesting agricultural, forest-
ed, riparian, and urban areas. Figure 4 shows the dis-
persal of C. entrerianus in the U.S.A., where by
2003 it was documented in 43 counties from Florida
and southern Georgia into southeastern Texas. In the
southern U.S.A., C. entrerianus reproduces copious-
ly from achenes and spreads vegetatively and peren-
nates from short rhizomes. Cyperus entrerianus is a
prolific seed producer, with the number of seeds per
inflorescence ranging from 1000–20,000+ depend-
ing on the size and maturity of plants and mature
plants (> 1 year old) producing 10–100+ inflores-
cences per year (Carter & Bryson, 1996; Bryson et
al., 2003a). Preliminary seed germination studies
indicate moderate to high viability (55%–95%)
(Carter & Bryson, 1996). In the southeastern U.S.A.,
C. entrerianus flowers and fruits from June until
frost in November or December (Carter, 1990;
Carter & Jones, 1991; Bryson & Carter, 1994).
Cyperus entrerianus continues to spread at an alarm-
ing rate and threatens agricultural and natural areas.
Also, preliminary studies suggest that populations
will potentially spread northward into Arkansas,
North Carolina, South Carolina, Tennessee, and
Virginia. Additional research is needed to determine
more effective methods of prevention and control.
Cyperus acuminatus Torr. & Hook., C. era-
grostis,C. luzulae,C. ochraceus,C. pseudovegetus,
C. reflexus,C. surinamensis, and C. virens are cited
as weeds (Appendix 2) and are classified with C.
entrerianus in Cyperus sect. Luzuloidei Kunth
(Kükenthal, 1935–1936; Denton, 1978). Cyperus
acuminatus,C. pseudovegetus, and C. virens are all
native to North America, where they are currently rel-
atively minor weeds; however, they could become
problems if introduced beyond their native ranges.
Cyperus pseudovegetus is widely distributed in east-
ern North America (Denton, 1978; Tucker et al.,
2002). In the U.S.A., C. pseudovegetus and C. virens
are common in disturbed, intermittently wet soils,
e.g., roadside ditches, margins of ponds, and swales
in fields, pastures, and grasslands. Cyperus virens is
widely distributed in the New World, ranging from
South America, Central America, the Caribbean
Islands, Mexico, and the southern U.S.A. (Denton,
1978), and is recently introduced into Hawaii with
the first collection made in 1976 (Wagner et al.,
1990). Several infraspecific taxa have been recog-
nized (Denton, 1978), including C. virens var. drum-
mondii. Carter et al. (1999) showed that C. drum-
mondii is specifically distinct from C. virens and in
the southeastern U.S.A. has a more restricted distri-
bution and habitat and is less weedy than C. virens.
Cyperus reflexus occurs in temperate South America,
Central America, Mexico, and in the U.S.A. (Denton,
1978; Tucker, 1994). It is introduced in Australia,
where it is naturalized near Sydney (Wilson, 1993).
In the U.S.A., C. reflexus is most common in south-
eastern Texas and Louisiana, where it is found in
intermittently wet, disturbed soils of ditches, fields,
and grasslands (Denton, 1978; Carter, pers. obs.); it
has also been reported in western Florida (Wunderlin,
1998). Additional research is needed to elucidate the
relationship between C. fraternus Kunth and C.
reflexus,which has been treated as C. reflexus var.
fraternus (Kunth) Kuntze (Kükenthal, 1935–1936;
Denton, 1978). Cyperus ochraceus is widespread in
the New World and is known from South America,
Central America, Mexico, the U.S.A., and the
Caribbean Islands (Denton, 1978). It is currently only
a minor weed in the southern U.S.A., where it is
found in disturbed, intermittently wet soils and is
most common in Texas and Louisiana but has dis-
persed to scattered sites elsewhere (Denton, 1978;
Tucker et al., 2002; Carter, pers. obs.).

38 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Cyperus surinamensis is widely distributed in
the New World, ranging from South America,
Central America, Mexico, and the Caribbean Islands
into southeastern and south-central U.S.A. (Denton,
1978). Readily identified by its retrorsely scabrid
culms, C. surinamensis has been cited as a weed in
both North and South America (WSSA, 1989;
Kissman, 1997). In warmer parts of the southeastern
U.S.A., it is a common weed in a variety of open dis-
turbed sites with hydric soils.
Cyperus eragrostis occurs naturally in South
America and in California, Oregon, Washington, and
British Columbia in North America (Denton, 1978).
It has been used ornamentally, which in part
accounts for its introduction into other parts of the
world (Tucker, 1987; Sell & Murrell, 1996; Darke,
1999; Petrík, 2003). It occurs sporadically in the
eastern U.S.A., where it is introduced and appears to
be spreading (Bryson & Carter, 1994; Bryson et al.,
1996; Tucker et al., 2002). Cyperus eragrostis is nat-
uralized in Australia and has expanded its range and
frequency there, where it is a weed of rice and
ephemerally wet, disturbed sites (Wilson, 1993). In
reporting C. eragrostis new to the Czech Republic,
Petrík (2003) provides a thorough account of its
invasion of Europe, where it is widely distributed
and has been known since the mid-1800s. Given its
association with rice as a weed in Australia, C. era-
grostis could become a problem in rice agriculture in
the southeastern U.S.A. and elsewhere. Additional
research is needed to determine more about the dis-
tribution and dispersal of C. eragrostis,its potential
to become an agricultural pest, and its control.
Cyperus oxylepis Nees ex Steud. and C. elegans
L. are widely distributed in tropical, subtropical, and
warm temperate regions of the New World. Both
species have viscid foliage and are markedly aromat-
ic, with the fragrance of cedar wood (Juniperus vir-
giniana L.) sometimes sensed in the field before the
plants are seen. The floral scales of C. oxylepis are
golden brown and those of C. elegans are greenish
tan. Cyperus oxylepis,listed as a weed (WSSA,
1989), is apparently expanding its range in coastal
areas of the southeastern U.S.A. (O’Neill, 1938b;
Thieret, 1964; Tucker, 1987; Bryson & Carter, 1992;
Bryson et al., 1996), where it is found in disturbed
clay soils of salt marshes.
A number of aquatic Cyperus species cultivated
in ponds and water gardens have become naturalized.
All have the potential to become invasive weeds in
aquatic and wetland habitats in tropical and subtropi-
cal areas, and at least one, C. prolifer, is invasive in the
U.S.A. (Carter et al., 1996). Trade and importation of
these species should be carefully regulated to prevent
further impact. Cyperus alternifolius subsp. flabelli-
formis has been used as an ornamental in water gar-
dens and as a potted plant for more than 200 years
(Bailey & Bailey, 1976) and is widely naturalized
from cultivation in the tropics and subtropics and
other warm areas (Bailey, 1935, 1949; O’Neill, 1946;
Kern, 1974; DeFilipps, 1980c; Koyama, 1985;
Wagner et al., 1990; Sell & Murrell, 1996). It has been
variously known as C. alternifolius subsp. flabelli-
formis Kük.; C. flabelliformis Rottb., nom. illeg.; and
C. involucratus Rottb. In the U.S.A. it is naturalized in
Florida, Louisiana, Texas, and California, where it is
occasionally found in moist to hydric soils of roadside
ditches, stream banks, vacant lots, and other disturbed
sites (Carter,pers. obs.; Tucker et al., 2002). In his
worldwide monograph of Cyperus, Kükenthal
(1935–1936) recognized two subspecies: C. alterni-
folius subsp. alternifolius and C. alternifolius subsp.
flabelliformis. Baijnath (1975) treated these as species
and stated that C. alternifolius is rare and mostly
restricted to Madagascar where it is native and that C.
involucratus [= C. alternifolius subsp. flabelliformis]
is the correct name for the widely naturalized cultivat-
ed plant indigenous to Africa. More recently, Gordon-
Gray (1995) adopted Kükenthal’s taxonomy, indicat-
ing the need for additional critical investigation of this
complex in southern Africa, which also includes the
related cultivated aquatics C. sexangularis and C. tex-
tilis. Until further research elucidates the relationships
among these taxa, we have adopted the more conser-
vative taxonomy of Kükenthal (1935–1936) and
Gordon-Gray (1995), recognizing two subspecies
within C. alternifolius. Although popular in water gar-
dens in southern Africa, C. sexangularis survives
under drier conditions in the absence of extended
water stress (Gordon-Gray, 1995), and C. textilis is
naturalized in the Azores (DeFilipps, 1980c). Thus, it
would appear that C. alternifolius,C. sexangularis,
and C. textilis have the potential to become invasive
pests in a variety of aquatic, wetland, and terrestrial
habitats in tropical and subtropical regions.
Cyperus prolifer is sold as an ornamental for
water gardens (Bailey & Bailey, 1976; Simpson,
1994) and has been variously listed as Cyperus has-

The Significance of Cyperaceae as Weeds 39
pan cv. ‘viviparus’ (Watkins & Sheehan, 1975; Graf,
1985), C. papyrus cv. ‘nanus’ (Bailey & Bailey,
1976), and C. isocladus Kunth (Bailey & Bailey,
1976; Everett, 1980–1982). It has been confused
with C. haspan, from which it is readily distin-
guished by its thick rhizome and inflorescence of 50
to 100 rays of more or less uniform length. Cyperus
prolifer is indigenous to eastern Africa where it
inhabits marshes, marshy shores, and swampy
stream banks (Kükenthal, 1935–1936; Haines &
Lye, 1983). Although Simpson (1994) stated that it
was not a weed, C. prolifer has become naturalized
from cultivation in the U.S.A. in central Florida
where it has invaded the margins of lakes (Carter et
al., 1996) and in Hawaii (Strong & Wagner, 1997). In
Florida, C. prolifer grows in floating mats and along
margins of natural limesink lakes, where it is associ-
ated with Oxycaryum cubense (Carter et al., 1996).
One extensive population of C. prolifer in Lake
Huntley, Florida, was established after dispersing
from an adjacent water garden during eight years of
cultivation (Carter et al., 1996).
Cyperus papyrus is a remarkable plant. Because
of its use in the manufacture of the first paper by the
ancient Egyptians, it is perhaps the best known of the
sedges (Schery, 1972). It is found in central and
southern Africa and the Nile River valley and is nat-
uralized in Sicily (Kükenthal, 1935–1936;
DeFilipps, 1980c; Gordon-Gray, 1995). Cyperus
papyrus forms dense stands in aquatic and wetland
habitats and dominates swamps with low biodiversi-
ty in northern Uganda (Mabberley, 1997). Plants
may grow to 5 m high, making it one of the largest
sedges (Koyama, 1985), and it is cultivated as an
ornamental and curiosity in greenhouses and out-
doors in ponds and water gardens in tropical and
subtropical regions of the world (Bailey, 1935, 1949;
Bailey & Bailey, 1976). Cyperus papyrus is natural-
ized in Florida, U.S.A. (Wunderlin, 1998), where it
is evidently not yet invasive, but would appear to
have the potential to invade aquatic and wetland
habitats in tropical and subtropical areas given its
dominance in swamps of northern Uganda.
Cyperus alopecuroides is a widely distributed
aquatic sedge in tropical and subtropical regions of
the Old World, e.g., northern and tropical Africa,
Madagascar,India, Ceylon, Indo-China, Malaysia,
and northern Australia; in the New World it is known
only from Guadeloupe in the West Indies and
Florida, U.S.A. (Kükenthal, 1935–1936; Koyama,
1985; Carter et al., 1996). It is a robust plant to 1.5
m high and in its habit and general inflorescence pat-
tern resembles the tropical species C. imbricatus
Retz. Both taxa were placed in section Exaltati
Benth. by Kükenthal (1935–1936). Its size in combi-
nation with other characters make C. imbricatus a
striking plant in the field: broad bracts and leaf
blades (to 15 mm wide) with contrasting surfaces
(adaxial light green, abaxial glaucous), and a
branched inflorescence with spikes of densely clus-
tered golden-brown spikelets (Carter et al., 1996).
Although it seems to be more clearly allied with sub-
genus Cyperus (Kükenthal, 1935–1936; Koyama,
1985), it has characteristics that seem to defy place-
ment there: namely, a bicarpellate gynoecium with
two stigmas and a lenticular achene with face adja-
cent to rachilla. When taken alone, the gynoecium
and fruit characteristics seem to indicate a relation-
ship with subgenus Juncellus (Clarke, 1908); howev-
er, both bi- and trigynous pistils have been found in
the same inflorescence (Koyama, 1985), which sup-
ports inclusion in subgenus Cyperus. In central
Florida, where this emergent aquatic sedge was
probably introduced with nursery stock used to
revegetate a reclamation wetland in an abandoned
phosphate pit, C. alopecuroides exhibits invasive
characteristics, forming extensive stands in shallow
water and floating mats (Carter et al., 1996).
According to Kantor (1999), C. alopecuroides was
cultivated by the ancient Egyptians and its inflores-
cence was widely depicted in one of the characteris-
tic motifs of their decorative art. Additional research
on C. alopecuroides is needed to understand better
its potential for becoming an invasive weed and its
phylogenetic relationships.
Cyperus odoratus is widely distributed in trop-
ical and subtropical regions around the world
(Kükenthal, 1935–1936; Kern, 1974) and is general-
ly found in disturbed hydric soils and wetlands. It is
frequently cited as a weed and has been listed as a
pest of rice (Appendix 2). In the southeastern
U.S.A., C. odoratus is commonly found in wet dis-
turbed sites, e.g., ditches, stream banks, swamps,
wetlands, and edges of ponds, but it is not of major
economic importance. Cyperus odoratus is classified
in subgenus Diclidium (Schrad. ex Nees) C. B.
Clarke [= Torulinium] characterized by spikelets that
disarticulate into one-fruited segments (Kükenthal,

40 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
1935–1936), and its achenes, enclasped within corky
rachilla segments, are dispersed by water (Kern,
1974; Haines & Lye, 1983). Jones et al. (1996) rec-
ognized several infraspecific taxa of this variable
species in North America. In the U.S.A., C. odoratus
is frequently associated with C. erythrorhizos Muhl.,
which is also listed as a weed (Holm et al., 1979;
WSSA, 1989). Cyperus erythrorhizos, a widespread
annual sedge restricted to North America
(Kükenthal, 1935–1936; Tucker et al., 2002), inhab-
its disturbed hydric soils of wetlands, ditches, stream
banks, floodplains, edges of ponds and swamps,
swales in fields and pastures, and occasionally rice
fields. It is of minor economic importance. Cyperus
digitatus Roxb. is closely related to C. erythrorhizos,
but it is perennial and generally a much larger plant.
Cyperus digitatus is widely distributed in tropical
and subtropical regions of both the Eastern and
Western hemispheres (Kükenthal, 1935–1936; Kern,
1974; Koyama, 1985) and,as can be seen in
Appendix 2, is frequently cited as a weed. Because it
is much more wide-ranging and cited as a pest of rice
in the Eastern Hemisphere (Kern, 1974), C. digitatus
is probably of greater economic significance than C.
erythrorhizos.
Cyperus articulatus L. ranges widely in tropi-
cal, subtropical, and warm temperate regions around
the world (Kükenthal, 1935–1936). It is a rhizoma-
tous perennial with a reed-like habit, septate culms,
and bladeless (usually) leaves. In the southeastern
U.S.A., C. articulatus occurs near the coast in
marshes, ditches, or other open disturbed sites, and
populations usually appear as scattered, solitary aer-
ial stems. As shown in Appendix 2, C. articulatus is
widely reported as a weed (Holm et al., 1979; Kühn,
1982; WSSA, 1989; Kissman, 1997).
Cyperus compressus is widely distributed in
tropical, subtropical, and warm temperate regions
around the world (Kükenthal, 1935–1936). It is fre-
quently cited as a weed and is found in a variety of
habitats disturbed and altered by humans, e.g., waste
places, grasslands, lawns, crops, roadsides, fallow
rice fields (Ohwi, 1965; Lin, 1968; Kern, 1974;
Kühn, 1982; Koyama, 1985; WSSA, 1989; Ravi &
Mohanan, 2002). In warmer parts of the southeastern
U.S.A., it is a common weed in sandy loam soils of
agricultural fields, roadsides, gardens, and other dis-
turbed sites. According to Bailey (1935) and Huxley
(1992), C. compressus has been cultivated as an
ornamental, which probably partly accounts for its
wide distribution.
Cyperus pilosus is a weed of tropical, subtropi-
cal, and warm temperate areas in Asia, western
Africa, and Australia (Kükenthal, 1935–1936;
Koyama, 1985; Wilson, 1993) and is commonly
cited as a weed of rice (McGivney, 1938; Kern,
1974; Wagner et al., 1990). It has been collected in
Hawaii, where it was possibly introduced with rice
agriculture, but has not been found there since 1916
(Wagner et al., 1990). Cyperus pilosus has been
known in the southeastern U.S.A. since 1938, where
it was probably introduced through the cultivation of
rice (McGivney, 1938; O’Neill, 1938a). In the south-
eastern U.S.A., it is found in rice fields, wet ditches,
and other wet disturbed sites and is apparently
spreading, having been reported new to several states
in recent years (Burkhalter, 1985; Bryson & Carter,
1992; Tucker et al., 2002). Cyperus procerus Rottb.
is related to C. pilosus. It is known from tropical and
subtropical regions of western Africa, India, Asia,
Malaysia, and Australia (Koyama, 1985; Wilson,
1993) and has been cited as a weed of rice fields in
Asia and western Africa (Hooper & Napper, 1972;
Kern, 1974). Cyperus pilosus and C. procerus share
several characteristics that distinguish them from
most other Cyperus spp.: stoloniferous habit, tri-
quetrous culm, and hispidulous rachis.
Cyperus sphacelatus Rottb. is widely distrib-
uted in the tropics and subtropics from eastern
Africa, Ceylon, Malaysia, northern Australia
(Queensland), Tahiti, South America, Central
America, and the Caribbean (Clarke, 1900; Uittien,
1932; Kükenthal, 1935–1936; Haines & Lye, 1983;
Tucker, 1983; Koyama, 1985). It is a heliophyte of
moist disturbed sites, beaches, riverbanks, fields, and
roadsides (Reed, 1977; Tucker, 1983; Carter et al.,
1996), and, in Malaysia, C. sphacelatus is reported-
ly a common weed on airstrips (Kern, 1974), which
suggests dispersal via air traffic. Mohr (1901) report-
ed C. sphacelatus from ballast heaps in Mobile,
Alabama, U.S.A., and more recently naturalized
populations have been found in southern Florida,
U.S.A. (Carter et al., 1996). An analysis of floral
scale length on herbarium specimens indicates that
the populations in Florida probably originated from
the West Indies (Carter et al., 1996). The recent dis-
covery of naturalized populations in peninsular
Florida suggests that C. sphacelatus is currently

The Significance of Cyperaceae as Weeds 41
undergoing range expansion in the southeastern
U.S.A. Field botanists and weed scientists should be
vigilant to detect additional populations of this intro-
duced pest, and appropriate governmental agencies
should initiate measures to survey for and eradicate
populations of C. sphacelatus in the U.S.A. before it
spreads further. The following combination of char-
acteristics distinguishes C. sphacelatus from other
Cyperus spp.: annual caespitose habit, triquetrous
achene, diffuse inflorescence with flattened
spikelets, and variegated floral scales pale, nearly
white, each with two conspicuous reddish patches.
Cyperus distans L. f. is a pantropical weed of
marshes, canal banks, ditches, agricultural crops,
and grasslands in Africa, India, Sri Lanka, southeast-
ern Asia, Malaysia, southern China, the Philippines,
the Caribbean islands, Central America, Mexico, and
tropical South America (Clarke, 1900; Uittien, 1932;
Kükenthal, 1935–1936; Koyama, 1985; Adams,
1994; Tucker, 1994). Cyperus distans is frequently
cited as a weed in the Eastern Hemisphere, where
aquatic biotypes are known, and it is a pest of rice
fields and grasslands (Appendix 2). It occurs sporad-
ically in the southeastern U.S.A. and has been
reported from coastal North Carolina, Georgia, and
Florida (Small, 1933; Kükenthal, 1935–1936;
McGivney, 1938; Radford et al., 1968; Beal, 1977;
Carter et al., 1996). The recent report (Carter et al.,
1996) from Florida, U.S.A., suggests that C. distans
is expanding its range there. The following combina-
tion of characteristics distinguishes C. distans from
other Cyperus spp.: rhizomes; scales ascending,
remote, with 3- to 5-nerved greenish keels, san-
guineous to reddish brown nerveless sides, and with
scarious emarginate tips. Field botanists and weed
scientists should seek and report additional popula-
tions, and appropriate state and federal agencies
should undertake eradication measures to ensure
early control of this potentially invasive pest in the
southeastern U.S.A.
A number of species classified by Kükenthal
(1935–1936) in Cyperus sect. Umbellati C. B.
Clarke are listed as weeds in Appendix 2. Cyperus
cyperinus (Retz.) Suringar and C. cyperoides (L.)
Kuntze are broadly distributed in warm parts of the
Eastern Hemisphere (Kükenthal, 1935–1936; Kern,
1974). Cyperus cyperoides is introduced in the West
Indies (Kükenthal, 1935–1936; Kern, 1974), and C.
cyperinus has been reported as a wool alien in Great
Britain (Sell & Murrell, 1996). Both species have
frequently been cited as weeds (Appendix 2). The
variation within these species is complex and poorly
understood, with numerous infraspecific taxa recog-
nized, and the synonymy is formidable (cf.
Kükenthal, 1935–1936; Kern, 1974; Haines & Lye,
1983; Koyama, 1985). No thorough systematic
review of this group has been done since Kükenthal
(1935–1936). Additional research to elucidate the
relationships of infraspecific taxa and their relation-
ships with one another and with related species, e.g.,
C. paniceus Boeckeler, is needed. Such research
with North American species of section Umbellati
has been productive, resulting in substantial taxo-
nomic and nomenclatural clarification (Carter, 1984;
Carter & Jarvis, 1986; Carter & Kral, 1990; Carter &
Jones, 1997).
Cyperus croceus Vahl, C. echinatus (L.) A. W.
Wood, and C. retrorsus Chapm. are listed as weeds
by WSSA (1989). All are caespitose perennials with
umbelliform inflorescences of simple spikes of
densely clustered spikelets, classified by Kükenthal
(1935–1936) in section Umbellati. These taxa are
native and widely distributed in the southeastern
U.S.A., where they are found in lawns, pastures,
roadsides, waste places, disturbed woodlands, and
other ruderal sites (Carter, 1984). Cyperus croceus
and C. echinatus were long known as C. globulosus
auct. non Aubl. and C. ovularis (Michx.) Torr.,
respectively (Carter & Kral, 1990). Cyperus croceus
also occurs in the Caribbean Islands (Carter, 1984).
Cyperus croceus and C. echinatus have been report-
ed as wool aliens in Great Britain (Sell & Murrell,
1996), and C. croceus has been associated with bal-
last (Britton, 1886). Cyperus echinatus is reportedly
naturalized in the Azores (DeFilipps, 1980c). Carter
(1984, in prep.) shows that plants with ascending
yellowish scales are distinct from C. retrorsus and
should be called C. ovatus Baldwin. Cyperus ovatus
is a coastal species in the southeastern U.S.A., which
is found in slightly wetter sites than related C. retror-
sus,e.g., moist ditches, disturbed sites in moist
sandy, loamy, or peaty soils in coastal flatwoods
(Carter, 1984, in prep.). Although not listed by
WSSA (1989), we include the related taxa C.
retroflexus Buckley and C. floribundus (Kük.) R.
Carter & S. D. Jones in Appendix 2, because they are
commonly weeds of roadsides, poorly kept lawns,
pastures, disturbed grasslands, and agricultural

42 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
fields; see Carter and Jones (1997) for clarification
of the taxonomy of C. retroflexus and its allies.
Cyperus floribundus is native to northeastern Mexico
and southern Texas (Carter, in prep.). Cyperus
retroflexus ranges from northern Mexico north
through Texas into Oklahoma and east into western
Mississippi, Arkansas, and Missouri and is apparent-
ly expanding its range eastward into Alabama,
Mississippi, and Missouri (Carter et al., 1987; Carter
& Bryson, 1991a, b). Cyperus plukenetii also
belongs to section Umbellati. It has spikelet modifi-
cations facilitating animal dispersal (Carter, 1993)
and is endemic to the eastern U.S.A., where it is well
adapted to open xeric sands of the Coastal Plain
(Carter, 1984, in prep.). Cyperus plukenetii currently
does not appear to be invasive in its natural range;
however, because of its specialized dispersal mecha-
nism and adaptation to dry soils, it could potentially
become an invasive weed if introduced into suitable
habitat elsewhere.
Cyperus aggregatus is frequently cited as a
weed (Appendix 2) and has been classified in section
Umbellati (Kükenthal, 1935–1936). The species was
previously called C. flavus (Vahl) Nees, nom. illeg.,
non J. Presl & C. Presl and C. cayennensis (Lam.)
Britton, non. illeg., non Willd. ex Link (Tucker,
1985). Cyperus aggregatus is native to Central and
South America, was introduced with ballast in the
U.S.A. (Britton, 1886; Mohr, 1901; Horvat, 1941;
Tucker et al., 2002), and is also introduced and
weedy in Australia (Wilson, 1993). Cyperus aggre-
gatus occurs sporadically in the Coastal Plain of the
southeastern U.S.A., where it can be locally abun-
dant and somewhat invasive on open, disturbed
sandy soils (Bryson & Carter, 1992; Tucker et al.,
2002; Carter, pers. obs.). It is likely to continue to
expand its range in warmer parts of the southeastern
U.S.A. and elsewhere.
Cyperus ligularis L. is occasionally cited as a
weed (Appendix 2). It is widely distributed in the
West Indies, Mexico, Central America, and South
America and is introduced in Africa and the south-
eastern U.S.A. (O’Neill, 1946; Tucker et al., 2002).
It is a frequent weed of disturbed sites in southern
peninsular Florida, U.S.A. (Wunderlin, 1998), and
Mohr (1901) reported that it was collected in 1891
on ballast at Mobile,Alabama, U.S.A., where it has
apparently not survived. Thus, C. ligularis is appar-
ently not tolerant of prolonged cold temperatures.
Cyperus ligularis is readily identified by its robust
caespitose habit; coarse lacerating leaves; grayish
green foliage; umbelliform inflorescence of dense,
oblong-cylindric, often branched spikes; and reddish
brown floral scales.
A number of Cyperus species colonize coastal
or inland sand dunes by forming extensive rhizomes.
Cyperus dentatus Torr., C. lecontei Torr. ex Steud.,
and C. onerosus M. C. Johnst. are related North
American species sharing similar growth forms and
habitats, spreading vegetatively through growth of
rhizomes. Cyperus lecontei is listed as a weed by the
WSSA (1989), and we have observed it colonizing
disturbed sandy roadsides and other open sandy
areas along the Gulf Coast. Cyperus lecontei and C.
dentatus are coastal fringe species. Cyperus lecontei
is found on exposed sands of coastal dunes and
swales in the southeastern U.S.A., and C. dentatus
occupies similar habitats but with a more northerly
distribution from the mid-Atlantic states north into
the maritime provinces of Canada (Tucker et al.,
2002). Cyperus onerosus is a related species endem-
ic to swales and pools far inland in nonmaritime
dune formations in western Texas (Carter, pers. obs.;
Tucker et al., 2002). Rhizomes of C. dentatus form
tubers, but do not in C. lecontei and C. onerosus
(Tucker et al., 2002). Cyperus arenarius Retz. ranges
from southern Iran through Pakistan, India, and
Ceylon into Indochina (Kükenthal, 1935–1936;
Koyama, 1985) and has been introduced into south-
ern Australia and southern Africa (Kukkonen, 2001).
Simpson and Inglis (2001) listed it as a weed, and
Koyama (1985: 209) described it as a coastal species
in Ceylon commenting that its “extensive rhizome
system … forms a large pure community.” The
endemic C. crassipes Vahl from coastal southeastern
Africa has a similar habit and habitat: sandy
seashores and riverbanks (Gordon-Gray, 1995).
Cyperus stoloniferus Retz., another vegetative colo-
nizer of coastal sands, ranges from Pakistan and
India to China and northern Australia and is also
known from Mauritius and Madagascar (Kukkonen,
2001). Although only C. arenarius and C. lecontei
are listed in Appendix 2, we think these ecologically
similar species have great potential to invade coastal
dunes or other open sandy areas, if introduced out-
side their natural ranges, as the alien Carex kobomu-
gihas done along the mid-Atlantic coast of North
America (Standley, 1983).

The Significance of Cyperaceae as Weeds 43
Cyperus fuscus is native to Eurasia and the
Mediterranean region of northern Africa and has
spread in Asia and Africa and into Greenland,
Iceland (Kükenthal, 1935–1936; DeFilipps, 1980c),
and North America (Smith, 1867; Britton, 1886;
Knowllton et al., 1911; McGivney, 1938; McKenzie
et al., 1998). This small caespitose annual produces
large numbers of tiny achenes. It is reported as a
weed in rice-producing areas of Asia and Portugal
and is a common weed in Afghanistan and Israel
(Weedon & Stephens, 1969; Holm et al., 1977;
Zhirong et al., 1990). Early records of C. fuscus in
North America were mostly associated with ballast
waste and wharf areas (Britton, 1886; Rhoads &
Klein, 1993). Cyperus fuscus seems to be expanding
its range in the U.S.A. (McKenzie et al., 1998), where
it is possibly still in the lag phase and could pose
problems in the future for rice agriculture.
Cyperus amabilis Vahl, C. cuspidatus Kunth,
and C. squarrosus L. are widely distributed in tropi-
cal, subtropical, and warm temperate regions of both
the Eastern and Western hemispheres (Kükenthal,
1935–1936; Kern, 1974). All three are diminutive
sedges listed in Appendix 2, and both C. amabilis
and C. cuspidatus have prominently cuspidate floral
scales. Cyperus amabilis has been reported as a
weed (Healy & Edgar, 1980; Kühn, 1982) and is
known from Africa, Asia, South America, Central
America, and North America (Kükenthal,
1935–1936; Tucker et al., 2002). As shown in
Appendix 2, C. cuspidatus and C. squarrosus are
listed as weeds of rice and are also found in waste
places, disturbed sites, sandy fields, and grasslands.
In warmer parts of the southeastern U.S.A., C. cusp-
idatus is sometimes locally abundant in disturbed
sandy soils in and around agricultural fields and has
also been observed as a weed in container-grown
plants and plant nurseries (Carter, pers. obs.).
Cyperus squarrosus is characterized by the distinc-
tive aroma of fenugreek (Trigonella foenum-grae-
cum L.), which is shared by C. fuscus,C. hyalinus
Vahl, and C. setigerus Torr. & Hook. (McKenzie et
al., 1998; Carter & Mears, 2000). Kern (1974)
showed that C. aristatus Rottb. is a synonym of C.
squarrosus, and, subsequently, contemporary work-
ers have followed Kern without reviewing the status
of a number of varieties and forms of C. aristatus
recognized by Kükenthal (1935–1936). North
American plants have been segregated as C. inflexus
Muhl. or C. aristatus var. inflexus (Muhl.) Boeckeler.
Preliminary research (Carter, unpubl. data) indicates
that C. inflexus is a smaller plant with smaller
spikelets and scales and supports its recognition as a
distinct endemic North American taxon. The names
C. inflexus and C. squarrosus var. runyonii (O’Neill)
S. D. Jones & Wipff were placed into synonymy,
without justification, under C. squarrosus by Tucker
et al. (2002). Cyperus granitophilus McVaugh is an
autotetraploid derivative of C. squarrosus, endemic
to granite and sandstone outcrops in the Piedmont
region from Virginia to Georgia, U.S.A. (Garoni &
Murdy, 1964; Tucker et al., 2002). Preliminary
observations (Carter, unpubl. data) indicate that C.
granitophilus is a coarser plant than the more com-
mon widespread C. inflexus and is morphologically
more similar to C. squarrosus. Although new taxa
have been described and other major nomenclatural
changes have occurred, the entire complex has not
been studied since Kükenthal (1935–1936). A sys-
tematic review worldwide of C. squarrosus and
related taxa is needed to achieve a consistent treat-
ment of these and other infraspecific taxa not
accounted for by contemporary authors.
Cyperus gracilis R. Br., yet another diminutive
sedge cited as a weed (Holm et al., 1979), is native
to Australia, where it grows in open woodlands and
grasslands in drier sites than related species (Wilson,
1993). It was once promoted for use as a ground
cover in Hawaii, where it is naturalized and common
in disturbed sites, lawns, and roadsides (Hughes,
1995). It is also introduced in California, U.S.A.
(Tucker et al., 2002).
Cyperus subg. Pycreus is characterized by per-
sistent rachillae, bifid styles, and lenticular achenes
with the achene angle adjacent to the rachilla
(Kükenthal, 1935–1936; Tucker et al., 2002), which
some treat as genus Pycreus (e.g., Koyama, 1985;
Adams, 1994; Gordon-Gray,1995). Weeds belong-
ing to subgenus Pycreus include C. flavescens L., C.
flavicomus Michx., C. flavidus Retz., C. lanceolatus
Poir., C. polystachyos Rottb., C. puncticulatus Vahl,
C. pumilus L., C. sanguinolentus, and C. substramineus
Kük.
Cyperus sanguinolentus has been frequently
cited as a weed (Holm et al., 1977; Reed, 1977;
Kühn, 1982; Zhirong et al., 1990). It is widely dis-
tributed in the Eastern Hemisphere,where it is
known from northeastern Africa, the Middle East,

44 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
India, Sri Lanka, central Asia, southeastern Asia,
China, Taiwan, Japan, Korea, the Philippines,
Indonesia, Malaysia, and Australia (Clarke, 1894;
Kükenthal, 1935–1936; Ohwi, 1965; Kern, 1974;
Holm et al., 1977; Reed, 1977; Kühn, 1982; Haines
& Lye, 1983; Zhirong et al., 1990; Wilson, 1993). In
the Western Hemisphere, it has been reported from
Hawaii (Wagner et al., 1990) and from the Coastal
Plain of the southeastern U.S.A. in North America
(Carter & Bryson, 2000b, 2001). Cyperus sanguino-
lentus is a weed in paddy and damp, low-dryland
crop fields in Asia (Holm et al., 1977; Zhirong et al.,
1990). In the southeastern U.S.A. it is a locally com-
mon annual weed in periodically disturbed sites with
hydric soils, e.g., road ditches and margins of artifi-
cial ponds (Carter & Bryson, 2000b). Its introduc-
tion into the southeastern U.S.A. is associated with
the cultivation of rice, and its dispersal and range
expansion there are associated with road construc-
tion and maintenance activities (Carter & Bryson,
2000b). Cyperus louisianensis Thieret, once thought
to be a rare endemic species in southern Louisiana,
U.S.A. (Thieret, 1977), has been shown to be the
nonindigenous weed C. sanguinolentus (Carter &
Bryson, 2000b).
Cyperus flavescens is widely distributed in both
Old and New Worlds (Kükenthal, 1935–1936;
O’Neill, 1946; Barros, 1960; Haines & Lye, 1983;
Tucker et al., 2002). It is a common weed of seeps,
roadside ditches, and disturbed wet sites in Natal
Province (now KwaZulu-Natal Province), South
Africa (Gordon-Gray, 1995). In the U.S.A. it is a
common tuft-forming weed in drainage ditches, dis-
turbed hydric sites, and moist lawns and fields
(Carter, pers. obs.), ranging widely from Florida
north into southern Canada and west to Texas and
Missouri (Tucker et al., 2002). Cyperus lanceolatus
is similar to C. flavescens and frequently occurs in
the same habitats in the southeastern U.S.A. Both
taxa have similar habits (dense tufts) and differ pri-
marily in the color of their achenes: black in C.
flavescens,brown in C. lanceolatus. Apparently less
tolerant of cold winter temperatures, C. lanceolatus
is restricted to the warmest parts of the southeastern
U.S.A., ranging from Florida north into southern
Georgia then west along the coast to Texas (Bryson
et al., 1996; Tucker et al., 2002). Cyperus lanceola-
tus also occurs in the West Indies, Mexico, Central
and South America, and Africa (O’Neill, 1946;
Barros, 1960; Haines & Lye, 1983; Tucker, 1994).
Although the type locality is Georgia, U.S.A.
(Elliott, 1821), C. fasciculatus Elliott is not cited by
contemporary American authors (e.g., Tucker, 1994;
Tucker et al., 2002) but is cited as a weed in Asia
(Appendix 2). This problem should be researched to
determine how the name C. fasciculatus impinges on
nomenclature of the North American plants current-
ly known as C. flavescens and C. lanceolatus.
Cyperus polystachyos is a cosmopolitan weed,
widely distributed in tropical, subtropical, and warm
temperate areas (Uittien, 1932; Kükenthal,
1935–1936; Barros, 1960; Kern, 1974; DeFilipps,
1980c; Haines & Lye, 1983; Tucker, 1983, 1994;
Koyama, 1985; Wilson, 1993; Adams, 1994;
Gordon-Gray, 1995). Cyperus polystachyos is cited
as a weed of hydric soils in ditches, waste
places, grasslands, and disturbed agricultural areas
and fields, including rice fields (Kern, 1974; Haines
& Lye, 1983). Cyperus polystachyos is taxonom-
ically and nomenclaturally complex. Kükenthal
(1935–1936) segregated 16 infraspecific taxa from
C. polystachyos (11 varieties, 5 forms). Also, the rela-
tionships between C. polystachyos and related taxa
like the North American species C. filicinus Vahl and
C. fugax Liebm. are poorly understood and are in
need of clarification. We have observed at least three
entities passing as C. polystachyos in the southeast-
ern U.S.A., with the greatest diversity centered along
the Gulf Coast. This group is in need of critical tax-
onomic review on a worldwide basis. Cyperus poly-
stachyos is cited as a weed of hydric soils in ditches,
waste places, grasslands, and disturbed agricultural
areas and fields, including rice fields (Kern, 1974;
Haines & Lye, 1983).
Cyperus flavicomus is found in North America,
South America, and Africa (Kükenthal, 1935–1936;
Barros, 1960; Tucker, 1994) and in Appendix 2 is
cited as a weed of waste places, rice fields and vari-
ous other crops, pastures, and turf. Cyperus flavico-
mus occurs sporadically on moist soil in and around
agricultural fields in the southeastern U.S.A., where
it is of minor importance as a weed. Cyperus pumilus
is a diminutive, densely tufted annual and a minor
weed of disturbed sandy soils of rice fields and fal-
low fields (Kern, 1974; Haines & Lye, 1983; Carter,
pers. obs.). As shown in Appendix 2 it is widespread
in the Old World. In the New World, C. pumilus is
known from the West Indies and the U.S.A.

The Significance of Cyperaceae as Weeds 45
(Kükenthal, 1935–1936; Kern, 1974; Haines & Lye,
1983; Koyama, 1985; Gordon-Gray, 1995). In the
U.S.A., C. pumilus has long been known from
Florida (Chapman, 1889 [as C. divergens Kunth];
Small, 1933; Long & Lakela, 1971; Godfrey &
Wooten, 1979; Clewell, 1985; Wunderlin, 1998) and
was reported in 1996 in southern Georgia (Bryson et
al., 1996). Cyperus pumilus appears to be spreading
in the southeastern U.S.A., as plants were found in
northern Georgia in 2003 (M. Czarnota s.n.,29
January 2003, VSC).
Cyperus hyalinus is transitional between
Cyperus and Kyllinga and is pragmatically treated
here in subgenus Queenslandiella (Domin) Govind.
Its taxonomic affinities are unclear, and it has been
variously placed in Pycreus,Kyllinga,Cyperus, and
the monotypic genus Queenslandiella based on mor-
phological traits (Clarke, 1884; Kükenthal,
1935–1936; Kern, 1974; Govindarajalu, 1975; Haines
& Lye, 1983). However, recent molecular evidence
suggests that Kyllinga,Pycreus, and, by extension,
Queenslandiella should be included in Cyperus
(Muasya et al., 2002). Cyperus hyalinus is known
from eastern Africa, Madagascar, Mauritius, India, Sri
Lanka, tropical Australia (Queensland), and Malaysia
(Kükenthal, 1935–1936; Kern, 1974; Haines & Lye,
1983; Koyama, 1985) and has recently been found in
southern Florida, where it was apparently introduced
by air traffic (Carter & Mears, 2000). Because Haines
and Lye (1983: 293) described it as “a weed of sandy
soils, near sea level” in eastern Africa and it is similar
in habit and habitat to certain weeds in Kyllinga, we
suspect that C. hyalinus could become a pest in turf,
flowerbeds, and containerized nursery plants in the
southeastern U.S.A. Additional research is needed to
determine its potential as a weed and to clarify its tax-
onomic relationships.
Cyperus laevigatus L. and C. serotinus Rottb.
are frequently cited as weeds (Appendix 2). Both
species have lenticular achenes with the achene face
adjacent to the rachilla and, thus, are classified into
subgenus Juncellus. Cyperus laevigatus is cosmo-
politan in tropical and warm temperate regions
(Kükenthal, 1935–1936; DeFilipps, 1980c). Aquatic
biotypes have been reported by Kühn (1982), and
this highly variable perennial sedge generally grows
in saline,alkaline,or mineral soils associated with
salt marshes in coastal areas or salt lakes, hot
springs, or artesian wells or along rivers inland
(Kükenthal, 1935–1936; Haines & Lye, 1983;
Wilson, 1993; Gordon-Gray, 1995; Tucker et al.,
2002). Cyperus laevigatus was collected along the
coast of North Carolina, U.S.A., where it was report-
edly introduced with ballast, but it apparently no
longer exists there (Radford et al., 1968). A number
of varieties were recognized by Kükenthal
(1935–1936), which contemporary authors ignore.
Wilson (1993) noted the presence of three forms of
C. laevigatus in Australia and stated the need for its
taxonomic study on a worldwide basis. Cyperus
serotinus occurs from the Mediterranean region of
southern Europe through much of Eurasia
(Kükenthal, 1935–1936; DeFilipps, 1980c), and it is
introduced sparingly in salt marshes along the mid-
Atlantic coast of North America (Tucker et al.,
2002). Kühn (1982) reported aquatic biotypes in C.
serotinus, indicating it as a weed of rice fields in
Asia. If introduced more widely, C. serotinus could
pose problems for rice agriculture in North America
and elsewhere.
ELEOCHARIS
Eleocharis is a genus of ca. 200 species world-
wide (Smith et al., 2002), about half of which are
aquatic or semi-aquatic (Holm et al., 1997).
Appendix 2 lists 53 species of Eleocharis as weeds.
Of 118 species of Eleocharis studied by Ueno et al.
(1989), all but six were shown to have C3photosyn-
thesis. Holm et al. (1997) considered E. acicularis
(L.) Roem. & Schult., E. dulcis Trin. ex Hensch., and
E. palustris (L.) Roem. & Schult. to be among the
world’s worst weeds and cite E. acicularis among
the five most troublesome weeds in Asian rice pad-
dies. Elliott (1821: 79) described E. quadrangulata
(Scirpus quadrangulatus Michx.) as “very injurious”
in rice fields of Georgia and South Carolina, U.S.A.
The tubers of E. dulcis are consumed as Chinese
water chestnuts, and the species is widely cultivated
in Asia (Kern, 1974; Mabberley, 1997). Sculthorpe
(1967) cited E. acicularis and E. palustris among the
most broadly distributed aquatic plants in the world,
and Svenson (1957) cited the cosmopolitan weed E.
geniculata (L.) Roem. & Schult. (as E. caribaea
(Rottb.) S. F. Blake) as the most widespread
Eleocharis species. As shown in Appendix 2, E.
geniculata is frequently cited as a weed and has been
reported as a pest in rice (Kern, 1974) and taro pad-
dies (Wagner et al., 1990). Eleocharis radicans

46 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
(Poir.) Kunth is reportedly naturalized in taro pad-
dies in Hawaii (Wagner et al., 1990). Walters (1980)
reported the South American species E. bonariensis
Nees as naturalized on banks of estuarine rivers in
France, with no indication of it being a weed.
Eleocharis macrostachya,E. mamillata H.
Lindb., and E. palustris belong to a taxonomically
perplexing complex and are widely distributed
around the world, found in hydric soils in a variety
of habitats, e.g., pond margins, marshes, ditches, and
wet meadows (Svenson, 1957; Smith et al., 2002).
All three taxa are listed as weeds in Appendix 2.
Additionally, the related taxon, E. erythropoda
Steud., is sometimes associated with disturbance
(Smith et al., 2002) and, therefore, might be consid-
ered a weed. Eleocharis palustris and E. mamillata
are found in both the Eastern and Western hemi-
spheres, whereas E. macrostachya and E. erythropo-
da are restricted to the New World (Smith et al.,
2002). Eleocharis macrostachya has an essentially
western distribution in the U.S.A., ranging from
Alaska, south through British Columbia and
California, east to Mississippi, Illinois, and
Wisconsin; it is also in Mexico and South America
(Svenson, 1957; Smith et al., 2002). Eleocharis
macrostachya seems to be dispersing eastward in the
U.S.A. and was only recently reported from western
Mississippi where it was locally abundant in hydric
soils in a roadside ditch along a major highway
(Bryson et al., 1996).
Although none of the primary sources used to
compile Appendix 2 lists either Eleocharis montevi-
densis Kunth or E. montana (Kunth) Roem. &
Schult. as weeds, we have included them based upon
observations made in the southeastern U.S.A.
Eleocharis montevidensis is widely distributed in
North and South America and restricted to the
Western Hemisphere (Svenson, 1957; Smith et al.,
2002). In the southeastern U.S.A., this rhizomatous
perennial is sometimes locally abundant and weedy
in hydric soils of ditches, roadsides, or other dis-
turbed sites (Carter,pers. obs.). Eleocharis montana
is a perennial restricted to the Western Hemisphere.
It is probably indigenous to South America and is
known from the Gulf coastal states of the southern
U.S.A., the Caribbean, and throughout much of
South America (Svenson, 1957; Smith et al., 2002).
In the southeastern U.S.A., E. montana is a weed of
hydric soils in disturbed areas and roadside ditches
and seems to be most common on fine-textured soils
in the rice-growing areas of southern Louisiana and
eastern Texas (Carter, pers. obs.).
Eleocharis albida is common along the Gulf
and Atlantic coasts in the southeastern U.S.A.; it also
occurs in Mexico and the Caribbean (Smith et al.,
2002). It is often locally abundant in hydric brackish
soils of disturbed open sites and ditches adjacent to
salt marshes (Carter, 2005). Extensive coastal real
estate development has undoubtedly facilitated the
expansion of E. albida in the southeastern U.S.A.
Although we include it in Appendix 2 because of its
propensity to form extensive stands following distur-
bance, we do this with some reservation, because it
is indigenous and is apparently invasive only in areas
where humans have severely altered the habitat.
Eleocharis parvula is frequently associated with E.
albida along the coast in the southeastern U.S.A.
(Carter, 2005); however, it is much more widely dis-
tributed, ranging throughout much of eastern North
America from eastern Canada southward into
Central America; it also occurs along the West Coast
of North America and in Eurasia (Smith et al., 2002).
Like E. albida,E. parvula can be locally abundant in
disturbed, hydric, brackish soils in coastal areas;
however, E. parvula also occurs sporadically inland.
A number of salt marsh species, including the sedges
E. parvula and Bolboschoenus robustus [= Scirpus
robustus Pursh], occur 400 km from the coast in
western Virginia, U.S.A., on saline soils formed by
the pumping of brine wells and are thought to have
been dispersed there by birds (Sauer, 1988).
Eleocharis baldwinii and E. vivipara are listed as
weeds (WSSA, 1989). Both species are profuse in their
vegetative proliferation and also reproduce from
achenes. Eleocharis baldwinii is common in parts of
the southeastern U.S.A., where it may be locally abun-
dant in dense mats on disturbed moist sandy or peaty
soils or floating in ponds or ditches (Carter,pers.
obs.). Eleocharis vivipara spreads vegetatively, form-
ing dense clumps on moist soil or tangled mats in ponds
and ditches (Carter, pers. obs.). In the U.S.A., both E.
baldwinii and E. vivipara are currently of minor eco-
nomic importance as weeds and are probably only
opportunistically weeds following disturbance by
humans. However, because of their ability to proliferate
vegetatively and to reproduce sexually from achenes,
we suspect they could become invasive if introduced
into suitable habitats outside their natural ranges.

The Significance of Cyperaceae as Weeds 47
Eleocharis ovata (Roth) Roem. & Schult. and
E. obtusa (Willd.) Schult. are cited as weeds
(Appendix 2; WSSA, 1989; Callahan et al., 1995) in
North America. Eleocharis ovata ranges broadly in
Eurasia and throughout much of Canada and the
northern U.S.A. (Svenson, 1957). Eleocharis obtusa
is common throughout much of North America and
is naturalized in Hawaii (Svenson, 1957; Smith,
2002c) and in rice fields in southern Europe
(Walters, 1980). Both E. ovata and E. obtusa are
closely related caespitose annuals (rarely perenni-
als), found in seasonally wet disturbed sites, margins
of ponds, and ditches (Svenson, 1957). Tufts of these
plants continue to increase in diameter, producing
new fertile culms throughout the growing season so
long as there is sufficient moisture (Bryson, pers.
obs.). Eleocharis engelmannii Steud. and E. lanceo-
lata Fernald are related taxa, similar to and easily
confused with E. ovata and E. obtusa in habitat and
growth characteristics. Eleocharis engelmannii
occurs sporadically throughout much of the range of
E. obtusa and differs from that species primarily in
its shorter tubercle. Eleocharis lanceolata is found in
the south-central U.S.A. and was collected in 1949
in California as a weed in a rice field (Smith, 2002c).
Hybrids between E. lanceolata and closely related E.
obtusa are known (e.g., Carr 13969, VSC). Its narrow-
er, more cylindrical spikelet, acute scale, and narrower
tubercle distinguish E. lanceolata from E. obtusa
(Svenson, 1957; Smith, 2002c). Eleocharis flavescens
(Poir.) Urb. var. flavescens and E. flavescens var. oli-
vacea (Torr.) Gleason, like their New World relative E.
obtusa,have become naturalized in rice fields of south-
ern Europe (Walters, 1980).
Eleocharis sect. Limnochloa (P. Beauv. ex T.
Lestib.) Torr. is a group of robust (for Eleocharis)
emergent aquatics. These perennial species show con-
siderable variation in the shape of their stems in trans-
verse section, from terete,to triquetrous, to quadran-
gular (Svenson, 1957). As shown in Appendix 2, a
number of species in this group are cited as weeds,
including E. acutangula (Roxb.) Schult., E. cellulosa
Torr., E. dulcis,E. interstincta (Vahl) Roem. &
Schult., E. mutata (L.) Roem. & Schult., E. philip-
pinensis Svenson, and E. quadrangulata. Eleocharis
acutangula and E. mutata are widely distributed in
both hemispheres (Svenson, 1957; Koyama, 1985),
whereas E. cellulosa,E. interstincta,and E. quadran-
gulata are exclusively New World species (Svenson,
1957). Eleocharis philippinensis and E. dulcis are
wide-ranging in the Eastern Hemisphere, where E.
dulcis is widely introduced and naturalized from cul-
tivation for its tubers (Chinese water chestnuts) (Kern,
1974; Koyama, 1985). Several species are cited as
pests in rice fields, and given their aquatic habitat and
emergent habit, it would appear that all have the
potential to be weeds of rice agriculture or invasive
pests of wetlands in natural areas (Kern, 1974; Holm
et al., 1979; Koyama, 1985). As discussed in the
Dispersal section, there is considerable potential for
achenes of these species to be disseminated long
distances by waterfowl.
FIMBRISTYLIS
There are more than 100 species of Fimbristylis
worldwide (Kral, 2002b), and 46 are listed as weeds
in Appendix 2. Fimbristylis dichotoma (L.) Vahl and
F. miliacea (L.) Vahl are co-ranked as the world’s
40th worst complex of weeds (Holm et al., 1977).
Fimbristylis dichotoma is a rapidly growing annual
or perennial that thrives in poorly aerated soils with
high moisture content (Holm et al., 1977). It has
been reported as a weed of paddy crops, old rice
fields, ditches, lawns, open wetland pastures and
meadows, roadsides, cultivated lands, and along for-
est margins in 21 countries throughout the tropical
and semitropical regions of the world including
Africa, Asia, the Pacific Islands, and North and
South America (Holm et al., 1977). Fimbristylis
dichotoma is cited as a weed in pineapple, rice,
roselle, teak, taro, and other upland row crops (Holm
et al., 1977). In the southeastern U.S.A., F.
dichotoma,F. caroliniana (Lam.) Fernald, and F.
castanea (Michx.) Vahl are frequently weeds follow-
ing mechanical disturbance of the soil (Kral, 1971).
Fimbristylis miliacea, a native to tropical
America, is now a troublesome weed in Africa, Asia,
Australia, and North and South America in 21 coun-
tries (Holm et al., 1977). It is considered a major
weed in rice in Asia, but it is also a weed of taro,
bananas, corn, sorghum, and sugarcane (Holm et al.,
1977). Fimbristylis miliacea, an annual or some-
times perennial in the tropics, is reported to produce
more than 1000 seeds per plant per year and without
dormancy (Holm et al., 1977). Seeds of F. miliacea
are easily dispersed and seedlings emerge rapidly on
moist soil (Holm et al., 1977). Infestations can con-
stitute 70% of all seedling weeds in agricultural

48 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
areas (Verga & Sierra, 1970), and in Malaysia, F.
miliacea is reported to be the first sedge emerging
after rice planting and the first sedge to recover fol-
lowing tillage (Burkill, 1935). Emergence of F. mili-
acea seedlings seems to be environmentally depend-
ent. In Japan, rice planted mid-season reduced the
number of emerging F. miliacea seedlings by 80%
when compared to rice planted early season, and
seedling emergence was even less in late-season rice
plantings (Noda & Eguchi, 1965).
Fimbristylis annua (All.) Roem. & Schult. and
F. autumnalis (L.) Roem. & Schult. are also listed as
weeds in North America (WSSA, 1989) but are not
as troublesome as F. miliacea in rice production in
the southeastern U.S.A. At least some of the forms of
F. annua were introduced into the U.S.A. with rice
agriculture (Kral, 1971). Fimbristylis decipiens Kral
was described from specimens collected in the
U.S.A. (Kral, 1971). Because it is morphologically
similar to and often occurs with F. annua and F.
dichotoma, herbarium specimens of these three
species are difficult to distinguish (Kral, 1971). A
number of Fimbristylis spp. are thought to have been
introduced in the U.S.A. and elsewhere around the
world with rice agriculture (Appendix 1). Fim-
bristylis aestivalis (Retz.) Vahl has been reported as
a weed of rice and in taro paddies in the Eastern
Hemisphere and in Hawaii (Kern, 1974; Wagner et
al., 1990; Ravi & Mohanan, 2002).
FUIRENA
The 30 species of Fuirena worldwide are near-
ly all heliophytic wetland plants of acidic soils in the
tropics and subtropics (Kral, 1980, 2002a). Eight
species are listed in Appendix 2, including two, F.
ciliaris (L.) Roxb. and F. umbellata Rottb., cited as
weeds of rice fields in the Eastern Hemisphere.
Fuirena breviseta (Coville) Coville, F. pumila (Torr.)
Spreng., F
. scirpoidea Michx., F
. simplexVahl, and
F. squarrosa Michx. are weeds in the U.S.A.
(WSSA, 1989), where they occur in wet soils of pas-
tures or along waterways and roadsides. None of the
Fuirena spp. is a major weed.
ISOLEPIS
Isolepis contains about 69 species worldwide,
predominately found in cool-tropical and temperate
regions of Africa and Australia (Smith, 2002d); a
single species is listed as a weed in Appendix 2.
Isolepis carinata [= Scirpus koilolepis (Steud.)
Gleason] is occasionally a weed on moist bare soils
in gardens, row crops, and natural areas, following
fire or tillage (Carter et al., 1990; Bryson & Hanks,
2001). It is usually not a particularly troublesome
weed in row crops because of its diminutive stature,
susceptibility to foliar herbicides, and early-season
phenology. Isolepis cernua is widely distributed
around the world primarily in temperate and subtrop-
ical regions, occurring in southern Africa (absent
from tropical Africa), Eurasia (absent from south-
eastern Asia), Australia and New Zealand, temperate
South America, and North America (Wilson, 1981;
Gordon-Gray, 1995; Smith, 2002d). It is apparently
a recent arrival (since 1888) in the U.S.A. and
Canada, where it is found primarily on the Pacific
coast in fresh to brackish water on beaches, dunes,
and marine bluffs (Smith, 2002d). It is also known
from Texas, where the earliest collection seen by
Smith (2002d) was from 1974. The taxonomy of I.
cernua and related species is in need of revision on a
worldwide basis to clarify relationships of taxa and
complex nomenclature (Wilson, 1981; Gordon-Gray,
1995). According to Smith (2002d), only I. cernua
var. cernua is known from North America. Although
no citations were found of I. cernua as a weed, it is
included here because of its apparent introduction
into the U.S.A. and its potential to be introduced and
naturalized elsewhere in temperate and subtropical
areas through the ornamental trade (Bailey, 1935;
Everett, 1980–1982; Grounds, 1989; Greenlee &
Fell, 1992; Huxley, 1992; Darke, 1999).
KYLLINGA
Kyllinga, a genus of short rhizomatous perennials
or caespitose annuals, consists of 40 to 45 species dis-
tributed in tropical, subtropical, and warm temperate
regions around the world (Tucker, 1984, 1987,
2002b). Appendix 2 lists 13 species as weeds, and K.
brevifolia is among the world’s worst weeds, having
been reported in 17 crops and 43 countries (Holm et
al., 1997). The maximum diversity of Kyllinga occurs
in tropical East Africa and Madagascar, where there
are 30 to 35 species (Kükenthal, 1935–1936; Haines
& Lye, 1983). An additional 11 to 12 Kyllinga species
occur in Asia and two occur in Australia; none is
native to Europe and only one is thought to be native
to North America. Kyllinga brevifolia,K. gracillima,
K. odorata Vahl, K. pumila Michx., and K. squamula-

The Significance of Cyperaceae as Weeds 49
ta are known from the continental U.S.A. (Kartesz,
1994). Kyllinga brevifolia and K. nemoralis (J. R.
Forst. & G. Forst.) Dandy ex Hutch. & Dalziel are
introduced weeds in Hawaii (Delahoussaye & Thieret,
1967; Holm et al., 1979; Tucker, 1987). Kyllinga poly-
phylla Willd. ex Kunth, a native of Africa, is intro-
duced into Samoa, Tahiti, and Fiji, where it is a weed
of disturbed places, pastures, and roadsides at eleva-
tions up to 700 m (Whistler, 1994). Spreading by rhi-
zomes, it is a particularly serious pest in pastures
because it displaces acceptable forage and is not eaten
by livestock (Whistler, 1994).
Kyllinga pumila, a weed of lawns and turf, was
initially described in the first North American flora by
Michaux (1803) and is evidently the only Kyllinga
species native to the continental U.S.A. Kyllinga brev-
ifolia,K. gracillima,K. odorata, and K. squamulata
are all pantropical species (Reed, 1977; Holm et al.,
1979; Tucker, 1984, 1987; Koyama, 1985) and were
apparently all introduced into the continental U.S.A.
from Asia. Although the precise time of their introduc-
tions is unknown, K. brevifolia was established in the
U.S.A. prior to 1821 (Elliott, 1821), and K. odorata
before 1836 (Torrey, 1836). Both are widespread in the
eastern U.S.A., especially in the southern Atlantic and
Gulf coastal plains, and are introduced weeds of South
America (Bryson et al., 1996; Kissmann, 1997). In the
U.S.A., distributions and recent range expansions indi-
cate later introductions for K. gracillima and K. squa-
mulata (Delahoussaye & Thieret, 1967; Sipple, 1978;
Ferren & Schuyler, 1980; Kral, 1981; Webb & Dennis,
1981; Webb et al., 1981; Wunderlin, 1982; Snyder,
1983, 1984; Naczi, 1984; Naczi et al., 1986; Sundell &
Thomas, 1988; Bryson & Carter, 1992, 1994; Mears &
Libby, 1995; Bryson et al., 1996). Kyllinga brevifolia
and K. odorata have continued to spread northward
and westward in the U.S.A., especially as weeds of
turf, pastures, and roadways (Bryson & Carter, 1992,
1994; Jones et al., 1993; Bryson et al., 1996), while K.
gracillima continues to spread south and westward
(Sipple, 1978; Ferren & Schuyler, 1980; Kral, 1981;
Webb & Dennis, 1981; Webb et al., 1981; Snyder,
1983, 1984; Naczi, 1984; Naczi et al., 1986; Sundell &
Thomas, 1988; Bryson & Carter, 1994; Mears &
Libby, 1995; Bryson et al., 1996, 1997).
The small achenes of the introduced Kyllinga
spp. could have arrived in the U.S.A. by a variety of
dispersal methods. Following introduction, Kyllinga
probably first naturalized along sandbars and dis-
turbed areas along streams or in open ruderal sites
with adequate moisture. Kyllinga spp. are common
weeds of highly maintained, frequently irrigated turf
in urban areas and on golf courses, and such sites
now provide excellent habitat for local proliferation,
dispersal, and range expansion of populations
(Yelverton, 1996). Kyllinga spp. are also frequent
weeds of mulched irrigated flowerbeds and con-
tainerized nursery plants (Whitwell & Smith, 1997).
Kyllinga brevifolia and K. gracillima are rhi-
zomatous perennials, and K. odorata,K. pumila, and
K. squamulata are annuals or short-lived perennials
in warmer climates. Kyllinga brevifolia flowers 10 to
12 weeks after germination and produces mature
seeds three weeks after flowering (Holm et al.,
1997). Kyllinga brevifolia seeds are disseminated by
wind and water and germinate without aging
(Sumaryono & Basuki, 1986), and human activities
result in the movement of whole plants, fragments,
or seeds in sod, soil, or grass clippings. A combina-
tion of frequent (often daily) irrigation and mowing
(3–6 times/week) without removal of clippings,
especially around golf course greens, enhances veg-
etative growth of perennial Kyllingaspecies
(Yelverton, 1996). Kyllinga brevifolia and K. gracil-
lima produce culms that produce fruit below most
turfgrass mowing heights (< 1.25 cm), resulting in a
reproductive advantage over many other weeds, and
they spread rapidly in turf via rhizome growth
(Yelverton, 1996). Factors contributing to the
increasing importance of Kyllinga species as weeds
include irrigation of turf, type and timing of herbi-
cide applications, use of fertilizer, and the expansion
in the container nursery plants and turfgrass industry
to meet the increasing demand for “instant,” well-
manicured flowerbeds, lawns, and golf courses
(Yelverton, 1996; Bryson et al., 1997).
Kyllinga brevifolia and K. gracillima are similar
in appearance and difficult, if not impossible, to dis-
tinguish vegetatively (Yelverton, 1996). Collections
of fruiting specimens of K. gracillima are primarily
from late August until frost, suggesting that the initi-
ation of flowering is dependent upon photoperiod.
The more northern distribution of K. gracillima in
the U.S.A. suggests that it can withstand cooler win-
ter temperatures. Kyllinga brevifolia,K. odorata,K.
pumila,and K. squamulata flower and produce fruit
during the frost-free months throughout their ranges
in the continental U.S.A. (Bryson et al., 1997).

50 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
LEPIDOSPERMA
Lepidosperma is a genus of ca. 60 species dis-
tributed in tropical and subtropical areas of China,
Malaysia, Australia, New Caledonia, and New
Zealand (Kern, 1974; Mabberley, 1997). One
species, L. chinense Nees & Meyen, grows in rice
fields in southern China (Kern, 1974) and is cited as
a weed in Appendix 2.
LEPIRONIA
Lepironia Rich. is a genus of five species dis-
tributed in Polynesia and Madagascar (Mabberley,
1997). Lepironia articulata (Retz.) Domin, cultivat-
ed for fibers used in sails and as packing material
(Mabberley, 1997), is a weed of rice fields in
Malaysia (Moody, 1989) and is cited in Appendix 2.
LIPOCARPHA
Lipocarpha R. Br. (including Hemicarpha
Nees) consists of ca. 35 species of wet pantropical
and warm temperate regions (Tucker, 2002c). In
Appendix 2, three species are cited as weeds of rice or
other wet agricultural fields: Lipocarpha chinensis
(Osbeck) J. Kern, L. microcephala (R. Br.) Kunth, and
L. squarrosa (L.) Goetgh. (Lin, 1968; Kern, 1974;
Holm et al., 1979; Kühn, 1982; Koyama, 1985).
Additionally, we have observed L. maculata (Michx.)
Torr. in the southeastern U.S.A. as a weed of disturbed
hydric soils, poorly kept moist lawns, roadsides, and
ditches. Based upon our observations of its habitat and
the citation of congeners as weeds in the Eastern
Hemisphere (Kern, 1974; Koyama, 1985), we suspect
that L. maculata could become a weed in rice fields in
the U.S.A. and elsewhere.
MAPANIA
Mapania (including Thoracostachyum Kurz) is
a genus of 73 species distributed in tropical and sub-
tropical areas of Asia (Mabberley, 1997). Mapania
cuspidata (Miq.) Uittien grows in rice fields in
Indonesia (Moody, 1989) and is cited as a weed in
Appendix 2.
OXYCARYUM
Oxycaryum Nees is a monotypic genus widely
distributed in the tropics and subtropics of Africa and
the Americas (Bruhl, 2002). The only species, O.
cubense, is in the West Indies (Kunth, 1837), South
and Central America (Nees von Esenbeck, 1842;
Adams, 1994), the southeastern U.S.A. (Chapman,
1889; Small, 1933; Godfrey & Wooten, 1979;
Tucker, 1987), and tropical Africa (Lye, 1971;
Hooper & Napper, 1972; Haines & Lye, 1983). In
the southeastern U.S.A., it occurs sporadically in
Florida (Chapman, 1889; Clewell, 1985; Wunderlin,
1998), southern Georgia (Bryson et al., 1996), south-
ern Alabama (Mohr, 1901; Lelong, 1988), Louisiana
(Thomas & Allen, 1993), and coastal Texas (Correll
& Johnston, 1970; Hatch et al., 1990; Jones et al.,
1997). Oxycaryum cubense has spirally arranged
scales and has been treated as Scirpus cubensis
Poepp. & Kunth (e.g., Correll & Johnston, 1970;
Godfrey & Wooten, 1979); however, its habit and
embryo resemble Cyperus (van der Veken, 1965;
Lye, 1971), and its taxonomic placement has been
disputed: Cypereae (Lye, 1971) and Scirpeae (Bruhl,
1995). The molecular analysis of Muasya et al.
(2002) supports classification of Oxycaryum in
Cypereae. Two forms differing only in gross inflo-
rescence features occur throughout the range of the
species. Plants with umbellate inflorescences are
called O. cubense f. cubense, while those with
monocephalous inflorescences are called O. cubense
f. paraguayense (Maury) Pedersen (Barros, 1960;
Pedersen, 1995). This aquatic species forms exten-
sive floating rafts in ponds, lakes, ditches, or
impounded swamps in the southeastern U.S.A. and
elsewhere (Haines & Lye, 1983; Bryson et al., 1996).
Oxycaryum cubense is one of the most vigorous
plants (along with Salvinia molesta D. S. Mitch. and
Pistia stratiotes L.) in forming sudds in African lakes
(Holm et al., 1977), thereby impeding navigation. In
the southeastern U.S.A., O. cubense appears to be
invasive,with floating mats covering large areas to
the exclusion of other aquatic vegetation (Bryson et
al., 1996); however, its sporadic distribution in the
U.S.A. suggests low fertility of achenes. Its corky
buoyant achenes are adapted to dispersal by moving
water, and its mat-forming, floating habit facilitates
asexual reproduction and transport of vegetative
fragments by moving water (Haines & Lye, 1983).
Oxycaryum cubense has been in the southeastern
U.S.A. for more than a century (Chapman, 1889;
Mohr, 1901), and we suspect that it was dispersed
into North America from the West Indies or South
America by migratory birds or with ballast. In order
to understand better its dispersal and potential to

The Significance of Cyperaceae as Weeds 51
invade wetland habitats, additional research into its
reproductive biology is needed to determine the
extent to which O. cubense reproduces sexually and
spreads from achenes.
RHYNCHOSPORA
Rhynchospora is a cosmopolitan genus of more
than 250 species, most of which inhabit wet, acidic
soils (Kral, 2002e). Rhynchospora spp. are of little
economic importance as weeds, and 20 species are
listed in Appendix 2. Although most Rhynchospora
spp. considered to be weeds are only secondarily or
occasionally so, R. corymbosa (L.) Britton, R.
holoschoenoides (Rich.) Herter, R. submarginata
Kük., and R. wightiana (Nees) Steud. are cited as
weeds of rice agriculture in the Eastern Hemisphere
(Kern, 1974; Simpson & Inglis, 2001). In the U.S.A.,
R. corniculata (Lam.) A. Gray and R. globularis
(Chapm.) Small are occasionally weeds (WSSA,
1989) along ground transportation routes but usually
do not cause economic losses, and dense stands of
the caespitose perennial, R. corniculata, along
waterways impede flow in canals associated with
rice production and can cause unwanted flooding of
agricultural fields. Several species related to R. cor-
niculata are sometimes locally abundant in roadside
ditches in the Coastal Plain of the eastern U.S.A.
Rhynchospora macrostachya Torr. ex A. Gray and R.
corniculata are found in hydric soils in a variety of
wetland habitats, including roadside ditches and
margins of artificial ponds, and both are caespitose
perennials of wide distribution in eastern North
America (Kral, 2002e). The related species, R. inun-
data Fernald and R. careyana Fernald, are emergent
rhizomatous perennials that form extensive stands in
shallow depressions in the flatwoods, including
roadside ditches (Kral, 2002e). Although only R.
corniculata is listed as a weed, we suspect that R.
careyana,R. inundata, and R. macrostachya might
be invasive, if introduced into similar habitats out-
side of their natural ranges. Rhynchospora caduca
Elliott, of little value as forage for livestock, is some-
times a weed in poorly maintained pastures in the
southeastern U.S.A. where it is native (Bryson, pers.
obs.) and is recently introduced and spreading rapid-
ly in Hawaii (Wagner et al., 1990; Wagner & Herbst,
1995). Rhynchospora globularis,another native of
the continental U.S.A., was collected in 1982 as an
introduction in Hawaii (Wagner et al., 1990) and also
occurs in northern California (Cranfil, 1993) where
it is perhaps introduced from the eastern U.S.A.
Other Rhynchospora spp. that opportunistically
spread into artificially disturbed sites within their
native ranges in the southeastern U.S.A. include R.
cephalantha A. Gray, R. debilis Gale, R. fascicularis
(Michx.) Vahl, R. glomerata (L.) Vahl, R. fernaldii
Gale, R. inexpansa (Michx.) Vahl, R. microcephala
(Britton) Britton ex Small, R. odorata C. Wright ex
Griseb., and R. torreyana A. Gray (Godfrey &
Wooten, 1979; Bryson & Carter, pers. obs.). We sus-
pect that such plants would likely become invasive if
introduced into suitable habitats elsewhere, as R.
caduca has in Hawaii.
SCHOENOPLECTUS
Schoenoplectus is a genus of 77 species world-
wide (Smith, 2002b), of which 20 are cited as weeds
in Appendix 2. Schoenoplectus mucronatus (L.)
Palla [= Scirpus mucronatus L.], considered to be
among the world’s worst weeds (Holm et al., 1997),
is a pest in rice and other row and tree crops in
Bangladesh, France, India, Malaysia, the
Philippines, Portugal, Spain, and the U.S.A. (Holm
et al., 1997). Schoenoplectus mucronatus is a greater
problem in paddy fields where hand labor is the pri-
mary method of weed control than in rice production
involving mechanical tillage and the use of herbi-
cides. Schoenoplectus grossus (L. f.) Palla [= Scirpus
grossus L. f.] is a weed of rice, riverbeds, reservoirs,
and irrigation systems in southeastern Asia including
regions of Vietnam, India, and the Philippines, and S.
tabernaemontani is also listed as a weed of rice in
China (Zhirong et al., 1990). Schoenoplectus jun-
coides (Roxb.) Palla is reportedly naturalized in rice
fields in Europe (DeFilipps, 1980a). Schoenoplectus
acutus and S. americanus (Pers.) Volkart ex Schinz
& R. Keller are weeds in wetland areas of North
America (WSSA, 1989; Callahan et al., 1995), while
S. californicus (C. A. Mey.) Soják is reported as a
weed in North America and Brazil (WSSA, 1989;
Kissmann, 1997).
SCIRPODENDRON
Scirpodendron Zipp. ex Kurz is a genus of two
species ranging from Sri Lanka and southeastern
Asia through Malesia to Australia and Polynesia
(Goetghebeur, 1998). Scirpodendron inhabits fresh-

52 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
water tidal swamps, tidal swamp forests, and forests
adjacent to mangroves, and its large fruits are dis-
persed by water (Kern, 1974). It is cultivated in
Sumatra for its leaves, which are used for thatching
and weaving mats and hats (Kern, 1974).
Scirpodendron ghaeri (Gaertn.) Merr. has been cited
as a weed of rice fields in Asia (Moody, 1989).
SCIRPUS
Scirpus is a genus of 35 species widely distrib-
uted in North America, Mexico, Eurasia, Australia,
and the Pacific Islands (Whittemore & Schuyler,
2002). Eight species are listed as weeds in Appendix
2, none of which is invasive in agricultural crop-
lands. Scirpus atrovirens Willd., S. pendulus Muhl.,
and S. cyperinus (L.) Kunth are native to North
America and cited as weeds there (WSSA, 1989;
Callahan et al., 1995). These Scirpus species are
occasional weeds along roadsides and waterways
and in wet pastureland but rarely cause economic
losses. Scirpus atrovirens and S. pendulus are natu-
ralized in Europe (DeFilipps, 1980a). In the U.S.A.,
where it is native, S. cyperinus sometimes forms
extensive stands dominating disturbed wetlands
(Carter, pers. obs.), and we strongly suspect it would
be an invasive pest if introduced into suitable habi-
tats outside its natural range.
SCLERIA
Scleria is widely distributed in tropical and sub-
tropical regions around the world and consists of ca.
200 species (Reznicek et al., 2002). As shown in
Appendix 2, 24 species are weeds, a number of
which are aquatics and known or potential weeds of
rice agriculture (e.g., Scleria biflora Roxb., S. laevis
Retz., S. lithosperma (L.) Sw., S. novae-hollandiae
Boeckeler). The non-native invasive weed S. lacus-
tris C. Wright has been found in freshwater marshes
of peninsular Florida, U.S.A., where it can be local-
ly abundant and dominant in water up to 1 m deep,
forming dense stands and displacing native vegeta-
tion (Tobe et al., 1998; Wunderlin, 1998; Jacono,
2001). Scleria lacustris seems to require recession of
standing water in order to become established
(Jacono, 2001). It is thought to be native in scattered
areas of the Neotropics, Africa, and Madagascar
(Core, 1933; Hennessy, 1985) and is known from
Brazil, Cuba, Costa Rica, French Guiana, Guyana,
Jamaica, Paraguay, Suriname, U.S.A., and six coun-
tries across tropical Africa (Jacono, 2001).
Additional research is needed to determine the eco-
logical range of S. lacustris and control strategies.
Scleria vaginata Steud. is an aggressive vine native
to Central and South America that was collected
once in southern Florida, U.S.A. (Reznicek et al.,
2002), and we suspect it could be invasive if intro-
duced into tropical and subtropical areas outside its
native range.
DISCUSSION
Cyperaceae is a large, diverse, cosmopolitan
family, and many of its species are biologically pre-
disposed to spread opportunistically into areas
altered by humans. Data compiled in Appendix 1
show that humans have played a tremendous role in
the dispersal of sedges, including many weeds.
Given the fundamental importance of dispersal and
habitat disturbance in the evolution and survival of
weeds and their intrinsic attributes favoring competi-
tion, colonization, and migration, it is not surprising
that many sedges have evolved and continue to
evolve as weeds. The magnitude of the human “foot-
print” on Earth is immense. Given the role that
humans play in destruction and conversion of natural
areas into disturbed and highly artificial ruderal
habitats and urban and agricultural systems, it is
axiomatic that the numbers of noxious weeds and
invasive plant species will increase in step with the
human population.
It is difficult to anticipate which species will
become weeds, and where and under what circum-
stances they will be invasive. Rhynchospora caduca,
a seemingly innocuous sedge native to the southeast-
ern U.S.A., has recently been reported as an invasive
weed in Hawaii (Wagner & Herbst, 1995).
Rhynchospora caduca is not extraordinary among
the beak-rushes in the southeastern U.S.A., which
suggests that any number of apparently harmless
species could pose similar problems in an alien envi-
ronment. Insular systems, such as the Hawaiian
Islands, have great potential as natural laboratories
for the study of invasion.
Appendix 2 is a list of 447 species of Cyperaceae
cited as weeds, which was compiled from more than
60 publications. Most cyperaceous weeds are from
tropical and subtropical regions, and the most trou-

The Significance of Cyperaceae as Weeds 53
blesome sedges (Cyperus rotundus,C. esculentus,C.
difformis, and C. iria) are native to Asia and Africa
but are now widely dispersed on other continents.
In order to examine the impact of humans on dis-
persal and introduction of cyperaceous weeds, we
noted commonalities in listings of species in
Appendices 1 and 2 and used these data to construct
Table 4. Thus, Table 4 shows the number of weed
species in each genus that are known or suspected to
have been anthropogenically dispersed. When Appen-
dices 1 and 2 are compared, 111 species are common
to both lists (Table 4) with the greatest number of
cyperaceous weeds known or suspected to be
dispersed by humans in Cyperus (43 spp., ca. 39%),
followed by Carex (24 spp., ca. 22%); Eleocharis
(9 spp., ca. 8%); Fimbristylis (8 spp., 7%); Kyllinga
and Schoenoplectus (6 spp. each, 5%); Scirpus (5 spp.
each, ca. 5%); Rhynchospora (3 spp., ca. 3%);
Fuirena (2 spp., ca. 2%); and Bolboschoenus, Bulbo-
stylis,Lepironia,Lipocarpha,and Mapania (1 spp.
each, ca. 1%).
Cyperus, by far, has been subject to greater
anthropogenic dispersal than the other cyperaceous
genera, which undoubtedly has been an important
factor. It is readily concluded from Appendix 1 that
Cyperus spp. have been mostly introduced uninten-
tionally through a variety of human activities, espe-
cially as contaminants of seeds (particularly rice),
wool, and dumping of ship’s ballast. It seems reason-
able to conclude from these data that systematic sur-
veys of flora in vicinity of ports of entry are needed
for early detection of new introductions and reintro-
ductions and to understand better the dynamics of
inadvertent importation of noxious weeds.
The role of rice agriculture in the introduction
of cyperaceous weeds has long been recognized and
is reinforced by data presented in Appendix 1. The
number of cyperaceous weeds associated with rice
agriculture in Appendix 2 is great, and, despite
advancements in the regulation of importation of
grain, there still exists the possibility of unintention-
al movement and introduction of other potentially
noxious sedges as contaminants in shipments of
seeds. These data indicate the need for continued
vigilance and regulation of movement and importa-
tion of sedges throughout the world.
Historically,Carexspp. have received little
attention as agricultural weeds. However, Kukkonen
(2001) includes rice fields in Pakistan as habitats of
Carex diandra,C. pycnostachya, and C. divisa
Huds., and he describes C. songorica Kar. & Kir., C.
diluta M. Bieb., and C. orbicularis Boott as growing
in irrigation channels. The latter set of species is not
listed in Appendix 2, which includes only entries
explicitly characterized as weeds or invasives or
directly associated with agricultural fields, gardens,
or turf. However, populations of native sedges that
have spread into irrigation canals associated with
agriculture have certainly adapted to human distur-
bance, and biotypes adapted to conditions in the
adjacent fields could easily evolve.
The large number of ornamental and cultivated
sedges (>150 spp.) listed in Appendix 1 was not
anticipated. Of particular interest is the increasing
horticultural usage of sedges, especially Carex spp.,
as ornamentals (Figs. 2 and 3). This indicates a need
for increased research into the reproductive biology,
physiology, and growth characteristics of ornamental
sedges to determine which species may be safely
used and where and which will likely become inva-
sive. There is also a need for greater awareness about
problems inherent in the unwise and irresponsible
use of ornamental sedges and additional measures
toward intervention to prevent the transportation and
importation of ornamental sedges.
Because of their distributions across vast latitu-
dinal, altitudinal, and climatic ranges and diverse
habitats, populations of widely distributed weeds
have been subject to a great array of environmental
factors resulting in much localized natural (and arti-
ficial) selection and diversification. Thus, in general,
the taxonomy of weeds is far more complex than of
other plants, which is evident in the complex nomen-
clature of the most widely distributed weeds, e.g.,
Cyperus esculentus,C. rotundus (Haines & Lye,
1983), C. polystachyos (cf. Kükenthal, 1935–1936),
and C. sanguinolentus (cf. Kükenthal, 1935–1936;
Kern, 1974). To resolve basic questions about rela-
tionships within these taxa, there is a great need for
additional morphometric, field-, and herbarium-
based research into the variation and taxonomy on a
worldwide basis. The increased use of molecular
techniques (e.g., Muasya et al., 2000a, b, 2002)
should help to stabilize nomenclature by resolving
the taxonomic status and rank of certain disputed
groups, e.g., the segregates of Cyperus and Scirpus.
In the future,the results of molecular research will
elucidate much about the pathways of introduction

54 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
and migration of invasive weeds. Introduction of new
weeds is increasingly a problem because of the fre-
quency and ease of long-distance and international
transportation, and advances in basic research will
result in molecular assays useful in detecting and
stopping weeds at ports of entry and in more accu-
rately diagnosing infestations of herbicide-resistant
biotypes of weeds.
Given the economics of weed control, including
indirect costs (e.g., increased cost of health care, reme-
diation of environmental damage), every precaution
should be taken to avoid tagging indigenous plants as
weeds without compelling supportive evidence.
Realistically and pragmatically, it is most certainly
advantageous and desirable for native plants to occupy
roadsides and other artificial habitats than alien weeds.
There is a great need for basic research to determine
the ecological tolerances and invasive potentials and
limits of indigenous and nonindigenous weeds. For
only through the results of such research will basic
knowledge be advanced sufficiently to allow applied
scientists, natural resource managers, and the public to
make informed, intelligent decisions about which
plants to promote,which to exclude, which to sup-
press, and when to suppress them.
LITERATURE CITED
Abad, P., B. Pascual, J. V. Maroto, S. Lopez-Galarza, A.
Sanbautista & J. Alagarda. 2000. Chufa (Cyperus
esculentus L. var. sativus Boeck.): An unconven-
tional crop. Studies related to application and cul-
tivation. Econ. Bot. 54: 439–448.
Adams, C. D. 1994. Cyperaceae. Pp. 262–485 in G.
Davidse, M. Sousa S. & A. O. Chater (editors),
Flora Mesoamericana, Vol. 6: Alismataceae a
Cyperaceae. Universidad Nacional Autónoma de
México, México, D.F.; Missouri Botanical Garden,
St. Louis; The Natural History Museum, London.
Akobundu, J. O. & C. W. Agyakwa. 1987. A Handbook
of West Africa Weeds. International Institute of
Tropical Agriculture, Ibadan, Nigeria.
Aleixo, M. de F. D. & I. F. M. Valio. 1976. Effect of
light, temperature and endogenous growth regula-
tors on the growth of buds of Cyperus rotundus L.
tubers. Z. Pflanzenphysiol. 80: 336–347.
Allan, M. 1978. Weeds: The Unbidden Guests in Our
Gardens. Viking Press, New York.
Baijnath, H. 1975. A study of Cyperus alternifolius L.,
sens. lat. (Cyperaceae). Kew Bull. 30: 521–526.
Bailey, L. H. 1935. The Standard Cyclopedia of
Horticulture, Vols. 1–3. The Macmillan Company,
New York.
Species Percent
Genus (incl. infrasp.) of Total
Cyperus243 39
Carex 24 22
Eleocharis 98
Fimbristylis 87
Kyllinga 65
Schoenoplectus 65
Scirpus 5 5
Rhynchospora 33
Fuirena 2 1
Bolboschoenus 11
Bulbostylis 11
Lepironia 11
Lipocarpha 11
Mapania 11
Abildgaardia 00
Caustis 0 0
Cladium 00
Courtoisina 0 0
Cymophyllus 00
Desmoschoenus 00
Eriophorum 00
Gahnia 00
Isolepis 00
Kobresia 00
Lepidosperma 0 0
Machaerina 00
Oreobolus 00
Oxycaryum 00
Schoenus 00
Scleria 00
Trichophorum 00
Uncinia 00
Total 111 ~100
Table 4. Numbers and percentages of species of
Cyperaceae by genus, which have been cited as weeds
and are known or suspected to be dispersed by humans
(data extracted from Appendices 1 and 2).1
1Authority names for genera in Table 4 not discussed
elsewhere in this paper are as follows: Caustis R. Br.;
Desmoschoenus Hook f.; Kobresia Willd.; Oreobolus R. Br.;
Schoenus L.; Trichophorum Pers.
2Includes Diclidium, Juncellus, Mariscus, Pycreus, and
Queenslandiella.

The Significance of Cyperaceae as Weeds 55
—. 1949. Manual of Cultivated Plants. Macmillan
Publishing Company, New York.
— & E. Z. Bailey. 1976. Hortus Third. Macmillan
Publishing Company, New York.
Baker, H. G. 1965. Characteristics and modes of origin
of weeds. Pp. 147–172 in H. G. Baker & G. L.
Stebbins (editors), The Genetics of Colonizing
Species. Academic Press, New York.
—. 1974. The evolution of weeds. Annual Rev. Ecol.
Evol. Syst. 5: 1–24.
Ball, P. W. & A. A. Reznicek. 2002. Carex. Pp. 254–273
in Flora of North America Editorial Committee
(editors), Flora of North America North of
Mexico, Vol. 23: Magnoliophyta: Commelinidae
(in part): Cyperaceae. Oxford Univ. Press, New
York.
—, — & D. F. Murray. 2002. Cyperaceae. Pp. 3–608 in
Flora of North America Editorial Committee (edi-
tors), Flora of North America North of Mexico,
Vol. 23: Magnoliophyta: Commelinidae (in part):
Cyperaceae. Oxford Univ. Press, New York.
Barrentine, W. L., C. E. Snipes & R. S. Baker. 1992.
Herbicide application technology. Pp. 439–514 in
C. G. McWhorter & J. R. Abernathy (editors),
Weeds of Cotton: Characterization and Control.
The Cotton Foundation, Memphis.
Barros, M. 1960. Las Cyperaceas del estado de Santa
Catalina. Sellowia 12: 181–450.
Baskin, J. M. & C. C. Baskin. 1978. A discussion of the
growth and competitive ability of C3and C4
plants. Castanea 43: 71–76.
Bayer AG. 1992. Important Crops of the World and
Their Weeds, 2nd ed. Business Group Crop
Protection, Bayer AG, Leverkusen, Germany.
Beal, E. O. 1977. A Manual of Marsh and Aquatic
Vascular Plants of North Carolina with Habitat
Data. Tech. Bull. 247. North Carolina Agricultural
Research Service, Raleigh.
Bellue, M. K. 1932. Weeds of California seed rice.
Monthly Bull. Calif. Dept. Agric. 21: 290–296.
Bendixen, L. E. & U. B. Nandihalli. 1987. Worldwide
distribution of purple and yellow nutsedge
(Cyperus rotundus and C. esculentus). Weed
Technol. 1: 61–65.
Benl, G. 1937. Eigenartige verbreitungseinrichtungen
bei der Cyperaceengattung Gahnia. Flora 131:
369–386.
Bernard, J. M. & F. K. Seischab. 1994. Life history of
shoots of Carex comosa F. Boott. Rhodora 96:
179–189.
Bertels, A. 1957. Contribuição ao conhecimento das
Ciperaceas de Pelotas (RS). Inst. Agronomico do
Sol. Bol. Tecn. 17: 22–28.
Black, C. C., T. M. Chen & R. H. Brown. 1969.
Biochemical basis for plant competition. Weed
Sci. 17: 338–344.
Boyette, C. D. 2000. The bioherbicide approach: Using
phytopathogens to control weeds. Pp. 134–152 in
A. H. Cobb & R. C. Kirkwood (editors),
Herbicides and Their Mechanisms of Action.
Sheffield Academic Press, Sheffield, United
Kingdom.
Brickell, C. & J. D. Zuk (editors). 1997. The American
Horticultural Society A–Z Encyclopedia of
Garden Plants, 1st ed. DK Publishing, Inc., New
York.
Britton, N. L. 1886. A preliminary list of N. A. species
of Cyperus, with descriptions of new forms. Bull.
Torrey Bot. Club 13: 205–216.
Brooklyn Botanical Garden. 1988. Ornamental Grasses.
Brooklyn Botanical Gardens, Inc., New York.
Brown, A. 1880. Ballast plants in New York City. Bull.
Torrey Bot. Club 7: 122–126.
Brown, A. D. H. & B. E. Marshall. 1981. Evolutionary
changes accompanying colonization in plants. Pp.
351–363 in G. C. E. Scudder & J. L. Reveal (edi-
tors), Evolution Today. Carnegie-Mellon Univ.,
Pittsburgh.
Brown, W. V. 1975. Variations in anatomy, associations,
and origins in kranz tissue. Amer. J. Bot. 62:
395–402.
Browning, J., K. D. Gordon-Gray & S. G. Smith. 1995.
Achene structure and taxonomy of North
American Bolboschoenus (Cyperaceae). Brittonia
47: 433–445.
Bruhl, J. 1993. Sedge Genera of the World, Vers. 2.01.
Intkey, Interactive Dataset. N.C.W. Beadle
Herbarium, Univ. of New England, Armidale.
—. 1995. Sedge genera of the world: Relationships and
a new classification of the Cyperaceae. Austral.
Syst. Bot. 8: 125–305.
—. 2002. Oxycaryum. P. 140 in Flora of North America
Editorial Committee (editors), Flora of North
America North of Mexico, Vol. 23: Magnoliophyta:
Commelinidae (in part): Cyperaceae. Oxford Univ.
Press, New York.
Bruton, D. F. & P. M. Catling. 1982. The slender sedge
(Carex praegracilis) new to Ottawa district: An
example of contemporary plant migration. Trail &
Landscape 16: 152–157.
Bryson, C. T. 1984. Weed alert: Smallflower umbrella
sedge (Cyperus difformis L.). S. Weed Sci. Soc.
Newslett. 7: 6.
—. 1985a. Carex weeds of Mississippi. J. Mississippi
Acad. Sci. 30: 10.
—. 1985b. Cyperus weeds of Mississippi cotton fields.
J. Mississippi Acad. Sci. 30: 10.
—. 1994. Effects of tillage and herbicide input levels on
purple nutsedge (Cyperus rotundus L.) control in
cotton. Proc. Weed Sci. Soc. Amer. 34: 75.
—. 1997. Costs and effects of tillage and herbicide
inputs for purple nutsedge (Cyperus rotundus)

56 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
control in cotton. Proc. Weed Sci. Soc. Amer.
37: 94.
—. 2003. Weed species. Pp. 1571–1588 in J. Plimmer
(editor), Encyclopedia Agrochemicals,Vol. 3. John
Wiley & Sons, New York.
— & R. Carter. 1992. Notes on Cyperus and Kyllinga
(Cyperaceae) in Mississippi with records of six
species new to the state. Sida 15: 119–124.
— & —. 1994. Notes on Carex,Cyperus, and Kyllinga
(Cyperaceae) in Mississippi with records of eight
species previously unreported to the state. Sida 16:
171–182.
— & —. 1995. Introduction and spread of weedy
Cyperus species in the United States. Proc. Weed
Sci. Soc. Amer. 35: 36.
— & —. 2004. Biology of pathways for invasive weeds.
Weed Technol. 18: 1216–1220.
— & J. E. Hanks. 1993. Effects of spray droplet size on
control of purple nutsedge (Cyperus rotundus).
Proc. S. Weed Sci. Soc. 46: 334.
— & —. 2001. Weed shifts in Deep Hollow watershed,
Leflore County, MS. Pp. 86–90 in R. A. Rebich &
S. Knight (editors), The Mississippi Delta
Management Systems Evaluation Area Project,
1995–1999. Mississippi Agricultural & Forestry
Experiment Station Bulletin 377, Mississippi State.
— & P. E. Keeley. 1992. Reduced tillage systems. Pp.
323–363 in C. G. McWhorter & J. R. Abernathy
(editors), Weeds of Cotton: Characterization and
Control. The Cotton Foundation, Memphis.
— & R. F. C. Naczi. 2002. Carex section Laxiflorae.
Pp. 431–440 in Flora of North America Editorial
Committee (editors), Flora of North America
North of Mexico, Vol. 23: Magnoliophyta:
Commelinidae (in part): Cyperaceae. Oxford Univ.
Press, New York.
— & G. D. Wills. 1991. Purple nutsedge (Cyperus
rotundus) control with low-volume technology in
cotton. Proc. S. Weed Sci. Soc. 40: 404.
—, — & P. C. Quimby Jr. 1990. Low volume applica-
tion of herbicides in an invert emulsion for purple
nutsedge (Cyperus rotundus) control. Weed
Technol. 4: 907–909.
—, R. F. C. Naczi & S. McDaniel. 1992a. Notes on
noteworthy records of Carex (Cyperaceae) from
the southeastern United States. Sida 15: 125–135.
—, G. D. Wills & J. E. Hanks. 1992b. Purple nutsedge
(Cyperus rotundus L.) control in cotton with low-
volume oil application technology. Proc. Weed
Sci. Soc. Amer. 32: 78.
—, J. E. Hanks & G. D. Wills. 1994a. Nutsedge
(Cyperus rotundus) control in reduced-tillage cot-
ton production with low-volume technology. Weed
Technol. 8: 28–31.
—, J. R. MacDonald & R. Warren. 1994b. Notes on
Carex (Cyperaceae), with C. godfreyi new to
Alabama and C. communis and C. scoparia new to
Mississippi. Sida 16: 355–361.
—, —, R. Carter & S. D. Jones. 1996. Noteworthy
Carex,Cyperus,Eleocharis,Kyllinga,and
Oxycaryum (Cyperaceae) from Alabama, Arkansas,
Georgia, Louisiana, Mississippi, North Carolina,
Tennessee, and Texas. Sida 17: 501–518.
—, R. Carter, L. B. McCarty & F. H. Yelverton. 1997.
Kyllinga, a genus of neglected weeds in the conti-
nental United States. Weed Technol. 11: 838–842.
—, N. C. Coile & J. H. Rettig. 1998. Friend or foe:
Identifying Cyperus. Palmetto 18: 16–19.
—, C. Salisbury & W. B. McCloskey. 1999. Weeds and
their control. Pp. 617–658 in C. W. Smith & J. T.
Cothren (editors), Cotton: Origin, History,
Technology and Production. John Wiley & Sons,
New York.
—, K. N. Reddy & W. T. Molin. 2002. Purple nutsedge
population dynamics in narrow row transgenic cot-
ton and soybean. Proc. S. Weed Sci. Soc. 55:
169–170.
—, R. Carter & D. J. Rosen. 2003a. Deeprooted Sedge
(Cyperus entrerianus). Proc. S.Weed Sci.Soc. 56: 370.
—, K. N. Reddy & W. T. Molin. 2003b. Purple nutsedge
(Cyperus rotundus) population dynamics in nar-
row row transgenic cotton (Gossypium hirsutum)
and soybean (Glycine max). Weed Technol. 17:
805–810.
Buchanan, G. A. 1992. Trends in weed control methods.
Pp. 47–72 in C. G. McWhorter & J. R. Abernathy
(editors), Weeds of Cotton: Characterization and
Control. The Cotton Foundation, Memphis.
Buchholtz, K. P. 1967. Definitions of terms used in
weed control. Weeds 15: 388–389.
Burkhalter, J. R. 1985. Aletris farinosa,Cyperus dif-
formis, and Cyperus pilosus new for Florida. Sida
11: 247–248.
Burkill, H. M. 1985. The Useful Plants of West Tropical
Africa, Vol. 1, 2nd ed. Royal Botanical Gardens, Kew.
Burkill, I. 1935. A Dictionary of the Economic Products
of the Malay Peninsula, Vols. 1 & 2. Governments of
the Straits Settlements and Federated Malay States,
Crown Agents for the Colonies, London.
Burns, E. R. & G. A. Buchanan. 1967. Controlling wolf-
tail (Carex cherokeensis) in pastures. Highlights
Agric. Res. Alabama Agric. Exp. Sta. 14: 4.
—, — & A. E. Hiltbold. 1969. Absorption and translo-
cation of 2,4-D by wolftail (Carex cherokeensis
Schwein.). Weed Sci. 17: 401–404.
Burr, R. J. & G. F. Warren. 1971. Enhancement of her-
bicide activity with an isoparaffinic oil carrier.
Weed Sci. 19: 701–705.
— & —. 1972. An oil carrier for increasing purple
nutsedge control. Weed Sci. 20: 324–327.

The Significance of Cyperaceae as Weeds 57
Byrd, J. D., Jr. 1995a. Report of the 1994 cotton weed
loss committee. Proc. Beltwide Cotton Conf.,
Cotton Weed Sci. Res. Conf. 18: 595–598.
—. 1995b. Mississippi cotton weed survey. Proc.
Beltwide Cotton Conf., Cotton Weed Sci. Res.
Conf. 18: 598–599.
Callahan, R. H., S. L. Carson & R. T. Dobbins. 1995.
NAWEEDS, Computer-aided Weed Identification
for North America. Illustrated User’s Guide plus
Computer Floppy Disk. Moscow, Idaho.
Cardenas, J., C. E. Reyes & J. D. Doll. 1972. Tropical
Weeds Maleza Tropicales, Vol. 1. Instituto
Colombiano Agropecuario, Bogota.
Carter, R. 1984. A Systematic Study of the New World
Species of Section Umbellati of Cyperus. Ph.D.
Dissertation, Vanderbilt Univ., Nashville.
—. 1990. Cyperus entrerianus (Cyperaceae), an over-
looked species in temperate North America. Sida
14: 69–77.
—. 1993. Animal dispersal in the North American
sedge, Cyperus plukenetii (Cyperaceae). Amer.
Midl. Naturalist 129: 352–356.
—. 2003. What to do with an unknown specimen:
Preparation and storage of vouchers. Proc. S.
Weed Sci. Soc. 56: 372.
—. 2005. An introduction to the sedges of Georgia.
Tipularia 20: 15–44.
— & C. T. Bryson. 1991a. A report of Cyperus
grayioides and Cyperus retroflexus (Cyperaceae)
new to Missouri and notes on other selected
Missouri Cyperus. Sida 14: 475–481.
— & —. 1991b. A second station and new county
record for Cyperus retroflexus (Cyperaceae) in
Missouri. Missouriensis 12: 33.
— & —. 1996. Cyperus entrerianus:A little known
aggressive sedge in the southeastern United States.
Weed Technol. 10: 232–235.
— & —. 2000a. Taxonomy of weedy Cyperus species.
P. 47 in Third International Weed Science
Congress, Abstracts. Foz do Iguaçu, Brazil.
— & —. 2000b. Notes on the distribution, ecology, and
taxonomy of Cyperus sanguinolentus (Cyperaceae)
new to the southeastern United States, and its
relation to the supposed endemic Cyperus louisia-
nensis. Sida 19: 325–343.
— & —. 2001. Bloodscale sedge (Cyperus sanguino-
lentus), a new weed in the United States. Proc.
Weed Sci. Soc. Amer. 41: 93.
— & C. E. Jarvis. 1986. Re-evaluation and lectotypifi-
cation of Scirpus echinatus L. Rhodora 88:
451–456.
— & S. D. Jones. 1991. Additional records of Cyperus
entrerianus (Cyperaceae) in the United States.
Sida 14: 615–616.
— & —. 1997. Notes on the Cyperus retroflexus com-
plex (Cyperaceae) with three nomenclatural pro-
posals. Rhodora 99: 319–334.
— & R. Kral. 1990. Cyperus echinatus and Cyperus
croceus, the correct names for North American
Cyperus ovularis and Cyperus globulosus. Taxon
39: 322–327.
— & R. L. Mears. 2000. Cyperus (subg.
Queenslandiella) hyalinus (Cyperaceae) new to
the United States and the western hemisphere.
Sida 19: 345–350.
—, C. T. Bryson & B. L. Lipscomb. 1987. Cyperus uni-
florus (Cyperaceae) east of the Mississippi River.
Sida 12: 250.
—, M. W. Morris & C. T. Bryson. 1990. Some rare or
otherwise interesting vascular plants from the
Delta Region of Mississippi. Castanea 55: 40–55.
—, R. L. Mears, K. C. Burks & C. T. Bryson. 1996. A
report of four exotic Cyperus species new to
Florida, U.S.A. Sida 17: 275–281.
—, D. K. Alexander, C. T. Bryson & A. Lazari. 1999.
The taxonomy of Cyperus virens and Cyperus
drummondii (Cyperaceae) in the southeastern
United States. Sida 18: 1049–1063.
Champlin, R. L. 1994. Notes on the Rhode Island flora.
Rhodora 96: 102–103.
Chandler, J. M. & F. T. Cooke. 1992. Economics of cot-
ton losses caused by weeds. Pp. 85–116 in C. G.
McWhorter & J. R. Abernathy (editors), Weeds of
Cotton: Characterization and Control. The Cotton
Foundation, Memphis.
—, A. S. Hamill & A. G. Thomas. 1984. Crop Losses
Due to Weeds in Canada and the United States.
Weed Science Society of America, Champaign,
Illinois.
Chapman, A. W. 1889. Flora of the Southern States, 2nd
ed. Ivison, Blakeman & Company, New York.
Chase, M. W., D. E. Soltis, P. S. Soltis, P. J. Rudall, M.
F. Fay, W. H. Hahn, S. Sullivan, J. Joseph, M.
Molvray, P. J. Kores, T. J. Givnish, K. J. Sytsma &
J. C. Pires. 2000. Higher-level systematics of the
monocotyledons: An assessment of current knowl-
edge and a new classification. Pp. 3–16 in K. L.
Wilson & D. A. Morrison (editors), Monocots:
Systematics and Evolution. CSIRO Publishing,
Collingwood, Victoria, Australia.
Chermezon, H. 1924. Sur la dissemination de quelques
Cyperacees. Bull. Soc. Bot. France 71: 849–861.
Clarke, C. B. 1884. On the Indian species of Cyperus. J.
Linn. Soc., Bot. 21: 1–2002.
—. 1894. Cyperaceae. Pp. 585–672 in J. D. Hooker
(editor), The Flora of British India, Vol. VI, Pt.
XIX. L. Reeve & Co., London.
—. 1900. Cyperaceae. Pp. 8–160 in I. Urban (editor),
Symbollae Antillanae, Vol. 2. Fratres Borntraeger,
Leipzig.

58 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
—. 1908. New genera and species of Cyperaceae. Bull.
Misc. Inform. Kew 8: 1–196.
Clewell, A. F. 1985. Guide to the Vascular Plants of the
Florida Panhandle. Univ. Presses of Florida,
Tallahassee.
Coble, H. D. & J. D. Byrd. 1992. Interference of weeds
in cotton. Pp. 73–84 in C. G. McWhorter & J. R.
Abernathy (editors), Weeds of Cotton: Character-
ization and Control. The Cotton Foundation,
Memphis.
Core, E. L. 1933. The American species of Scleria.
Brittonia 2: 1–105.
Correll, D. S. & M. C. Johnston. 1970. Manual of the
Vascular Plants of Texas. Texas Research
Foundation, Renner.
Cranfill, R. 1993. Rhynchospora. P. 1144 in J. C.
Hickman (editor), The Jepson Manual: Higher
Plants of California. Univ. of California Press,
Berkeley.
Cronk, Q. C. B. & J. L. Fuller. 1995. Plant Invaders.
Chapman & Hall, London.
Cronquist, A. 1981. An Integrated System of
Classification of Flowering Plants. Columbia
Univ. Press, New York.
Cusick, A. W. 1984. Carex praegracilis: A halophytic
sedge naturalized in Ohio. Michigan Botanist 23:
103–106.
Daehler, C. C. & D. R. Strong. 1997. Hybridization
between introduced smooth cordgrass (Spartina
alterniflora: Poaceae) and native California cord-
grass (S. foliosa) in San Francisco Bay, California,
USA. Amer. J. Bot. 84: 607–611.
Darby, W. J., P. Ghaliougui & L. Grivetii. 1977. Food:
The Gift of Osiris, Vol. 2. Academic Press,
London.
Darke, R. 1999. The Color Encyclopedia of Ornamental
Grasses: Sedges, Rushes, Restios, Cattails, and
Selected Bamboos. Timber Press, Portland.
Datta, S. & A. Benerjee. 1976. The weight and number
of seeds produced by rice-field weeds. P. A. N. S.
22: 257–263.
De Datta, S. & H. Jereza. 1976. The use of cropping
systems and land and water management to shift
weed species. Philipp. J. Crop Sci. 1: 173–178.
DeFelice, M. S. 2002. Yellow nutsedge Cyperus escu-
lentus L.—Snack food of the gods. Weed Technol.
16: 901–907.
— & C. T. Bryson. 2004. Interactive Encyclopedia of
North American Weeds, Vers. 3.0 (DVD-ROM).
Southern Weed Science Society, Champaign,
Illinois.
DeFilipps, R. A. 1980a. Scirpus. Pp. 277–280 in T. G.
Tutin,V. H. Heywood, N. A. Burges, D. M. Moore,
D. H. Valentine, S. M. Walters & D. A. Webb (edi-
tors), Flora Europaea, Vol. 5. Cambridge Univ.
Press, Cambridge.
—. 1980b. Fimbristylis. P. 284 in T. G. Tutin, V. H.
Heywood, N. A. Burges, D. M. Moore, D. H.
Valentine, S. M. Walters & D. A. Webb (editors),
Flora Europaea, Vol. 5. Cambridge Univ. Press,
Cambridge.
—. 1980c. Cyperus. Pp. 284–288 in T. G. Tutin, V. H.
Heywood, N. A. Burges, D. M. Moore, D. H.
Valentine, S. M. Walters & D. A. Webb (editors),
Flora Europaea, Vol. 5. Cambridge Univ. Press,
Cambridge.
Delahoussaye, A. J. & J. W. Thieret. 1967. Cyperus sub-
genus Kyllinga (Cypercaeae) in the continental
United States. Sida 3: 128–136.
Denton, M. F. 1978. A taxonomic treatement of the
Luzulae group of Cyperus. Contr. Univ. Michigan
Herb. 11: 197–271.
Derr, J. F. & J. W. Wilcut. 1993. Control of yellow and
purple nutsedge (Cyperus esculentus and C. rotun-
dus) in nursery crops. Weed Technol. 7: 112–117.
Dickoré, W. B. 1994. Flora Karakorumensis 1.
Angispermae, Monocotyledoneae. Stapfia 39:
1–298.
Dirven, J. 1970. Weed flora of fallow rice fields.
Surinam. Surinaamse Landb. 18: 47–63.
Dowler, C. C., E. W. Hauser & A. W. Johnson. 1974.
Crop-herbicide sequences on a southeastern
coastal plain soil. Weed Sci. 22: 500–505.
Duble, R. L., E. C. Holt & G. G. McBee. 1968. The
translocation of two organic arsenicals in purple
nutsedge. Weed Sci. 16: 421–424.
—, — & —. 1970. Effect of ANA on synthesis and uti-
lization of food reserves in purple nutsedge. Weed
Sci. 18: 174–179.
Duke, S. O. & C. D. Boyette. 2001. Biocontrol: Use of
natural foes. Pp. 31–44 in R. DePrado & J. V.
Jorrin (editors), Uso de Herbicidas en la
Agricultura del Siglo XXI. Univ. Cordoba Press,
Spain.
Elliott, S. 1821. A Sketch of the Botany of South-
Carolina and Georgia, Vol. 1. J. R. Schenck,
Charleston.
Elmore, C. D. 1984. Perennial Vines in the Delta of
Mississippi. Mississippi Agricultural and Forestry
Experiment Station Bulletin 927, Mississippi
State.
— & C. T. Bryson (editors). 1986–2001. Weed Identi-
fication Guide, Sets 1–7. Southern Weed Science
Society, Champaign, Illinois.
— & R. Paul. 1983. Composite list of C4weeds. Weed
Sci. 31: 686–692.
—, L. G. Heatherly & R. A. Wesley. 1989. Perennial
vine control in multiple cropping systems on a clay
soil. Weed Technol. 3: 282–287.
Everett, T. H. 1980–1982. The New York Botanical
Garden Illustrated Encyclopedia of Horticulture,
Vols. 1–10. Garland Publishing,Inc., New York.

The Significance of Cyperaceae as Weeds 59
Eyles, D. & J. Robertson. 1963. A Guide and Key to the
Aquatic Plants of the Southeastern United States.
U.S. Public Health Service Bulletin 186. U.S. Public
Health Service, Washington, D.C.
Fagotto, F. 1987. Sand-dune fixation in Somalia.
Environm. Conservation 14: 157–163.
Fernald, M. L. 1930. Carex macrocephala and C.
anthericoides. Rhodora 32: 9–11.
Ferren, W. R. & A. E. Schuyler. 1980. Intertidal vascu-
lar plants of river systems near Philadelphia. Proc.
Acad. Nat. Sci. Philadelphia 132: 86–120.
Fischer, B. B., A. H. Lange, J. McCaskill, B. Crampton
& B. Tabraham. 1978. Growers’ Weed
Identification Handbook. Univ. of California,
Berkeley.
Ford, B. A. & A. A. Reznicek. 2002. Carex section
Squarrosae. Pp. 518–519 in Flora of North America
Editorial Committee (editors), Flora of North
America North of Mexico, Vol. 23: Magnoliophyta:
Commelinidae (in part): Cyperaceae. Oxford Univ.
Press, New York.
Fosberg, F. R. 1988. Vegetation of Bikini Atoll. Atoll
Res. Bull. 315.
Frans, R. E. 1969. Changing ecology of weeds in cotton
fields. Proc. Beltwide Cotton Prod. Mech. Conf.
23: 24–30.
Frick, K. E. 1978. Purple nutsedge (Cyperus rotundus
L.): A case for augmentation. Pp. 141–151 in T. E.
Freeman (editor), Proceedings of the IV
International Symposium on Biological Control of
Weeds. Univ. of Florida, Gainesville.
Friedman, T. & M. Horowitz. 1971. Biologically active
substances in subterranean parts of purple
nutsedge. Weed Sci. 19: 398–401.
Gaddy, L. L. 1986. Twelve new ant-dispersed species
from the southern Appalachians. Bull. Torrey Bot.
Club 113: 247–251.
Garg, D. K., L. E. Bendixen & R. S. Anderson. 1967.
Rhizome differentiation in yellow nutsedge. Weed
Sci. 15: 124–128.
Garoni, L. W. & W. H. Murdy. 1964. Systematic rela-
tionship of granite outcrop endemic Cyperus gran-
itophilus McVaugh to C. inflexus Muhl. A. S. B.
Bull. 11: 43.
Glaze, N. C. 1987. Cultural and mechanical manipula-
tion of Cyperus spp. Weed Technol. 1: 82–83.
Godfrey,R. K. & J. W. Wooten. 1979. Aquatic and
Wetland Plants of Southeastern United States:
Monocotyledons. Univ. of Georgia Press, Athens.
Goetghebeur, P. 1998. Cyperaceae. Pp. 141–190 in K.
Kubitzki (editor), Families and Genera of Vascular
Plants, Vol. 4. Springer-Verlag. Berlin.
Gordon-Gray, K. D. 1995. Cyperaceae in Natal.
Strelitzia 2. National Botanical Institute, Pretoria,
South Africa.
Govindarajalu, E. 1975. Studies in Cyperaceae. XIV.
Endomorphic evidence for placing Cyperus hyali-
nus under the new subgenus Queenslandiella.
Reinwardtia 9: 187–195.
Graf, A. B. 1985. Exotica Series 4, Vols. 1 & 2, 12th ed.
Roehrs Company Inc., East Rutherford, New Jersey.
Greenlee, J. & D. Fell. 1992. The Encyclopedia of
Ornamental Grasses. Rodale Press, Emmaus,
Pennsylvania.
Grichar, W. J., P. R. Nester & A. E. Colburn. 1992.
Nutsedge (Cyperus spp.) control in peanuts
(Arachis hypogaea) with imazethapyr. Weed
Technol. 6: 396–400.
Grounds, R. 1989. Ornamental Grasses. Christopher
Helm, London.
Guppy, H. 1893. The river Thames as an agent in plant
dispersal. Bot. J. Linn. Soc. 30: 333–346.
Haines, R. W. & K. A. Lye. 1983. The Sedges and
Rushes of East Africa. East African Natural
History Society, Nairobi.
Hamilton, K. C. 1971. Repeated, foliar application of
MSMA on purple nutsedge. Weed Sci. 19:
675–677.
Handel, S. N. 1976. Dispersal ecology of Carex pedun-
culata (Cyperaceae), a new North American
myrmecochore. Amer. J. Bot. 63: 1071–1079.
—. 1978. New ant-dispersed species in the genera
Carex,Luzula, and Claytonia. Canad. J. Bot. 56:
2925–2927.
Hardcastle, W. S. & R. E. Wilkinson. 1971. Reactions of
several crops to dichlobenil. Weed Sci. 19: 655–658.
Hatch, S. L., K. N. Gandhi & L. E. Brown. 1990.
Checklist of the Vascular Plants of Texas. MP-
1655. Texas Agricultural Experiment Station,
College Station.
Hauser, E. W. 1962a. Establishment of nutsedge from
space-planted tubers. Weeds 10: 209–212.
—. 1962b. Development of purple nutsedge under field
conditions. Weeds 10: 315–321.
—. 1963a. Effects of amitrole on purple nutsedge at dif-
ferent growth stages. Weeds 11: 181–183.
—. 1963b. Response of purple nutsedge to amitrole,
2,4-D, and EPTC. Weeds 11: 251–252.
Healy,A. J. & E. Edgar. 1980. Flora of New Zealand,
Vol. 3. P. D. Hasselberg, Wellington.
Hennessy, E. F. F. 1985. The genus Scleria in southern
Africa. Bothalia 15: 505–530.
Hermann, F. J. 1970. Manual of the Carices of the
Rocky Mountains and Colorado Basin. Agriculture
Handbook 374. U.S. Department of Agriculture,
Washington, D.C.
Hesla, B. I., I. L. Tieszen & S. K. Imbamba. 1982. A
systematic survey of C3and C4photosynthesis in
the Cyperaceae of Kenya, East Africa.
Photosynthetica 16: 196–205.
Heywood, V. H. 1993. Flowering Plants of the World.
B. T. Batsford, London.

60 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Hill, J. E., J. R. Smith Jr. & D. E. Bayer. 1994. Rice
weed control: Current technology and emerging
issues in temperate rice. Austral. J. Exp. Agric. 34:
1021–1029.
Hollingsworth, E. B. 1969. Effects of soil incorporation
on control of Carex with herbicides. Proc. Weed
Sci. Soc. Amer. 19: 77.
Hollis, J. 1972. Nematocide-weed interaction in rice
fields. Pl. Dis. Reporter 56: 420–424.
Holm, L. G., D. L. Plucknett, J. V. Pancho & J. P.
Herberger. 1977. The World’s Worst Weeds:
Distribution and Biology. Univ. Press of Hawaii,
Honolulu.
—, J. V. Pancho, J. P. Herberger & D. L. Plucknett.
1979. A Geographical Atlas of World Weeds. John
Wiley & Sons, New York.
—, J. Doll, E. Holm, J. Pancho & J. Herberger. 1997.
World Weeds: Natural Histories and Distribution.
John Wiley & Sons, Inc., New York.
Holt, E. C., J. A. Long & W. W. Allen. 1962. The toxi-
city of EPTC to nutsedge. Weeds 10: 103–105.
Holt, J. S. & D. R. Orcutt. 1991. Functional relation-
ships of growth and competitiveness in perennial
weeds and cotton (Gossypium hirsutum). Weed
Sci. 39: 575–584.
— & —. 1996. Temperature thresholds for bud sprout-
ing in perennial weeds and seed germination in
cotton. Weed Sci. 44: 523–533.
Hooper, S. S. & D. M. Napper. 1972. Cyperaceae. Pp.
278–349 in F. N. Hepper (editor), Flora of West
Tropical Africa. Crown Agents for Oversea
Governments and Administrations, London.
Hopkins, W. G. & N. P. A. Hüner. 2004. Introduction to
Plant Physiology. John Wiley & Sons, Inc.,
Hoboken, New Jersey.
Horak, M. J. & J. S. Holt. 1986. Isozyme variability and
breeding systems in populations of yellow
nutsedge (Cyperus esculentus). Weed Sci. 34:
538–543.
—, — & N. C. Ellstrand. 1987. Genetic variability in
yellow nutsedge (Cyperus esculentus). Weed Sci.
35: 506–512.
Horowitz, M. 1972. Growth, tuber formation, and
spread of Cyperus rotundus L. from single tubers.
Weed Res. 12: 348–363.
Horvat, M. L. 1941. A revision of the subgenus
Mariscus found in the United States. Catholic
Univ. Amer. Biol. Ser. 33: 1–147.
Hughes, G. D. 1995. New Hawaiian plant records. II.
Records of the Hawaii Biological Survey for 1994,
Part 2: Notes. Bishop Mus. Occas. Pap. 42: 1–10.
Hutton, E. E. 1976. Dissemination of perigynia in
Carex pauciflora. Castanea 41: 346–348.
Huxley, A. 1992. The Royal Horticulture Society
Dictionary of Gardening. Royal Horticulture
Society,Wisley.
Jacometti, G. 1912. Le erbe che infestano le risaie ital-
iane. Congresso Risicolo Internazionale, Vercelli
4: 57–91.
Jacono, C. C. 2001. Scleria lacustris (Cyperaceae), an
aquatic and wetland sedge introduced to Florida.
Sida 19: 1163–1170.
James, L., J. Evans, M. Ralphs & R. Child (editors). 1991.
Noxious Range Weeds. Westview Press, Boulder.
Jansen, L. L. 1971. Morphology and photoperiodic
responses of yellow nutsedge. Weed Sci. 19:
210–219.
Johnson, D. E. 1997. Weeds of Rice in West Africa.
West Africa Rice Development Association.
Bouake, Cote d’Ivoire.
Jones, S. D., C. T. Bryson & J. E. Ubelaker. 1993. Carex
blanda and Kyllinga odorata (Cyperaceae) new to
New Mexico and a significant range extension of
Cyperus retrorsus. Sida 15: 252–253.
—, J. K. Wipff & R. Carter. 1996. Nomenclatural com-
binations in Cyperus (Cyperaceae). Phytologia 80:
288–290.
—, — & P. M. Montgomery. 1997. Vascular Plants of
Texas. Univ. of Texas Press, Austin.
Jordan, D. L. 1996. Adjuvants and growth stage affect
purple nutsedge (Cyperus rotundus) control with
chlorimuron and imazethpyr. Weed Technol. 10:
359–362.
Kantor, H. J. 1999. Cyperus alopecuroides Rottb. and
the Egyptian Volute. Pp. 169–198 in Plant
Ornament in the Ancient Near East. Oriental Insti-
tute, Univ. of Chicago. <http://www.oi.uchicago.
edu/research/library/dissertation/kantor.html>,
accessed 7 July 2008.
Kartesz, J. T. 1994. A Synonymized Checklist of the
Vascular Flora of the United States, Canada, and
Greenland, Vol. 1. Timber Press, Portland.
Keeley, P. E. 1987. Interference and interaction of pur-
ple and yellow nutsedges (Cyperus rotundus and
C. esculentus) with crops. Weed Technol. 1:
74–81.
— & R. J. Thullen. 1971. Control of nutsedge with
organic arsenical herbicides. Weed Sci. 19:
601–606.
—, C. H. Carter & J. H. Miller. 1972. Evaluation of the
relative phytotoxicity of herbicides to cotton and
nutsedge. Weed Sci. 20: 71–74.
Kern, J. H. 1974. Cyperaceae I. Pp. 435–753 in C. G. G.
van Steenis (editor), Flora Malesiana, Vol. 7.
Noordhaff, Leiden.
Kissmann, K. G. 1997. Plantas Infestantes e Nocivas:
Tomo 1, 2nd ed. BASF. Seaward Corporation,
West Orange, New Jersey.
Klingman, G. C., F. M. Ashton & L. J. Noordhoff. 1982.
Weed Science: Principles and Practices, 2nd ed.
John Wiley & Sons, Inc., New York.
Knowllton, C. H., J. A. Cushman, W. Deane & A. K.

The Significance of Cyperaceae as Weeds 61
Harrison. 1911. Reports on the flora of Boston
District-VIII. Rhodora 13: 27–29.
Koyama, T. 1985. Cyperaceae. Pp. 125–405 in M. D.
Dassanayake & F. R. Fosberg (editors), Flora of
Ceylon, Vol. 5. Amerind Publishing Co. Pvt. Ltd.,
New Delhi.
Kral, R. 1971. A treatment of Abildgaardia,Bulbostylis,
and Fimbristylis (Cyperaceae) for North America.
Sida 4: 58–277.
—. 1980. A synopsis of Fuirena (Cyperaceae) for the
Americas north of South America. Sida 7: 309–354.
—. 1981. Further additions to some notes on the flora
of the Southern States, particularly Alabama and
Middle Tennessee. Rhodora 83: 301–315.
—. 2002a. Fuirena. Pp. 32–37 in Flora of North
America Editorial Committee (editors), Flora of
North America North of Mexico, Vol. 23:
Magnoliophyta: Commelinidae (in part):
Cyperaceae. Oxford Univ. Press, New York.
—. 2002b. Fimbristylis. Pp. 121–131 in Flora of North
America Editorial Committee (editors), Flora of
North America North of Mexico, Vol. 23: Magno-
liophyta: Commelinidae (in part): Cyperaceae.
Oxford Univ. Press, New York.
—. 2002c. Bulbostylis. Pp. 131–136 in Flora of North
America Editorial Committee (editors), Flora of
North America North of Mexico, Vol. 23:
Magnoliophyta: Commelinidae (in part):
Cyperaceae. Oxford Univ. Press, New York.
—. 2002d. Abildgaardia. Pp. 136–137 in Flora of North
America Editorial Committee (editors), Flora of
North America North of Mexico, Vol. 23:
Magnoliophyta: Commelinidae (in part):
Cyperaceae. Oxford Univ. Press, New York.
—. 2002e. Rhynchospora. Pp. 200–239 in Flora of
North America Editorial Committee (editors),
Flora of North America North of Mexico, Vol. 23:
Magnoliophyta: Commelinidae (in part):
Cyperaceae. Oxford Univ. Press, New York.
Kühn, U. 1982. Cyperaceae. In E. Häfliger, U. Kühn, L.
Hämet-Ahti, C. D. K. Cook, R. Faden & F. Speta
(editors), Monocot Weeds 3. Ciba-Geigy Ltd.,
Basel, Switzerland.
Kükenthal, G. 1935–1936. Cyperaceae–Scirpoideae–
Cypereae. Pp. 1–671 in A. Engler (editor), Das
Pflanzenreich IV, 20 (Heft 101). Wilhelm
Englemann, Leipzig.
Kukkonen, I. 2001. Cyperaceae, No. 206. Pp. 1–277 in
S. I. Ali & M. Qaiser (editors), Flora of Pakistan.
Univ. of Karachi, Karachi; Missouri Botanical
Garden, St. Louis.
Kunth, C. S. 1837. Enumeratio plantarum, Vol. 2. J. G.
Cottae, Stuttgart & Tübingen.
Le Bourgeois, T. & H. Merlier. 1995. Les Adventices
d’Afrique Soudano-sahelienne. CIRAD-CA,
Montpellier.
LeBaron, H. M. 1991. Distribution and seriousness of
herbicide-resistant weed infestations worldwide.
Pp. 27–43 in J. C. Caseley, G. W. Cussans & R. K.
Atkins (editors), Herbicide Resistance in Weeds
and Crops. Butterworth-Heinemann, Ltd., Oxford.
Lelong, M. G. 1988. Noteworthy monocots of Mobile and
Baldwin counties, Alabama. Sida 13: 101–113.
Lin, C. I. 1968. Weeds Found on Cultivated Land in
Taiwan, Vol. 1. Dept. Agronomy, College of
Agriculture, National Taiwan Univ., Taipei.
Lipscomb, B. L. 1980. Cyperus difformis L.
(Cyperaceae) in North America. Sida 8: 320–327.
Long, J. A., W. W. Allen & E. C. Holt. 1962. Control of
nutsedge in bermudagrass turf. Weeds 10: 285–287.
Long, R. W. & O. Lakela. 1971. A Flora of Tropical
Florida. Univ. of Miami Press, Coral Gables.
Lorenzi, H. 1982. Plantas Daninhas do Brasil:
Terrestres, Aquaticas, Parasitas, Toxicas e
Medicinais. Nova Odessa, São Paulo.
Lye, K. A. 1971. Studies in African Cyperaceae 2. The
genus Oxycaryum Nees. Bot. Not. 124: 280–286.
—. 1973. Studies in African Cyperaceae 8. The taxo-
nomic position of Abildgaardia Vahl and Nemum
Hamilton. Bot. Not. 126: 325–329.
—. 1981. Studies in African Cyperaceae 19. The genera
of Anosporum Nees and Sorostachys Steudel.
Nordic J. Bot. 1: 186–191.
—. 1992. The history of the genus Mariscus. Lidia 3:
37–72.
—. 2000. Achene structure and function of structure in
Cyperaceae. Pp. 615–628 in K. L. Wilson & D. A.
Morrison (editors), Monocots: Systematics and
Evolution. CSIRO Publishing, Collingwood,
Victoria, Australia.
Mabberley, D. J. 1997. The Plant Book, 2nd ed.
Cambridge Univ. Press, Cambridge.
Mallik, M. & K. Tesfai. 1988. Alleopathic effect of
common weeds on soybean’s growth and soy-
bean–bradrhizobium symbiosis. Pl. & Soil 112:
177–182.
Martinez-Diaz, G. 1997. Allelopathy of Purple
Nutsedge (Cyperus rotundus L.) on Cotton
(Gossypium). Ph.D. Dissertation, Univ. of Arizona,
Tuscon.
Mastrogiuseppe, J. 2002. Carex section Macgocephalae.
Pp. 307–309 in Flora of North American Editorial
Committee (editors), Flora of North America
North of Mexico, Vol. 23: Magnoliophyta: Com-
melinidae (in part): Cyperaceae. Oxford Univ.
Press, New York.
McGivney, M. V. 1938. A revision of the subgenus
Eucyperus found in the United States. Catholic
Univ. Amer. Biol. Ser. 26: 1–74.
McKenzie, P. M., B. Jacobs, C. T. Bryson, G. C. Tucker
& R. Carter. 1998. Cyperus fuscus (Cyperaceae),
new to Missouri and Nevada, with comments on

62 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
its occurrence in North America. Sida 18:
325–333.
McNaughton, S. J. & L. L. Wolf. 1973. General Ecology.
Holt, Rinehart & Winston, Inc., New York.
McWhorter, C. G. 1982. The use of adjuvants. Pp.
10–21 in R. H. Hodgson (editor), Adjuvants for
Herbicides. Weed Science Society of America,
Champaign, Illinois.
— & C. T. Bryson. 1992. Herbicide use trends. Pp.
233–294 in C. G. McWhorter & J. R. Abernathy
(editors), Weeds of Cotton: Characterization and
Control. The Cotton Foundation, Memphis.
Mears, R. & G. W. Libby. 1995. New records of
Cyperus (Cyperaceae) from Kentucky. Castanea
60: 79–82.
Michaux, A. 1803. Flora Boreali-Americana. Levrault,
Paris.
Miles, J. E., K. K. Nishimoto & O. Kawabata. 1996.
Dirurnally altering temperatures stimulate sprout-
ing of purple nutsedge (Cyperus rotundus) tubers.
Weed Sci. 44: 122–125.
Miller, J. H. & K. V. Miller. 1999. Forest Plants of the
Southeast and Their Wildlife Uses. Southern Weed
Science Society, Champaign, Illinois.
Mohr, C. 1901. Plant life of Alabama. Contr. U.S.
National Herb. 6: 1–921.
Moody, K. 1981. Major Weeds of Rice in South and
Southeast Asia. International Rice Research
Institute, Los Banos, Laguna, Philippines.
—. 1989. Weeds Reported in Rice in South and
Southeast Asia. International Rice Research
Institute, Manila.
Moore, L. B. & E. Edgar. 1970. Flora of New Zealand,
Vol. 2. A. R. Shearer, Wellington.
Morales-Payan, J. P., R. Charudattan & W. M. Stall.
2005. Fungi for biological control of weedy
Cyperaceae, with emphasis on purple and yellow
nutsedges (Cyperus rotundus and C. esculentus).
Outlooks on Pest Management 16: 148–155.
Muasya, A. M., J. J. Bruhl, D. A. Simpson, A. Culham &
M. W. Chase. 2000a. Suprageneric phylogeny of
Cyperaceae: A combined analysis. Pp. 593–601 in
K. L. Wilson & D. A. Morrison (editors), Monocots:
Systematics and Evolution. CSIRO Publishing,
Collingwood, Victoria, Australia.
—, D. A. Simpson, M. W. Chase & A. Culham. 2000b.
Phylogenetic relationships within the heteroge-
nous Scirpus s. lat. (Cyperaceae) inferred from
rbcL and trnL-F sequence data. Pp. 610–614 in K.
L. Wilson & D. A. Morrison (editors), Monocots:
Systematics and Evolution. CSIRO Publishing,
Collingwood,Victoria, Australia.
—, — & —. 2002. Phylogenetic relationships in
Cyperus L. s.l. (Cyperaceae) inferred from plastid
DNA sequence data. Bot. J. Linn. Soc. 138:
145–153.
Muenscher, W. C. 1955. Weeds, 2nd ed. Cornell Univ.
Press, Ithaca.
Mühlenbach, V. 1979. Contributions to the synanthrop-
ic (adventive) flora of the railroads in St. Louis,
Missouri, U.S.A. Ann. Missouri Bot. Gard. 66:
1–108.
—. 1983. Supplement to the contributions to the synan-
thropic (adventive) flora of the railroads in St.
Louis, Missouri, U.S.A. Ann. Missouri Bot. Gard.
70: 170–178.
Murray, D. S., L. M. Verhalen & R. J. Tyrl. 1992. The
changing weed problem in cotton. Pp. 117–232 in
C. G. McWhorter & J. R. Abernathy (editors),
Weeds of Cotton: Characterization and Control.
The Cotton Foundation, Memphis.
Naczi, R. F. C. 1984. Rare sedges discovered and redis-
covered in Delaware. Bartonia 50: 31–35.
—, R. J. Driskill, E. L. Pennell, N. E. Seyfried, A. O.
Tucker & N. H. Dill. 1986. New records of some
rare DelMarVa sedges. Bartonia 52: 49–57.
Nees von Esenbeck, C. G. 1842. Cyperaceae. In C. F. P.
Martius (editor), Flora Brasiliensis, Vol. 2, Pt. 1. F.
Fleischer, Leipzig.
Negbi, M. 1992. A sweetmeat plant, perfume plant and
their relatives: A chapter in the history of Cyperus
esculentus L. and C. rotundus L. Econ. Bot. 46:
64–71.
Noda, K. & S. Eguchi. 1965. Studies on ecology of
weeds on arable lands: 1. Emergence patterns of
annual representative weeds which are commonly
found on the paddy rice fields of south-western
Japan. Bull. Kyushu Agric. Exp. Sta. 11: 153–170.
Ohwi, J. 1965. Flora of Japan. Smithsonian Institution,
Washington, D.C.
O’Neill, H. T. 1938a. Cyperus pilosus Vahl in the
United States. Rhodora 40: 74.
—. 1938b. Cyperus oxylepis Nees ex Steudel in the
United States. Rhodora 40: 358.
—. 1946. The genus Cyperus in Cuba. Contr. Ocas.
Mus. Hist. Nat. Colegio “De La Salle” 6: 1–15.
Ottesen, C. 1989. Ornamental Grasses. McGraw-Hill
Publishing Co., New York.
Palm, C. E., W. W. Dykstra, G. R. Ferguson, R.
Hansberry, W. J. Hayes Jr., L. W. Hazleton, J. G.
Horsfall, E. F. Knipling, L. D. Leach, L. W.
Lovvorn & G. A. Swanson. 1968. Principles of
Plant and Animal Pest Control, Vol. 2: Weed
Control. National Academy of Sciences,
Washington, D.C.
Pappas-Fader, T., J. F. Cook, T. Butler, P. J. Lana & J.
Hare. 1993. Resistance of California arrowhead
and smallflower umbrella sedge to sulfonylurea
herbicides. Proc. W. Soc. Weed Sci. 46: 76.
—, R. G. Turner, J. F. Cook, T. D. Butler, P. J. Lana &
M. Carriere. 1994. Resistance monitoring program
for aquatic weeds to sulfonylurea herbicides in

The Significance of Cyperaceae as Weeds 63
California rice fields. Proc. Rice Tech. Working
Group 25: 165.
Parsons, W. T. & E. G. Cuthbertson. 1992. Noxious
Weeds of Australia. Inkata Press, Melbourne.
Patterson, D. T. 1982. Shading responses of purple and
yellow nutsedges (Cyperus esculentus and C.
rotundus). Weed Sci. 30: 25–30.
Pedersen, T. M. 1995. Nueva combinacion in
Cyperaceae. Hickenia 2: 138.
Pereira, W., G. Crabtree & R. D. William. 1987.
Herbicide action on purple and yellow nutsedge
(Cyperus rotundus and C. esculentus). Weed
Technol. 1: 92–98.
Peterson, J. K. & H. F. Harrison Jr. 1995. Sweet potato
allelopathic substance inhibits growth of purple
nutsedge (Cyperus rotundus). Weed Technol. 9:
277–280.
Petrík, P. 2003. Cyperus eragrostis—A new alien
species for the Czech flora and the history of its
invasion of Europe. Preslia 75: 17–28.
Phatak, S. C., M. B. Calloway & C. S. Varrina. 1987.
Biological control and its integration in weed man-
agement systems for purple and yellow nutsedge
(Cyperus rotundus and C. esculentus). Weed
Technol. 1: 84–91.
Pijl, L. van der. 1982. Principles of Dispersal in Higher
Plants. Springer-Verlag, New York.
Pio Corrêa, M. 1926–1984. Dicionário das Plantas úteis
do Brasil e das Exoticas Cultivadas, Vols. 1–6.
Ministerio da Agricultura Instituto Brasileiro de
Desenvolvimento Forestal Brazil, Imprensa
Nacional, Rio de Janeiro.
Plowman, A. B. 1906. The comparative anatomy and
phylogeny of Cyperaceae. Ann. Bot. (Oxford) 20:
1–37.
Radford, A. E., H. A. Ahles & C. R. Bell. 1968. Manual
of the Vascular Flora of the Carolinas. Univ. of
North Carolina Press, Chapel Hill.
Radosevich, S. R. & J. S. Holt. 1984. Weed Ecology.
John Wiley & Sons, New York.
Ramos, G. S., V. W. Cárdenas, P. B. Aquilar & R. P.
Marín. 2004. Atlas de Malezas Arvenses del
Estado de Querétaro. Universidad Autonoma de
Querétaro, Querétaro, México.
Ranade, S. B. & W. Burns. 1925. The eradication of
Cyperus rotundus L. Mem. Dept. Agric. India,
Bot. Ser. 13: 99–192.
Randall, J. M. 1996. Weed control for the preservation
of biological diversity. Weed Technol. 10: 370–383.
—. 1997. Defining weeds of natural areas. Pp. 18–25 in
J. O. Luken & J. W. Thieret (editors), Management
of Plant Invasions. Springer-Verlag, New York.
Ratliff, R. D. & S. E. Westfall. 1992. Restoring plant
cover on high-elevation gravel areas, Sequoia
National Park, California. Biol. Conservation 60:
189–195.
Ravi, N. & N. Mohanan. 2002. Common Tropical and
Sub-tropical Sedges and Grasses. Science
Publishers, Inc., Enfield, New Hampshire.
Reed, C. F. 1977. Economically Important Foreign
Weeds. Agriculture Handbook No. 498. U.S.
Department of Agriculture, Washington, D.C.
Reichard, S. H. & C. W. Hamilton. 1997. Predicting
invasions of woody plants introduced into North
America. Conservation Biol. 11: 193–203.
Rejmanek, M. 1996. A theory of seed plant invasiveness:
The first sketch. Biol. Conservation 78: 171–181.
Reznicek, A. A. 2002. Carex section Rhynchosystis. Pp.
420–421 in Flora of North America Editorial
Committee (editors), Flora of North America
North of Mexico, Vol. 23: Magnoliophyta:
Commelinidae (in part): Cyperaceae. Oxford Univ.
Press, New York.
— & P. M. Catling. 1987. Carex praegracilis in eastern
North America: A remarkable case of rapid inva-
sion. Rhodora 89: 205–216.
— & —. 2002. Carex section Divisae. Pp. 302–306 in
Flora of North America Editorial Committee (edi-
tors), Flora of North America North of Mexico,
Vol. 23: Magnoliophyta: Commelinidae (in part):
Cyperaceae. Oxford Univ. Press, New York.
—, — & S. M. McKay. 1976. Carex praegracilis W.
Boott, recently adventive in southern Ontario.
Canad. Field-Naturalist 90: 180–183.
—, J. E. Fairey III & A. T. Whittemore. 2002. Scleria.
Pp. 242–251 in Flora of North America Editorial
Committee (editors), Flora of North America
North of Mexico, Vol. 23: Magnoliophyta:
Commelinidae (in part): Cyperaceae. Oxford Univ.
Press, New York.
Rhoads, A. F. & W. M. Klein Jr. 1993. The Vascular
Flora of Pennsylvania: Annotated Checklist and
Atlas. American Philosophical Society,
Philadelphia.
Richburg, J. S., III, J. W. Wilcut & G. R. Wehtje. 1993.
Toxicity of imazethpyr to purple (Cyperus rotun-
dus) and yellow nutsedge (C. esculentus). Weed
Technol. 7: 900–905.
—, — & —. 1994. Toxicity of AC 263,322 to purple
(Cyperus rotundus) and yellow nutsedge (C. escu-
lentus). Weed Sci. 42: 398–402.
Ridley, H. N. 1930. The Dispersal of Plants Throughout
the World. L. Reeve & Co., Ltd., Ashford, United
Kingdom.
Rosen, D. J., R. Carter & C. T. Bryson. 2006. The
spread of Cyperus entrerianus (Cyperaceae) in the
southeastern United States and its invasive poten-
tial in bottomland hardwood forests. S. E.
Naturalist 5: 333–344.
Rothrock, P. E. 1991. The identity of Carex albo-
lutescens,C. festucacea, and C. longii (Cyper-
aceae). Rhodora 93: 51–66.

64 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Ryang, H., J. Chun & Y. Moon. 1978. Control of the
perennial weed Scirpus maritimus in reclaimed
paddy fields of west seashore. II. Physiological
and ecological characteristics of S. maritimus. J.
Korean Soc. Crop Sci. 23: 64–73.
Rzedowski, G. C. de & J. Rzedowski. 2004. Manual de
Malezas de la Región de Salvatierra, Guanajuato.
Flora del Bajío y de Regiones Adyacentes,
Fascículo complementario XX. Instituto de
Ecología, Veracruz, México.
Sauer, J. D. 1988. Plant Migration. Univ. of California
Press, Berkeley.
Schery, R. W. 1972. Plants for Man, 2nd ed. Prentice-
Hall, Inc., Englewood Cliffs, New Jersey.
Schippers, P., S. J. ter Borg, J. M. van Groenendael &
B. Habekotte. 1993. What makes Cyperus esculen-
tus (yellow nutsedge) an invasive species? A spa-
tial model approach. Pp. 495–504 in Proceedings
of the Brighton Crop Protection Conference—
Weeds. Brighton Crop Protection Registered
Office, Farham, Surrey, England.
—, — & J. J. Bos. 1995. A revision of the infraspecific
taxonomy of Cyperus esculentus (yellow
nutsedge) with an experimentally evaluated char-
acter set. Syst. Bot. 20: 461–481.
Schroeder, J., S. H. Thomas & L. Murray. 1993. Yellow
and purple nutsedge and chile peppers host south-
ern root-knot nematode. Weed Sci. 41: 150–156.
Sculthorpe, C. 1967. The Biology of Aquatic Vascular
Plants. Edward Arnold Ltd., London.
Sell, P. & G. Murrell. 1996. Flora of Great Britain and
Ireland, Vol. 5. Cambridge Univ. Press,
Cambridge.
Shaner, D. L. & J. L. Lyon. 1980. Interaction of
glyphosate with aromatic amino acids on transpi-
ration in Phaseolus vulgaris. Weed Sci. 28: 31–35.
Shear, G. M. 1985. Introduction and history of limited
tillage. Pp. 1–14 in A. F. Wiese (editor), Weed
Control in Limited-tillage Systems. Weed Science
Society of America, Champaign, Illinois.
Simpson, D. A. 1992. A Revision of the Mapania.
Royal Botanic Gardens, Kew.
—. 1993. Plant portrait 225: Rhynchospora nervosa.
Bot. Mag. (Kew Mag.) 10: 117–121.
—. 1994. Plant portrait 236: Cyperus prolifer
(Cyperaceae). Bot. Mag. (Kew Mag.) 11: 6–8.
— & C. A. Inglis. 2001. Cyperaceae of economic, eth-
nobotanical, and horticultural importance: A
checklist. Kew Bull. 56: 257–360.
— & T. Koyama. 1998. Cyperaceae. Pp. 247–485 in T.
Santisuk & K. Larsen (editors), Flora of Thailand,
Vol. 6. The Forest Herbarium, Royal Forest
Department, Bangkok.
Sipple, W. S. 1978. An Atlas of Vascular Plant Species
Distribution Maps for Tidewater Maryland.
Wetland Publ. No. 1. Maryland Department of
Natural Resources, Annapolis.
Small, J. A. 1954. Carex kobomugi at Island Beach,
New Jersey. Ecology 35: 289–291.
Small, J. K. 1933. Manual of the Southeastern Flora.
Univ. of North Carolina Press, Chapel Hill.
Smith, A. H. 1867. On colonies of plants observed near
Philadelphia. Proc. Acad. Nat. Sci. Philadelphia
1867: 15–24.
Smith, S. G. 2002a. Bolboschoenus. Pp. 37–44 in Flora
of North America Editorial Committee (editors),
Flora of North America North of Mexico, Vol. 23:
Magnoliophyta: Commelinidae (in part):
Cyperaceae. Oxford Univ. Press, New York.
—. 2002b. Schoenoplectus. Pp. 44–60 in Flora of North
America Editorial Committee (editors), Flora of
North America North of Mexico, Vol. 23:
Magnoliophyta: Commelinidae (in part): Cyper-
aceae. Oxford Univ. Press, New York.
—. 2002c. Eleocharis ser. Ovatae. Pp. 102–105 in Flora
of North America Editorial Committee (editors),
Flora of North America North of Mexico, Vol. 23:
Magnoliophyta: Commelinidae (in part): Cyper-
aceae. Oxford Univ. Press, New York.
—. 2002d. Isolepis. Pp. 137–140 in Flora of North
America Editorial Committee (editors), Flora of
North America North of Mexico, Vol. 23:
Magnoliophyta: Commelinidae (in part): Cyper-
aceae. Oxford Univ. Press, New York.
—, A. E. Schuyler & W. J. Crins. 1993. Scirpus. Pp.
1146–1148 in J. C. Hickman (editor), The Jepson
Manual: Higher Plants of California. Univ. of
California Press, Berkeley.
—, J. J. Bruhl, M. S. Gonzalez-Elizondo & F. J.
Menapace. 2002. Eleocharis. Pp. 60–120 in Flora
of North America Editorial Committee (editors),
Flora of North America North of Mexico, Vol. 23:
Magnoliophyta: Commelinidae (in part): Cyper-
aceae. Oxford Univ. Press, New York.
Snyder, D. B. 1983. Rare New Jersey grasses and
sedges. Bartonia 49: 71–72.
—. 1984. Botanical discoveries of Vincent Abraitys.
Bartonia 50: 54–56.
Soejani, M., A. J. G. H. Kosteramans & G.
Tjitrosoepomo. 1987. Weeds of Rice in Indonesia.
Balai Pustaka, Jakarta.
Soros, C. L. & J. J. Bruhl. 2000. Multiple Evolutionary
Origins of C4Photosynthesis in the Cyperaceae.
Pp. 629–636 in K. L. Wilson & D. A. Morrison
(editors), Monocots: Systematics and Evolution.
CSIRO Publishing, Collingwood, Victoria,
Australia.
Stalter,R. 1980. Carex kobomugi at Sandy Hook, New
Jersey. Bull. Torrey Bot. Club 107: 431–432.
Standley, L. A. 1983. Carex kobomugi Ohwi, an adven-
tive sedge to New England. Rhodora 85: 265–267.
—. 2002. Carexsection Vulpinae. Pp. 273–278 in Flora

The Significance of Cyperaceae as Weeds 65
of North America Editorial Committee (editors),
Flora of North America North of Mexico, Vol. 23:
Magnoliophyta: Commelinidae (in part): Cyper-
aceae. Oxford Univ. Press, New York.
—, J. Cayouette & L. Bruederle. 2002. Carex section
Phacocystis. Pp. 379–401 in Flora of North
America Editorial Committee (editors), Flora of
North America North of Mexico, Vol. 23:
Magnoliophyta: Commelinidae (in part): Cyper-
aceae. Oxford Univ. Press, New York.
Stephens, J. M. 1994. Chufa Cyperus esculentus L. var.
sativus Boeck. Florida Cooperative Extension
Service Fact Sheet HS-583, Univ. of Florida,
Gainesville.
Steward, K. K. & W. H. Ornes. 1975. The autecology of
sawgrass in the Florida Everglades. Ecology 56:
162–171.
Stoller, E. W. & R. D. Sweet. 1987. Biology and life
cycle of purple and yellow nutsedges (Cyperus
rotundus and C. esculentus). Weed Technol. 1:
66–73.
—, D. P. Nema & V. M. Bhan. 1972. Yellow nutsedge
tuber germination and seedling development.
Weed Sci. 20: 93–97.
Strong, M. T. & W. L. Wagner. 1997. New and notewor-
thy Cyperaceae from the Hawaiian Islands. Bishop
Mus. Occas. Pap. 48: 37–50.
Stuckey, J. M., T. J. Monaco & A. D. Worsham. 1980.
Identifying Seedling and Mature Weeds Common
in the Southeastern United States. North Carolina
Agricultural Research Service and Agricultural
Extension Service, North Carolina State Univ.,
Raleigh.
Stuckey, R. L. & T. M. Barkley. 1993. Weeds. Pp.
193–198 in Flora of North America Editorial
Committee (editor), Flora of North America North
of Mexico, Vol. 1. Oxford Univ. Press, New York.
Sumaryono & Basuki. 1986. Growth and reproduction
of Cyperus kyllingia Endl. and Cyperus brevifolius
(Rottb.) Hassk. In J. V. Pancho, S. S. Sastroutomo
& S. Tjitrosemito (editors), Symposium in Weed
Science Special Publication, No. 24. Institute of
Tropical Biology, Bogor.
Sundell, E. & R. D. Thomas. 1988. Four new records of
Cyperus (Cyperaceae) in Arkansas. Sida 13:
259–261.
Svenson, H. K. 1957. Cyperaceae: Tribe Scirpeae. N.
Amer. Flora 18: 505–556.
—. 1979. The Flora of Cape Cod. Cape Cod Museum of
Natural History, Brewster.
Swink, F. & G. Wilhelm. 1979. Plants of the Chicago
Region, 3rd ed. Morton Arboretum, Lisle, Illinois.
Tachholm, V. & M. Drar. 1950. Flora of Egypt, Vol. 2.
Bull. Fac. Sci. Fouad I Univ. 28: 1–475.
Tadulingam C. & G. Venkatanaryana. 1955. A
Handbook of Some South Indian Weeds. Madras
Gov. Press, Serial No. 38/1954. Delhi, India.
Terry, P. J. 1976. Sedge Weeds of East Africa. 1.
Identification. E. African Agric. Forest. J. 42:
231–249.
—. 2001. The Cyperaceae—Still the world’s worst
weeds? Pp. 3–18 in C. R. Riches (editor), The
World’s Worst Weeds. British Crop Protection
Council, Farnham, United Kingdom.
Thieret, J. W. 1964. More additions to the Louisiana
flora. Sida 1: 294–295.
—. 1977. Cyperus louisianensis (Cyperaceae), a new
species from southern Louisiana. Proc. Louisiana
Acad. Sci. 40: 23–26.
Thomas, R. D. & C. M. Allen. 1993. Atlas of the
Vascular Flora of Louisiana, Vol. I: Ferns & Fern
Allies, Conifers, & Monocotyledons. Moran
Colorgraphic Printing, Baton Rouge.
Thomas, W. W. 1984a. Systematics of Rhynchospora
sect. Dichromena. Mem. New York Bot. Gard. 37:
1–116.
—. 1984b. Insect pollination of Cymophyllus fraseri
(Andrews) Mackenzie. Castanea 49: 94–95.
Thullen, R. J. & P. E. Keeley. 1979. Seed production
and germination in Cyperus esculentus and C.
rotundus. Weed Sci. 27: 502–505.
Thumbleson, M. E. & T. K. Kommedahl. 1961.
Reproductive potential of Cyperus esculentus by
tubers. Weeds 9: 646–653.
Tietz, H. M. 1972. An Index to the Described Life
Histories, Early Stages and Hosts of the
Macrolepidoptera of the Continental United States
and Canada. Allyn Museum of Entomology,
Sarasota.
Tobe, J. D., K. C. Burks, R. W. Cantrell, M. A. Garland,
M. E. Sweeley, D. W. Hall, P. Wallace, G. Anglin,
G. Nelson, J. R. Cooper, D. Bickner, K. Gilbert,
N. Aymond, K. Greenwood & N. Raymond.
1998. Wetland Plants: An Identification Manual.
Florida Department of Environmental Protection,
Tallahassee.
Torrey, J. 1836. Monograph of North American
Cyperaceae. Ann. Lyceum Nat. Hist. New York 3:
249–288.
Tucker, G. C. 1983. The taxonomy of Cyperus
(Cyperaceae) in Costa Rica and Panama. Syst.
Bot. Monogr. 2: 1–85.
—. 1984. A revision of the genus KyllingaRottb.
(Cyperaceae) in Mexico and Central America.
Rhodora 86: 507–538.
—. 1985. The correct name for Cyperus cayennensis
(C. flavus), Cyperaceae. S. W. Naturalist 30:
607–608.
—. 1987. The genera of Cyperaceae in the southeastern
United States. J. Arnold Arbor. 68: 361–445.
—. 1994. Revision of the Mexican species of Cyperus
(Cyperaceae). Syst. Bot. Monogr. 43: 1–213.

66 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
—. 2002a. Cladium. Pp. 44–60 in Flora of North
America Editorial Committee (editors), Flora of
North America North of Mexico, Vol. 23:
Magnoliophyta: Commelinidae (in part): Cyper-
aceae. Oxford Univ. Press, New York.
—. 2002b. Kyllinga. Pp. 193–194 in Flora of North
America Editorial Committee (editors), Flora of
North America North of Mexico, Vol. 23:
Magnoliophyta: Commelinidae (in part): Cyper-
aceae. Oxford Univ. Press, New York.
—. 2002c. Lipocarpha. Pp. 195–197 in Flora of North
America Editorial Committee (editors), Flora of
North America North of Mexico, Vol. 23:
Magnoliophyta: Commelinidae (in part): Cyper-
aceae. Oxford Univ. Press, New York.
—, B. G. Marcks & J. R. Carter. 2002. Cyperus. Pp.
141–191 in Flora of North America Editorial
Committee (editors), Flora of North America
North of Mexico, Vol. 23: Magnoliophyta:
Commelinidae (in part): Cyperaceae. Oxford Univ.
Press, New York.
Turner, R. J., Jr. & E. Wasson (editors). 1998. Botanica.
Random House Australia Pty. Ltd., Milsons Point,
Sydney.
Ueno, O., M. Samejima & T. Koyama. 1989. Distribu-
tion and evolution of C4syndrome in Eleocharis,
a sedge group inhabiting wet and aquatic envi-
ronments, based on culm anatomy and carbon
isotope ratios. Ann. Bot. (Oxford) 64: 425–438.
Uittien, H. 1932. Cyperaceae. Pp. 72–149 in A. Pulle
(editor), Flora of Suriname. E. J. Brill, Leiden.
U.S. Army Corps of Engineers. 1999. Central and
Southern Florida Project Comprehensive Review
Study: Final Integrated Feasibility Report and
Programmatic Environmental Impact Statement
(CD-ROM). U.S. Army Corps of Engineers,
Jacksonville District, Florida.
USDA. 1960. Index of Plant Diseases in the United
States. Agriculture Handbook 165. U.S. Department
of Agriculture Government Printing Office,
Washington, D.C.
—. 1970. Selected Weeds of the United States.
Agriculture Handbook 366. U.S. Department of
Agriculture, Washington, D.C.
USFWS. 1988. National List of Vascular Plant Species
that Occur in Wetlands. U.S. Fish and Wildlife
Service Biological Report 88 (24), Washington,
D.C.
Vaillant, A. 1967. Chemical control of annual weeds in
rice. World Crops 19: 38–44.
Veken, P. van der. 1965. Contribution a l’embryolgra-
phie systématique des Cyperaceae–Cyperoideae.
Bull. Jard. Bot. État. Bruxelles 35: 285–352.
Vencill, W. K., J. S. Richburg III, J. W. Wilcut & L. R.
Hawf. 1995. Effect of MON-12037 on purple
(Cyperus rotundus) and yellow (Cyperus esculen-
tus) nutsedge. Weed Technol. 9: 148–152.
Verga, B., K. Moody & R. Visperas. 1977. Autecology
of Scirpus maritimus, IV. Suggested control under
field conditions. Philipp. Weed Sci. Bull. 4: 7–12.
Verga, M. & J. Sierra. 1970. Population of weed seeds
in a lowland rice field. Philipp. Agric. 54: 1–7.
Vlaming, V. de & V. W. Proctor. 1968. Dispersal of
aquatic organisms: Viability of seeds recovered
from droppings of captive killdeer and mallard
ducks. Amer. J. Bot. 55: 20–26.
Vorster, P. 1996. Justification for the generic status of
Courtoisina (Cyperaceae). Bot. J. Linn. Soc. 121:
271–280.
Wagner, W. L. & D. R. Herbst. 1995. Contributions to
the flora of Hawaii. IV. New records and name
changes. Bishop Mus. Occas. Pap. 42: 13–27.
—, — & S. H. Sohmer. 1990. Manual of the Flowering
Plants of Hawaii, Vol. 2. Univ. of Hawaii Press,
Bishop Museum, Honolulu.
Walters, S. M. 1980. Eleocharis. Pp. 281–284 in T. G.
Tutin, V. H. Heywood, N. A. Burges, D. M. Moore,
D. H. Valentine, S. M. Walters & D. A. Webb (edi-
tors), Flora Europaea, Vol. 5. Cambridge Univ.
Press, Cambridge.
Waterhouse, D. F. 1997. The Major Invertebrate Pests
and Weeds of Agriculture and Plantation Forestry
in the Southern and Western Pacific. ACIAR
Monog. 44. Australian Centre for International
Agricultural Research, Canberra.
Watkins, J. V. & T. J. Sheehan. 1975. Florida Landscape
Plants. Univ. Presses of Florida, Gainesville.
Webb, D. H. & W. M. Dennis. 1981. Additions to the
flora of Tennessee. Sida 9: 184–185.
—, — & T. S. Patrick. 1981. Distribution and natural-
ization of Cyperus brevifolioides (Cyperaceae) in
eastern United States. Sida 9: 188–190.
Weedon, R. R. & H. A. Stephens. 1969. Cyperus fuscus
in Nebraska and South Dakota. Rhodora 71: 433.
Westbrooks, R. G. 1998. Invasive Plants, Changing the
Landscape of America: Fact Book. Federal
Interagency Committee for the Management of
Noxious and Exotic Weeds, Washington, D.C.
—. 2001. Invasive species, coming to America:
New strategies for biological protection through
prescreening, early warning, and rapid response.
Wildland Weeds 4: 5–11.
Whistler,W. A. 1994. Wayside Plants of the Islands:
A Guide to the Lowland Flora of the Pacific
Islands Including Hawaii, Samoa, Tonga, Tahiti,
Fiji, Guam and Belau. Isle Botanica, Honolulu.
Whittemore, A. T. & A. F. Schuyler. 2002. Scirpus. Pp.
8–21 in Flora of North America Editorial
Committee (editors), Flora of North America
North of Mexico, Vol. 23: Magnoliophyta:
Commelinidae (in part): Cyperaceae. Oxford
Univ. Press, New York.

The Significance of Cyperaceae as Weeds 67
Whitwell, T. & R. Smith. 1997. Perennial kyllinga
control in nursery crops. Proc. Weed Sci. Soc.
Amer. 37: 28.
Wiese, A. F. 1986. Herbicide application. Pp. 1–27 in
N. D. Camper (editor), Research Methods in Weed
Science. Southern Weed Science Society,
Champaign, Illinois.
Wilcut, J. W., J. S. Richburg III, E. F. Eastin, G. R.
Wiley, F. R. Walls Jr. & S. Newell. 1994.
Imazethapyr and paraquat systems for weed man-
agement in peanut (Arachis hypogaea). Weed Sci.
42: 601–607.
Williams, R. D. 1978. Photoperiod effects on the repro-
ductive biology of purple nutsedge (Cyperus
rotundus). Weed Sci. 26: 539–542.
Wills, G. D. 1972. Sugars, phosphorus, and iron in pur-
ple nutsedge. Weed Sci. 20: 348–350.
—. 1987. Description of purple and yellow nutsedge
(Cyperus rotundus and C. esculentus). Weed
Technol. 1: 2–9.
—. 1998. Comparison of purple nutsedge (Cyperus
rotundus) from around the world. Weed Technol.
12: 491–503.
— & G. A. Briscoe. 1970. Anatomy of purple nutsedge.
Weed Sci. 18: 631–635.
— & C. G. McWhorter. 1988. Effect of inorganic salts
on the toxicity and translocation of glyphosate and
MSMA in purple nutsedge (Cyperus rotundus).
Weed Sci. 33: 755–761.
—, C. T. Bryson & J. E. Hanks. 1991. New low-volume
application technology for purple nutsedge
(Cyperus rotundus) control. Proc. Beltwide Cotton
Prod. Res. Conf. 15: 959.
Wilson, K. L. 1981. A synopsis of the genus Scirpus
sens. lat. in Australia. Telopea 2: 153–172.
—. 1993. Cyperaceae. Pp. 293–396 in G. J. Harden
(editor), Flora of New South Wales, Vol. 4. New
South Wales Univ. Press, Kensington.
WSSA. 1989. Composite List of Weeds. Weed Science
Society of America, Champaign, Illinois.
Wunderlin, R. P. 1982. Guide to the Vascular Plants of
Central Florida. Univ. Presses of Florida, Tampa.
—. 1998. Guide to the Vascular Plants of Florida. Univ.
Press Florida, Gainesville.
Yelverton, F. 1996. Know your sedges. Golf Course
Management 64: 56–60.
Zandstra, B. H. & R. K. Nishimoto. 1977. Movement
and activity of glyphosate in purple nutsedge.
Weed Sci. 25: 268–274.
—, C. K. H. Teo & R. K. Nishimoto. 1974. Response of
purple nutsedge to repeated applications of
glyphosate. Weed Sci. 22: 230–232.
Zeven, A. C. & P. M. Zhukovsky. 1975. Dictionary of
Cultivated Plants and Their Centers of Diversity.
Centre for Agricultural Publishing and Docu-
mentation, Wageningen, The Netherlands.
Zhirong, W., X. Mingyuan, M. Dehui, S. Shunzu,
W. Xianfeng, Y. Chunbo, Z. Dianjing, F. Weizhuo,
M. Enhui & C. Jixian. 1990. Farmland Weeds
in China: A Collection of Coloured Illustrative
Plates. Agricultural Publishing House, Shanghai.
Zimdahl, R. L. 1995. Introduction. Pp. 1–18 in A. E.
Smith (editor), Handbook of Weed Management
Systems. Marcel Dekker, Inc., New York.

68 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
APPENDIX 1
Known and suspected anthropogenic dispersal in Cyperaceae.
Species1Method of dispersal Source
Bolboschoenus glaucus (Lam.) planted as waterfowl food, Browning et al., 1995; Smith, 2002a
S. G. Sm. rice agriculture
Bolboschoenus maritimus (L.) Palla rice agriculture Holm et al., 1997
Bolboschoenus maritimus subsp. planted as waterfowl food Smith, 2002a
paludosus (A. Nelson) T. Koyama
Bolboschoenus robustus (Pursh) Soják ornamental Everett, 1980–1982
Bulbostylis humilis (Kunth) C. B. Clarke wool alien Sell & Murrell, 1996
Bulbostylis striatella C. B. Clarke wool alien Sell & Murrell, 1996
Carex acuta L. ornamental Grounds, 1989; Huxley, 1992
Carex acutiformis Ehrh. ornamental Huxley, 1992
Carex alba Scop. ornamental Huxley, 1992
Carex albula Allan ornamental Greenlee & Fell, 1992; Turner & Wasson,
1998; Darke, 1999
Carex appressa R. Br. erosion control, wool alien Huxley, 1992; Sell & Murrell, 1996;
Simpson & Inglis, 2001
Carex arenaria L. ornamental Huxley, 1992
Carex atrata L. ornamental Grounds, 1989; Huxley, 1992
Carex austrina Mack. railroad adventive Mühlenbach, 1983
Carex baccans Nees ornamental Bailey, 1935; Greenlee & Fell, 1992; Huxley,
1992; Darke, 1999
Carex baldensis L. ornamental Huxley, 1992
Carex baltzellii Chapm. ex Dewey ornamental Darke, 1999
Carex berggreni Petrie ornamental Grounds, 1989; Huxley, 1992; Darke, 1999
Carex brevior (Dewey) Mack. ex Lunell contaminated grass seed Bryson et al., 1992
Carex brunnea Thunb. ornamental Grounds, 1989; Huxley, 1992
Carex buchananii Berggr. ornamental Bailey, 1935; Brooklyn Botanical Garden,
1988; Ottesen, 1989; Greenlee & Fell, 1992;
Darke, 1999
Carex caryophyllea Latourr. ornamental Huxley, 1992; Darke, 1999
Carex cherokeensis Schwein. hay Bryson, pers. obs.
Carex comans Berggr. ornamental Everett, 1980–1982; Greenlee & Fell, 1992;
Turner & Wasson, 1998; Darke, 1999
Carex conica Boott ornamental Bailey & Bailey, 1976; Ottesen, 1989;
Greenlee & Fell, 1992; Darke, 1999
Carex crawfordii Fernald railroad adventive Mühlenbach, 1983
Carex crinita Lam. ornamental Darke, 1999
Carex curvula All. ornamental Huxley, 1992
Carex devia Cheeseman wool alien Sell & Murrell, 1996

The Significance of Cyperaceae as Weeds 69
Carex deweyana Schwein. wool alien Sell & Murrell, 1996
Carex diandra Schrank ornamental Huxley, 1992
Carex digitata L. ornamental Huxley, 1992; Darke, 1999
Carex dipsacea Berggr. ornamental Grounds, 1989; Huxley, 1992;
Turner & Wasson, 1998
Carex dissita Sol. ex Hook. f. ornamental Huxley, 1992
Carex divulsa Stokes subsp. leersii ornamental Grounds, 1989
(Kneuck.) W. Koch
Carex dolichostachya Hayata ornamental Darke, 1999
Carex eburnea Boott in Hook. ornamental Darke, 1999
Carex elata All. ornamental Grounds, 1989; Ottesen, 1989; Greenlee
&Fell, 1992; Huxley, 1992; Turner & Wasson,
1998; Darke, 1999
Carex exserta Mack. revegetation Ratliff & Westfall, 1992
Carex firma Host ornamental Grounds, 1989; Huxley, 1992; Darke, 1999
Carex flacca Schreb. ornamental Grounds, 1989; Huxley, 1992; Darke, 1999
Carex flaccosperma Dewey ornamental Darke, 1999
Carex flagellifera Colenso ornamental, wool alien Sell & Murrell, 1996; Ottesen, 1989;
Greenlee & Fell, 1992; Turner & Wasson,1998
Carex flava L. ornamental Ottesen, 1989
Carex gallaecica H. Lév. & Vaniot ornamental Bailey, 1935
Carex gaudichaudiana Kük. ornamental Bailey,1935; Huxley,1992
Carex geyeri Boott erosion control Hermann, 1970
Carex grayi J. Carey ornamental Bailey & Bailey, 1976; Grounds, 1989;
Greenlee & Fell, 1992; Turner & Wasson,
1998; Darke, 1999
Carex hachijoensis Akiyama ornamental Grounds, 1989; Greenlee & Fell, 1992;
Huxley, 1992; Darke, 1999
Carex hirta L. ballast Brown, 1880
Carex hoodii Boott in Hook. erosion control Hermann, 1970
Carex hubbardii Nelmes wool alien Sell & Murrell, 1996
Carex humilis Leyss. ornamental Huxley, 1992
Carex intumescens Rudge ornamental Bailey, 1935; Huxley, 1992
Carex inversa R. Br. wool alien Sell & Murrell, 1996
Carex kaloides Petrie ornamental Huxley, 1992
Carex kobomugi Ohwi ballast, planted for dune Champlin, 1994; Mastrogiuseppe, 2002
stabilization
Carex longebrachiata Boeckeler wool alien Sell & Murrell, 1996
Carex longii Mack. hay, pine-bark mulch Bryson, pers. obs.
Carex lupulina Muhl. ex Willd. ornamental Darke, 1999
Carex montana L. ornamental Huxley, 1992; Darke, 1999
Species1Method of dispersal Source

70 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Carex morrowii Boott ornamental Bailey, 1935; Bailey & Bailey, 1976; Everett,
1980–1982; Grounds, 1989; Ottesen, 1989;
Greenlee & Fell, 1992; Huxley, 1992; Turner
&Wasson, 1998; Darke, 1999
Carex muskingumensis Schwein. ornamental Brooklyn Botanical Garden, 1988; Grounds,
1989; Ottesen, 1989; Greenlee & Fell, 1992;
Huxley, 1992; Darke, 1999
Carex nebrascensis Dewey railroad adventive Mühlenbach, 1979
Carex nigra (L.) Reichard ornamental Ottesen, 1989; Greenlee & Fell, 1992;
Huxley, 1992; Darke, 1999
Carex nudata W.Boott in S. Watson ornamental Greenlee & Fell, 1992; Turner & Wasson,
1998; Darke, 1999
Carex oklahomensis Mack. hay, highway construction Bryson et al., 1992, 1996
Carex ornithopoda Willd. ornamental Grounds, 1989; Ottesen, 1989; Greenlee
&Fell, 1992; Huxley,1992; Darke, 1999
Carex oshimensis Nakai ornamental Grounds, 1989; Darke, 1999
Carex pallescens L. ornamental Darke, 1999
Carex paniculata L. ornamental Huxley,1992; Heywood, 1993
Carex pansa L. H. Bailey ornamental Greenlee & Fell, 1992; Darke, 1999
Carex pendula Huds. ornamental Bailey & Bailey, 1976; Everett, 1980–1982;
Grounds, 1989; Ottesen, 1989; Greenlee
&Fell, 1992; Huxley,1992; Darke, 1999;
Reznicek, 2002
Carex pensylvanica Lam. ornamental Darke, 1999
Carex petriei Cheeseman ornamental Grounds, 1989; Greenlee & Fell, 1992;
Huxley, 1992; Darke, 1999
Carex phyllocephala T.Koyama ornamental Grounds, 1989; Greenlee & Fell, 1992;
Darke, 1999
Carex pilulifera L. ornamental Grounds, 1989; Darke, 1999
Carex plantaginea Lam. ornamental Bailey & Bailey, 1976; Grounds, 1989;
Greenlee &Fell, 1992; Huxley,1992
Carex praegracilis W. Boott highway construction and Reznicek et al., 1976; Bruton & Catling,1982;
maintenance, ornamental Cusick, 1984; Reznicek & Catling, 1987;
Darke, 1999
Carex pseudocyperus L. ornamental Brooklyn Botanical Garden, 1988; Greenlee
&Fell, 1992; Huxley, 1992; Darke, 1999
Carex riparia Curtis ornamental Bailey, 1935; Everett, 1980–1982; Grounds,
1989; Huxley,1992; Darke, 1999
Carex scaposa C. B. Clarke ornamental Huxley, 1992
Carex secta Boott wool alien Sell & Murrell, 1996
Carex siderosticta Hance ornamental Grounds, 1989; Greenlee & Fell, 1992;
Huxley, 1992; Darke, 1999
Carex solandri Boott ornamental, wool alien Sell & Murrell, 1996; Darke, 1999
Species1Method of dispersal Source
Appendix 1. Continued.

The Significance of Cyperaceae as Weeds 71
Carex spectabilis Dewey erosion control Hermann, 1970
Carex spissa L. H. Bailey ornamental Greenlee & Fell, 1992; Turner & Wasson,
1998; Darke, 1999
Carex stricta Lam. ornamental Ottesen, 1989; Darke, 1999
Carex sylvatica Huds. ornamental Brooklyn Botanical Garden, 1988; Greenlee
&Fell, 1992; Huxley, 1992
Carex temnolepis Franch. ornamental Greenlee & Fell, 1992
Carex tereticaulis F. Muell. wool alien Sell & Murrell, 1996
Carex testacea Sol. ex Boott ornamental Greenlee & Fell, 1992; Turner & Wasson,
1998; Darke, 1999
Carex texensis (Torr.) L. H. Bailey ornamental Greenlee & Fell, 1992
Carex trifida Cav. ornamental Huxley, 1992
Carex tumulicola Mack. ornamental Greenlee & Fell, 1992; Turner & Wasson,
1998; Darke, 1999
Carex umbrosa Host ornamental Huxley, 1992
Carex uncifolia Cheeseman ornamental Grounds, 1989; Huxley, 1992
Carex uruguensis Boeckeler erosion control Pio Corrêa, 1926–1984
Carex vilmorini Mottet ornamental Greenlee & Fell, 1992
Carex virgata Boott ex Hook. f. wool alien Sell & Murrell, 1996
Carex vulpina L. ornamental Huxley, 1992
Carex vulpinoidea Michx. possibly introduced with Sell & Murrell, 1996
fodder or other seed,
wool alien
Caustis dioica R. Br. ornamental Simpson & Inglis, 2001
Cymophyllus fraserianus (Ker Gawl.) ornamental Bailey, 1935; Bailey & Bailey, 1976; Everett,
Kartesz & Gandhi 1980–1982; Grounds, 1989; Huxley, 1992;
Turner & Wasson, 1998; Darke, 1999
Cyperus adenophorus Schrad. ornamental Everett, 1980–1982
Cyperus aggregatus (Willd.) Endl. ballast, wool alien Britton, 1886; Mohr, 1901; Horvat, 1941;
Sell & Murrell, 1996; Tucker et al., 2002
Cyperus albostriatus Schrad. ornamental, naturalized Bailey, 1935; Bailey & Bailey, 1976; Everett,
ornamental 1980–1982; Greenlee & Fell, 1992; Wilson,
1993; Brickell & Zuk, 1997; Turner &
Wasson, 1998; Darke, 1999
Cyperus alopecuroides Rottb. contaminant of nursery stock Carter et al., 1996
Cyperus alternifolius L. garden escape, naturalized Bailey, 1935; Kern, 1974; Bailey & Bailey,
subsp. flabelliformis Kük. ornamental 1976; Brickell & Zuk,1997; Everett, 1980–
1982; Burkill, 1985; Koyama, 1985; Wagner
et al., 1990; Greenlee & Fell, 1992; Huxley,
1992; Sell & Murrell, 1996; Turner & Wasson,
1998; Darke, 1999
Cyperus bulbosus Vahl ornamental Simpson & Inglis, 2001
Cyperus capitatus Vand. erosion control Simpson & Inglis, 2001
Cyperus chordorrhizus Chiov. erosion control, revegetation Simpson & Inglis, 2001
Species1Method of dispersal Source

72 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Cyperus clarus S. T. Blake wool alien Sell & Murrell, 1996
Cyperus compressus L. ballast, ornamental Smith, 1867; Britton, 1886; Bailey, 1935;
Huxley, 1992; Gordon-Gray, 1995
Cyperus congestus Vahl ornamental, wool alien Huxley, 1992; Sell & Murrell, 1996
Cyperus conglomeratus Rottb. erosion control, ornamental Bailey, 1935; Burkill, 1985
Cyperus croceus Vahl ballast, wool alien Smith, 1867; Britton, 1886; Sell & Murrell,1996
Cyperus cyperinus (Retz.) Suringar ornamental, wool alien Huxley, 1992; Sell & Murrell, 1996
Cyperus dactylotes Benth. wool alien Sell & Murrell, 1996
Cyperus difformis L. rice agriculture Holm et al., 1977; Lipscomb, 1980;
Wagner et al., 1990
Cyperus echinatus (L.) A. W. Wood wool alien Sell & Murrell, 1996
Cyperus elegans L. ornamental Darke, 1999
Cyperus entrerianus Boeckeler highway construction and Carter,1990; Carter & Bryson, 1996
maintenance, rice agriculture
Cyperus eragrostis Lam. naturalized ornamental, Grounds, 1989; Huxley, 1992; Sell & Murrell,
wool and grass-seed alien 1996; Brickell & Zuk, 1997; Darke, 1999
Cyperus erythrorhizos Muhl. ornamental Huxley, 1992
Cyperus esculentus L. cultivated for tubers, Bailey,1935; Bailey & Bailey,1976; Holm et al.,
wool alien 1977; Sell & Murrell, 1996; Turner & Wasson,
1998; Darke, 1999; Miller & Miller, 1999
Cyperus fertilis Boeckeler ornamental Bailey,1935; Huxley,1992
Cyperus filicinus Vahl ornamental Huxley, 1992
Cyperus fuscus L. ballast Smith, 1867; Britton, 1886
Cyperus giganteus Vahl water purification Pio Corrêa, 1926–1984
Cyperus gracilis R. Br. ground cover Hughes, 1995
Cyperus gunnii Hook. f. wool alien Sell & Murrell, 1996
Cyperus haspan L. ornamental, Holm et al., 1997; Everett, 1980–1982;
rice agriculture Darke, 1999
Cyperus hyalinus Vahl air traffic Carter & Mears, 2000
Cyperus imbricatus Retz. ballast McGivney, 1938
Cyperus iria L. rice agriculture Holm et al., 1977; Koyama, 1985
Cyperus jeminicus Rottb. erosion control Simpson & Inglis, 2001
Cyperus laevigatus L. ballast Radford et al., 1968
Cyperus ligularis L. ballast Mohr, 1901; Horvat, 1941
Cyperus longus L. ornamental Bailey,1935; Brickell & Zuk,1997; Darke,1999
Cyperus lucidus R. Br. ornamental Bailey, 1935
Cyperus luzulae (L.) Rottb. ex Retz. wool alien Sell & Murrell, 1996
Cyperus natalensis Hochst. ornamental Bailey, 1935
Species1Method of dispersal Source
Appendix 1. Continued.

The Significance of Cyperaceae as Weeds 73
Cyperus odoratus L. railroad adventive Mühlenbach, 1983
Cyperus owanii Boeckeler naturalized ornamental Bailey & Bailey, 1976; Huxley, 1992
Cyperus papyrus L. naturalized ornamental Bailey, 1935; Bailey & Bailey, 1976; Everett,
1980–1982; Grounds, 1989; Wagner et al.,
1990; Greenlee & Fell, 1992; Huxley, 1992;
Wilson, 1993; Brickell & Zuk, 1997; Turner
&Wasson, 1998; Darke, 1999
Cyperus pilosus Vahl rice agriculture McGivney, 1938; Wagner et al., 1990
Cyperus planifolius Rich. ballast Horvat, 1941
Cyperus plukenetii Fernald attachment to clothing Carter, 1993
Cyperus prolifer Lam. naturalized ornamental Bailey & Bailey, 1976; Everett, 1980–1982;
Greenlee & Fell, 1992; Huxley,1992; Carter
et al.,1996; Brickell & Zuk,1997; Darke, 1999
Cyperus pulcher Thunb. recommended for Gordon-Gray, 1995
cultivation in water
gardens
Cyperus reflexus Vahl wool alien Sell & Murrell, 1996
Cyperus retroflexus Buckley ballast? Horvat, 1941
Cyperus rigidifolius Steud. wool alien Sell & Murrell, 1996
Cyperus rotundus L. agriculture, animals, ballast, Smith, 1867; Britton, 1886; Holm et al.,1977;
machinery, wool alien Sell & Murrell, 1996; Miller & Miller,1999
Cyperus rutilans (C. B. Clarke) wool alien Sell & Murrell, 1996
Maiden & Betche
Cyperus sanguinolentus Vahl highway construction Carter & Bryson, 2000b
and maintenance, rice
agriculture
Cyperus sexangularis Nees water gardens Gordon-Gray, 1995
Cyperus sphacelatus Rottb. ballast Britton, 1886; Mohr, 1901; McGivney,1938
Cyperus sporobolus R. Br.wool alien Sell & Murrell, 1996
Cyperus stoloniferus Retz. erosion control Burkill, 1935
Cyperus strigosus L. ornamental Bailey, 1935
Cyperus subumbellatus Kük. ornamental Simpson & Inglis, 2001
Cyperus surinamensis Rottb. ballast Britton, 1886
Cyperus tenuis Sw. wool alien Sell & Murrell, 1996
Cyperus textilis Thunb. cultivated, presumably Gordon-Gray, 1995
as an ornamental
Cyperus umbellatus Benth. ballast Brown, 1880
Cyperus ustulatus A. Rich. wool alien Sell & Murrell, 1996
Cyperus vaginatus R. Br.wool alien Sell & Murrell, 1996
Desmoschoenus spiralis (A.Rich.) Hook. f.
ornamental Grounds, 1989
Eleocharis acicularis (L.) Roem. rice agriculture, Bailey, 1935; Bailey & Bailey, 1976; Everett,
&Schult. ornamental 1980–1982; Holm et al., 1997; Turner &
Wasson, 1998; Darke, 1999
Species1Method of dispersal Source

74 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Eleocharis dulcis Trin. ex Hensch. cultivated for tubers Kern, 1974; Bailey & Bailey, 1976; Everett,
(Chinese water chestnut), 1980–1982; Gordon-Gray, 1995; Brickell &
rice agriculture Zuk, 1997; Holm et al., 1997; Huxley, 1992;
Turner & Wasson, 1998; Darke, 1999
Eleocharis interstincta (Vahl) ornamental Bailey, 1935
Roem. & Schult.
Eleocharis lanceolata Fernald rice agriculture Smith, 2002c
Eleocharis macrostachya Britton construction equipment Bryson, pers. obs.
Eleocharis montevidensis Kunth ornamental Brickell & Zuk, 1997
Eleocharis nodulosa (Roth) Schult. wool casual Sell & Murrell, 1996
Eleocharis ovata (Roth) Roem. ornamental Bailey, 1935
&Schult.
Eleocharis pachycarpa E. Desv.sheep industry Svenson, 1957
in C. Gay
Eleocharis palustris (L.) Roem. ornamental, rice Huxley, 1992; Holm et al., 1997
&Schult. agriculture
Eleocharis parvula (Roem. & Schult.) ornamental Everett, 1980–1982
Link ex Bluff, Nees & Schauer
Eleocharis pusilla R. Br. ornamental Grounds, 1989
Eleocharis vivipara Link ornamental Everett, 1980–1982; Huxley, 1992
Eriophorum angustifolium Honck. ornamental Everett, 1980–1982; Grounds, 1989; Huxley,
1992; Turner & Wasson,1998; Darke, 1999
Eriophorum chamissonis C. A. Mey.ornamental Huxley, 1992
Eriophorum gracile W.D. J. Koch ornamental Darke, 1999
ex Roth
Eriophorum latifolium Hoppe ornamental Everett, 1980–1982
Eriophorum vaginatum L. ornamental Grounds, 1989; Huxley,1992; Darke,1999
Eriophorum virdicarinatum ornamental Everett, 1980–1982; Grounds, 1989; Huxley,
(Engelm.) Fernald 1992; Darke, 1999
Eriophorum virginicum L. ornamental Darke, 1999
Eriophorum scheuchzeri Hoppe ornamental Huxley,1992
Fimbristylis annua (All.) Roem. rice agriculture Kral, 1971; Holm et al., 1977; Kral, 2002b
&Schult.
Fimbristylis cymosa R. Br. revegetation Fosberg, 1988
Fimbristylis decipiens Kral rice agriculture Kral, 1971
Fimbristylis dichotoma (L.) Vahl rice agriculture Kral, 2002b
Fimbristylis miliacea (L.) Vahl rice agriculture, Burkill, 1935; Kral, 1971; Holm et al., 1977;
soil improvement Koyama, 1985; Kral, 2002b
Fimbristylis pauciflora R. Br.soil improvement Burkill, 1935
Fimbristylis spadicea Vahl ballast Smith, 1867
Fimbristylis squarrosa Vahl ballast Kral, 2002b
Species1Method of dispersal Source
Appendix 1. Continued.

The Significance of Cyperaceae as Weeds 75
Fimbristylis tomentosa Vahl rice agriculture Kral, 2002b
Fimbristylis umbellaris (Lam.) Vahl soil improvement Burkill, 1935
Fimbristylis vahlii (Lam.) Link ballast Smith, 1867
Fuirena squarrosa Michx. ballast Smith, 1867
Fuirena umbellata Rottb. erosion control Burkill, 1935
Gahnia procera J. R. Forst. & G. Forst. ornamental Grounds, 1989
Isolepis cernua (Vahl) Roem. & Schult. ornamental Bailey, 1935; Everett, 1980–1982; Grounds,
1989; Greenlee & Fell, 1992; Huxley, 1992;
Turner & Wasson, 1998; Darke, 1999
Isolepis nodosa (Rottb.) R. Br. ornamental Turner & Wasson, 1998
Isolepis prolifera (Rottb.) R. Br. ornamental Huxley, 1992
Isolepis setacea (L.) R. Br.ornamental Huxley, 1992
Kobresia pygmaea C. B. Clarke erosion control Dickoré, 1994
in Hook. f.
Kyllinga brevifolia Rottb. pine-bark mulch, rice Koyama, 1985; Bryson & Carter, 1992;
agriculture, turfgrass sod, Sell & Murrell, 1996; Holm et al., 1997;
wool alien Bryson et al., 1997
Kyllinga erecta Schumach. wool alien Sell & Murrell, 1996
Kyllinga gracillima Miq. turfgrass sod Bryson et al., 1997
Kyllinga nemoralis (J. R. Forst. &ornamental Bailey, 1935
G. Forst.) Dandy ex Hutch. & Dalziel
Kyllinga odorata Vahl wool alien, turfgrass sod Sell & Murrell, 1996; Bryson et al., 1997
Kyllinga squamulata Thonn. ex Vahl turfgrass sod Bryson et al., 1997
Lepironia articulata (Retz.) Domin fibers in sails and as Mabberley, 1997
packing material
Lipocarpha maculata (Michx.) Torr. ballast Smith, 1867
Lipocarpha micrantha (Vahl) ballast Smith, 1867
G. C. Tucker
Machaerina sinclairii (Hook. f.) Koyama
ornamental Grounds, 1989
Mapania cuspidata (Miq.) Uittien ornamental Bailey, 1935; Simpson, 1992;
Simpson & Inglis, 2001
Mapania mannii C. B. Clarke ornamental Simpson, 1992; Simpson & Inglis, 2001
Mapania palustris (Hassk. ex Steud.) ornamental Simpson, 1992; Simpson & Inglis, 2001
Fern.-Vill.
Mapania pandanophylla (F. Muell.) ornamental Bailey, 1935
K. Schum.
Oreobolus pectinatus Hook. f. ornamental Grounds, 1989
Rhynchospora alba (L.) Vahl ornamental Bailey, 1935
Rhynchospora colorata (L.) H. Pfeiff. ornamental Simpson & Inglis, 2001
Rhynchospora corymbosa (L.) Britton revegetation, soil Burkill, 1935
improvement
Species1Method of dispersal Source

76 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Rhynchospora fusca (L.) W. T. Aiton ornamental Bailey, 1935
Rhynchospora nervosa (Vahl) Boeckeler
ornamental Huxley, 1992; Simpson 1993;
Simpson & Inglis, 2001
Rhynchospora nervosa ornamental Huxley, 1992
subsp. ciliata T. Koyama
Schoenoplectus acutus ornamental Everett, 1980–1982
(Muhl. ex J. M. Bigelow)
Á. Löve & D. Löve
Schoenoplectus californicus erosion control Smith et al., 1993
(C. A. Mey.) Soják
Schoenoplectus grossus (L. f.) Palla rice agriculture Holm et al., 1997
Schoenoplectus heterochaetus ornamental Everett, 1980–1982
(Chase) Soják
Schoenoplectus lacustris (L.) Palla ornamental Bailey, 1935; Everett, 1980–1982
Schoenoplectus lacustris ornamental Everett, 1980–1982;
subsp. validus (Vahl) T. Koyama Turner & Wasson, 1998
Schoenoplectus mucronatus (L.) Palla rice agriculture, planted as Holm et al., 1997; Smith, 2002b
waterfowl food
Schoenoplectus tabernaemontani ornamental Everett, 1980–1982; Grounds, 1989;
(C. C. Gmel.) Palla Greenlee & Fell, 1992; Huxley, 1992;
Turner & Wasson, 1998; Darke, 1999
Schoenus pauciflorus (Hook. f.) Hook. f.
ornamental Grounds, 1989; Huxley, 1992
Scirpus atrovirens Willd.ornamental Bailey,1935; Darke, 1999
Scirpus cyperinus (L.) Kunth ornamental Everett, 1980–1982; Huxley, 1992; Darke,1999
Scirpus divaricatus Elliott railroad adventive Mühlenbach, 1979
Scirpus georgianus R. M. Harper railroad adventive Mühlenbach, 1983
Scirpus holoschoenus L. ornamental Bailey, 1935; Huxley, 1992; Brickell & Zuk,1997
Scirpus pallidus (Britton) Fernald accidental transport Whittemore & Schuyler, 2002
Scirpus pendulus Muhl. accidental transport Whittemore & Schuyler, 2002
Scirpus sylvaticus L. ornamental Huxley, 1992
Trichophorum alpinum (L.) Pers. ornamental Huxley, 1992
Uncinia divaricata W. Boott ornamental Grounds, 1989
Uncinia egmontiana Hamlin ornamental Grounds, 1989; Greenlee & Fell, 1992;
Huxley, 1992
Uncinia rubra Colenso ex Boott ornamental Grounds, 1989; Huxley, 1992;
Brickell & Zuk, 1997
Uncinia uncinata (L. f.) Kük. ornamental Grounds, 1989; Greenlee & Fell, 1992;
Brickell & Zuk, 1997; Turner & Wasson, 1998
1Plant nomenclaturefollows Flora of North America, volume 23; plant names werealso verified through the Missouri Botanical
Garden w3TROPICOS VAST database (rev.1.5) (http://mobot.mobot.org/W3T/Search/vast.html) and the International Plant Names
Index (http://www.ipni.org/index.html). A moreinclusive list of names cited in the references is available from the authors.
Species1Method of dispersal Source
Appendix 1. Continued.

The Significance of Cyperaceae as Weeds 77
APPENDIX 2
Cyperaceous weeds of the world with data on habit, habitat, and distribution.
Species1Source Habit 2Habitat Distribution3
Abildgaardia ovata (Burm. f.) Kral Holm et al., 1979; P pastures, AFR, ASI, AUS,
Soerjani et al., 1987; rice fields CAR, EUR, IND,
Moody, 1989; Kukkonen, NA, PI, SA
2001
Bolboschoenus affinis (Roth) Drobow Kukkonen, 2001 P rice fields EUR, IND
Bolboschoenus caldwellii (V. J. Cook) Kern, 1974; P aquatic, AUS, PI
Soják Simpson & Inglis, 2001 irrigation ditches
Bolboschoenus fluviatilis (Torr.) Soják Holm et al., 1979; P aquatic ASI, AUS, NA
WSSA, 1989
Bolboschoenus maritimus (L.) Palla Kern, 1974; Reed, 1977; P aquatic, crops, AFR, ASI, CAR,
Kühn, 1982; Moody, 1989; rice fields EUR, IND, NA,
Holm et al., 1997; Johnson, PI, SA
1997; Kissman, 1997
Bolboschoenus planiculmis (F. Schmidt) Zhirong et al., 1990 P wetlands, ASI
T. V. Egorova rice fields
Bulbostylis barbata (Rottb.) C. B. Clarke Ohwi, 1965; Lin, 1968; A crops, cultivated AFR, ASI, AUS,
Reed, 1977; Godfrey & fields, fallow CAR, IND, NA,
Wooten, 1979; Holm et al., fields, rice fields, PI, SA
1979; Kühn, 1982; Moody, waste places
1989; Le Bourgeois &
Merlier, 1995; Simpson
&Inglis, 2001
Bulbostylis capillaris (L.) C. B. Clarke Godfrey & Wooten, 1979; A roadsides, NA, SA
Lorenzi, 1982; Moody, waste places
1989; Kissman, 1997
Bulbostylis ciliatifolia (Elliott) Fernald Godfrey & Wooten, 1979 A fallow fields, CAR, NA
roadsides,
waste places
Bulbostylis densa (Wall.) Hand.-Mazz. Ohwi, 1965; Reed, 1977; Aaquatic biotypes, AFR, ASI, AUS,
Kühn, 1982; Moody,1989; crops, cultivated IND, PI
Kukkonen, 2001; Simpson fields, rice fields,
&Inglis, 2001 waste places
Bulbostylis filamentosa (Vahl) C.B. Clarke Healy & Edgar, 1980 P AFR
Bulbostylis hispidula (Vahl) R. W. Haines Kühn, 1982; Le Bourgeois A aquatic biotypes, AFR
&Merlier, 1995; Simpson crops, cultivated
&Inglis, 2001 fields, grasslands
Bulbostylis hispidula subsp.pyriformis Gordon-Gray, 1995 A pioneers or AFR
(Lye) R. W. Haines exposed areas,
weeds of cultivation
Bulbostylis humilis (Kunth) C. B. Clarke Gordon-Gray, 1995 A gardens, AFR
potted plants
Bulbostylis puberula (Poir.) Kunth Holm et al., 1979; rice fields IND
Soerjani et al., 1987;
Moody, 1989
Carex acuta L. Holm et al., 1979 P EUR
Carex albolutescens Schwein. WSSA, 1989 Pmoist soils NA
Carex albula Allan Moore & Edgar, 1970; P crops, grasslands PI
Simpson & Inglis, 2001

78 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Carex aquatilis Wahlenb. Holm et al., 1979; P stream margins, ASI, NA
WSSA, 1989 wetlands
Carex atherodes Spreng. Holm et al., 1979; P wetlands NA
WSSA, 1989
Carex aureolensis Steud. Bryson, pers. obs. P crop borders, NA
lawns, pastures,
waste places
Carex baccans Nees Holm et al., 1979 P PI
Carex biwensis Franch. Reed, 1977 P aquatic ASI
Carex blanda Dewey Bryson, 1985a; DeFelice P lawns, waste NA
&Bryson, 2004 places
Carex bonariensis Desf. Holm et al., 1979; P SA
Kissman, 1997
Carex breviculmis R. Br. Moore & Edgar, 1970; P rice fields, ASI, AUS, PI
Moody, 1989; Simpson & gardens,
Inglis, 2001 grasslands
Carex brevicuspis C. B. Clarke Lin, 1968; Reed, 1977 P ASI
Carex brizoides L. Reed, 1977; Kühn, 1982 P crops, grasslands ASI, EUR
Carex brongniartii Kunth Holm et al., 1979 P SA
Carex buchananii Berggr. Moore & Edgar, 1970; P grasslands PI
Simpson & Inglis, 2001
Carex canescens L. Holm et al., 1979 P wetlands EUR
Carex cherokeensis Schwein. WSSA, 1989; DeFelice & P lawns, pastures NA
Bryson, 2004
Carex comans Berggr. Moore & Edgar, 1970; P gardens, pastures PI
Simpson & Inglis, 2001
Carex comosa Boott Bryson, pers. obs. P wetlands NA
Carex coriacca Hamlin Moore & Edgar, 1970; P grasslands PI
Simpson & Inglis, 2001
Carex dietrichiae Boeckeler Holm et al., 1979 P SA
Carex dimorpholepis Steud. Ohwi, 1965 P wet fields ASI
Carex dispalata Boott ex A. Gray Reed, 1977 P aquatic, ASI
wet/low places
Carex disticha Huds. Holm et al., 1979 P EUR
Carex divisa Huds. Holm et al., 1979; P gardens, rice fields AFR, ASI, AUS
Kukkonen, 2001 (New Zealand),
EUR, IND, NA
Carex divulsa Stokes Holm et al., 1979 P grasslands EUR, PI, SA
Carex eurycarpa T. Holm Holm et al., 1979; P NA
WSSA, 1989
Carex fedia Nees Moody, 1989 P rice fields ASI
Carex flagellifera Colenso Healy & Edgar, 1980; P pastures PI
Parsons & Cuthbertson,1992;
Simpson & Inglis, 2001
Species1Source Habit 2Habitat Distribution3
Appendix 2. Continued.

The Significance of Cyperaceae as Weeds 79
Carex foliosa D. Don Moody, 1989; P rice fields ASI, IND
Kukkonen, 2001
Carex frankii Kunth WSSA, 1989 P crop borders, NA
lawns, pastures,
waste places
Carex gayana E. Desv. Kissman, 1997 P SA
Carex glauca Scop. Holm et al., 1979 P ASI
Carex glaucescens Elliott WSSA, 1989 P pastures, roadsides, NA
waste places
Carex graeffeana Boeckeler Holm et al., 1979 P PI
Carex heterostachya Bunge Zhirong et al., 1990 P field borders, ASI
wetlands
Carex hirta L. Kühn, 1982 P crops, grasslands, AFR, ASI, EUR,
waste places NA
Carex hudsonii A. Benn. Holm et al., 1979 P EUR
Carex inversa R. Br. Healey & Edgar, 1980; P gardens PI
Simpson & Inglis, 2001
Carex iynx Nelmes Healy & Edgar, 1980; P grasslands PI
Simpson & Inglis, 2001
Carex kobomugi Ohwi Small,1954; Svenson,1979; P sandy beaches ASI, NA
Stalter,1980; Standley, 1983
Carex lacustris Willd. Holm et al., 1979; P ditches, roadsides, NA
WSSA, 1989 wetlands
Carex lasiocarpa Ehrh. Holm et al., 1979; Kühn, P aquatic biotypes ASI, EUR, NA
1982; WSSA, 1989;
Simpson & Inglis, 2001
Carex leavenworthii Dewey Bryson, 1985a Plawns, waste places NA
Carex leporina L. Holmet al.,1979; Kühn, P grasslands, AFR, ASI, EUR
1982 waste places
Carex longebrachiata Boeckeler Reed,1977; Simpson & P grasslands, AUS, PI
Inglis, 2001 wet places
Carex longii Mack. Bryson, 1985a P lawns, pastures, NA
waste places
Carex louisianica L. H. Bailey WSSA, 1989 Pditches, right-of- NA
ways, roadsides,
wetlands
Carex lucida Boott Reed, 1977 P low places/ PI
elevations
Carex lupulina Muhl. ex Willd. Holm et al., 1979; P ditches, roadsides, NA
WSSA, 1989 wetlands
Carex macrorrhiza Boeckeler Kissman, 1997 PSA
Carex maorica Hamlin Moore & Edgar,1970; P aquatic, PI
Simpson & Inglis, 2001 irrigation ditches
Carex maximowiczii Miq. Reed, 1977 P wet places ASI
Carex myosurus Nees Holm et al., 1979 P AUS, PI
Species1Source Habit 2Habitat Distribution3

Carex nebrascensis Dewey USDA, 1970; Holm et al., P pastures, roadsides NA
1979; WSSA, 1989
Carex nigra (L.) Reichard Holm et al., 1979 P stream and lake EUR, NA
margins, wetlands
Carex notha Kunth Moody, 1989 P rice fields ASI
Carex nubigena D. Don ex Tilloch & Taylor
Holm et al.,1979; Moody,1989
P rice fields ASI
Carex oklahomensis Mack. Bryson et al., 1992, 1994b, P pastures, roadsides, NA
1996; Standley, 2002 waste places
Carex ovalis Gooden. Kühn,1982; Simpson & P forests, grasslands, EUR, NA
Inglis, 2001 waste places
Carex pallescens L. WSSA, 1989 P lawns, waste places ASI, EUR, NA, PI
Carex panicea L. Kühn, 1982 P aquatic biotypes, AFR, ASI, EUR,
crops, grasslands NA, PI
Carex paniculata L.
Reed,1977; Holm et al.,
P aquatic biotypes, AFR, ASI, AUS,
1979;
Kühn, 1982; Simpson waste places EUR
&Inglis, 2001
Carex philocrena V. I. Krecz Moody, 1989 P rice fields ASI
Carex praegracilis W. Boott Reznicek et al., 1976; P waste places, NA
Swink & Wilhelm, 1979; roadsides
Bruton & Catling, 1982;
Reznicek & Catling, 1987
Carex pruinosa Boott Moody, 1989 P rice fields ASI
Carex pumila Thunb. Lin, 1968; Holm et al.,1979 PASI, PI
Carex pycnostachya Kar. & Kir. Kukkonen, 2001 P rice fields ASI, IND
Carex remota L. Holm et al., 1979 P ASI
Carex rigescens (Franch.) V.I. Krecz. Zhirong et al., 1990 P lawns, orchards, ASI
waste places
Carex riparia Curtis Holm et al., 1979 PEUR
Carex rostrata Stokes ex With. WSSA, 1989 P wetlands
Carex sahnii Ghildyal & U. C. Bhattach. Moody,1989 Price fields ASI
Carex senta Boott Holm et al., 1979; WSSA, P stream and river NA
1989 margins, wetlands
Carex sororia Kunth Kissman, 1997 P SA
Carex spicata Huds. Bryson, pers. obs. P ASI, EUR, NA
Carex testacea Sol. ex Boott Parsons & Cuthbertson, P pastures PI
1992; Simpson & Inglis,2001
Carex thunbergii Steud. Ohwi, 1965; Reed, 1977 P aquatic, rice fields ASI
Carex uruguensis Boeckeler Kissman, 1997 P pastures, roadsides SA
Carex verrucosa Muhl. WSSA, 1989 P roadsides, wet areas NA
Carex vulpina L. Holm et al., 1979 P EUR
Carex vulpinoidea Michx. Moore & Edgar, 1970; P pastures, old fields, EUR, NA, PI, SA
Godfrey & Wooten, 1979; waste places
Simpson & Inglis, 2001
80 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Species1Source Habit 2Habitat Distribution3
Appendix 2. Continued.

The Significance of Cyperaceae as Weeds 81
Cladium jamaicense Crantz Holm et al., 1979 P aquatic NA
Cladium mariscus (L.) Pohl Holm et al.,1979; Kühn, P aquatic biotypes, AFR, ASI, EUR,
1982; Moody,1989; rice fields IND, NA, PI, SA
Simpson & Inglis, 2001
Courtoisina cyperoides (Roxb.) Soják Simpson & Koyama, 1998; A rice fields AFR, ASI, IND
Simpson & Inglis, 2001
Cyperus acuminatus Torr. & Hook. DeFelice & Bryson, 2004 A field borders, NA
pastures, roadsides,
wet clay soils
Cyperus aggregatus (Willd.) Endl. Holm et al., 1979; Kühn, P crops, sandy soil, AUS, CAR, NA,
1982; Lorenzi, 1982; waste places SA
WSSA, 1989; Wilson,
1993; Kissman, 1997;
Simpson & Inglis, 2001
Cyperus albostriatus Schrad. Healy & Edgar, 1980; P gardens, AUS, PI
Simpson & Inglis, 2001 waste places
Cyperus alopecuroides Rottb. Holm et al., 1979; Kühn, Paquatic biotypes, AFR, ASI, AUS,
1982; Moody,1989; rice fields IND, NA, PI, SA
Carter et al., 1996;
Bryson et al., 1998;
Simpson & Inglis, 2001
Cyperus alternifolius L. subsp.
Kern,1974; Holm et al.,1979;
P aquatic biotypes, AFR, ASI, EUR,
flabelliformis Kük. Kühn, 1982; Moody, 1989; rice fields, IND, NA, PI, SA
Kissman, 1997; Simpson & waste places
Koyama, 1998; Simpson &
Inglis, 2001
Cyperus alulatus J. Kern
Moody,1989; Kukkonen,2001
A rice fields ASI, IND
Cyperus amabilis Vahl Holm et al., 1979; Kühn, A grasslands, AFR, ASI, IND,
1982; Le Bourgeois & Merlier, waste places SA
1995; Simpson & Inglis,2001
Cyperus amuricus Maxim. Ohwi, 1965; Reed, 1977; A/P cultivated fields, ASI
Holm et al., 1979 waste places,
wet places
Cyperus arenarius Retz. Simpson & Inglis, 2001 P unspecified ASI, IND
Cyperus articulatus L. Holm et al., 1979; Kühn, Paquatic biotypes, AFR, ASI, AUS,
1982; Moody, 1989; crops, rice fields IND, NA, SA
WSSA, 1989; Kissman,
1997; Bryson et al.,1998;
Simpson & Inglis, 2001
Cyperus babakan Steud. Kern, 1974; Holm et al., P rice fields ASI, IND, PI
1979; Soerjani et al.,
1987; Moody, 1989
Cyperus bifax C. B. Clarke Koyama, 1985; Moody, P ditches, irrigated ASI, AUS, IND
1989; Wilson, 1993 cultivation, open wet
ground, rice fields
Cyperus boreohemisphaericus Lye Simpson & Inglis, 2001 crops AFR
Cyperus bulbosus Vahl Terry, 1976; Reed, 1977; P grasslands, AFR, ASI, AUS,
Holm et al., 1979; Moody,rice fields IND
1989; Kukkonen, 2001;
Simpson & Inglis, 2001
Cyperus castaneus Willd. Holm et al., 1979; A rice fields ASI
Moody, 1989
Species1Source Habit 2Habitat Distribution3

Cyperus cephalotes Vahl Soerjani et al., 1987; P rice fields ASI, IND
Moody, 1989
Cyperus compactus Retz. Kern,1974; Holm et al., 1979; P aquatic biotypes, ASI, AUS, IND,
Kühn, 1982; Soerjani et al., crops, ditches, PI
1987; Moody, 1989; Kukkonen, rice fields, waste
2001; Simpson & Inglis, 2001 places, wet places
Cyperus compressus L. Ohwi,1965; Lin,1968; Kern, A crops, fallow rice AFR, ASI, AUS,
1974; Godfrey & Wooten, fields, gardens, IND, NA, PI, SA
1979; Kühn, 1982; Koyama, grasslands, lawns,
1985; Soerjani et al., 1987; roadsides, waste
Moody, 1989; WSSA, 1989; places
Wagner & Herbst, 1995;
Bryson et al.,1998; Kukkonen,
2001; Simpson & Inglis,2001;
Ravi & Mohanan, 2002;
DeFelice & Bryson, 2004
Cyperus congestus Vahl Healey & Edgar, 1980; P cultivation, damp AFR, AUS
Wilson, 1993; Gordon-Gray,ground, disturbed
1995; Simpson & Inglis, areas, ditches,
2001 gardens, roadsides
Cyperus conglomeratus Rottb. Moody,1989; Simpson & P crops, rice fields AFR, IND
Inglis, 2001
Cyperus corymbosus Rottb. Holm et al., 1979; Moody, P rice fields ASI, IND, SA
1989; Simpson & Inglis, 2001
Cyperus crassipes Vahl Reed,1977; Holm et al.,1979 P AFR
Cyperus croceus Vahl Godfrey & Wooten, 1979; P pastures, turf, CAR, NA, SA
WSSA, 1989; Bryson et al., waste places
1998
Cyperus cuspidatus Kunth Kern, 1974; Godfrey & A aquatic biotypes, AFR, ASI, AUS,
Wooten, 1979; Kühn,1982; crops, fallow IND, NA, PI, SA
Moody, 1989; WSSA, fields, rice fields,
1989; Kukkonen, 2001; sandy fields, waste
Simpson & Inglis, 2001 places, wet places
Cyperus cyperinus (Retz.) Suringar Lin, 1968; Reed, 1977; P cultivated lands, ASI, IND, PI
Simpson & Koyama, 1998; gardens, rice
Simpson & Inglis, 2001 fields, waste places
Cyperus cyperoides (L.) Kuntze Kern, 1974; Terry, 1976; P crops, disturbed AFR, ASI, AUS,
Kühn, 1982; Soerjani et al., sites, fallow fields, CAR, IND, PI
1987;Moody, 1989; Zhirong fields, gardens,
et al., 1990; Gordon-Gray, grasslands, rice
1995; Le Bourgeois & Merlier, fields, roadsides,
1995; Johnson,1997; Simpson waste places
&Koyama,1998; Kukkonen,
2001; Simpson & Inglis, 2001
Cyperus cyperoides subsp. macrocarpus Terry, 1976 P crops AFR
(Kunth) Lye
Cyperus denudatus L. f. Simpson & Inglis, 2001 P rice fields AFR
Cyperus diandrus Torr.Holm et al., 1979 Awet areas NA
Cyperus diaphanus Schrad. ex Kern, 1974; Moody, 1989 A rice fields ASI, IND, PI
Roem. & Schult.
82 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Species1Source Habit 2Habitat Distribution3
Appendix 2. Continued.

The Significance of Cyperaceae as Weeds 83
Cyperus difformis L. Ohwi, 1965; Lin,1968; Kern, A aquatic biotypes, AFR, ASI, AUS,
1974; Terry, 1976; Reed, crops, grasslands, EUR, IND, NA,
1977; Holm et al., 1977, rice fields PI, SA
1979; DeFilipps, 1980c;
Moody, 1981; Kühn,1982;
Koyama, 1985; Akobundu
&Agyakwa, 1987; Soerjani
et al.,1987; Moody, 1989;
WSSA,1989; Zhirong et al.,
1990; Gordon-Gray, 1995;
Johnson,1997; Kissman,1997;
Bryson et al.,1998; Kukkonen,
2001; Simpson & Inglis,
2001; Ravi & Mohanan,
2002; Carter, 2005
Cyperus diffusus Vahl Cardenas et al., 1972; P gardens, low ASI, CAR, IND,
Reed, 1977; Holm et al., elevations, rice PI, SA
1979; Moody,1989; fields, warm
Kissman, 1997; Simpson regions
&Koyama, 1998; Simpson
&Inglis, 2001; DeFelice
&Bryson, 2004
Cyperus digitatus Roxb. Kern, 1974; Holm et al., P aquatic biotypes, AFR, ASI, AUS,
1979; Kühn, 1982; Soerjani crops, rice fields, IND, NA, PI, SA
et al., 1987; Moody, 1989; waste places
Simpson & Koyama, 1998;
Kukkonen, 2001; Simpson
&Inglis, 2001
Cyperus dilatatus Schumach. Simpson & Inglis, 2001 P cultivated fields, AFR
gardens
Cyperus distans L. f. Kern, 1974; Terry, 1976; P aquatic biotypes, AFR, ASI, AUS,
Holm et al., 1979; Kühn, crops, grasslands, CAR, EUR, IND,
1982; Soerjani et al., 1987; rice fields, waste NA, PI, SA
Moody,1989; Gordon-Gray,places
1995; Johnson,1997; Kissman,
1997; Simpson & Koyama,
1998; Simpson & Inglis, 2001
Cyperus distinctus Steud. Carter, pers. obs. P ditches, roadsides, CAR, NA
waste places
Cyperus dives Delile Gordon-Gray, 1995 P sugarcane fields AFR
Cyperus dubius Rottb. Moody, 1989; Gordon-Gray, P open rice fields, AFR, IND
1995; Simpson & Inglis, 2001 sandy sites
Cyperus duclouxii E. G. Camus Zhirong et al., 1990 P ASI
Cyperus echinatus (L.) A. W. Wood WSSA, 1989; Bryson et al., P ditches, pastures, NA
1998 roadsides, waste
places
Cyperus elatus L. Kern, 1974; Soerjani et al., P rice fields ASI, IND, PI
1987; Moody,1989; Simpson
&Inglis, 2001
Cyperus elegans L. Bryson et al., 1998 Pwaste places CAR, NA
Cyperus entrerianus Boeckeler Kissman, 1997; Carter, 1990; P crops, pastures, NA, SA
Carter & Jones, 1991; Bryson roadsides
&Carter, 1994, 1996; Bryson
et al.,1998; Simpson & Inglis,
2001; DeFelice & Bryson, 2004
Species1Source Habit 2Habitat Distribution3

Cyperus eragrostis Lam. Holm et al., 1979; Parsons P/A? disturbed or AFR, AUS, EUR,
&Cuthbertson, 1992; fallow areas, NA, PI, SA
Wilson, 1993; Gordon-Gray, pastures, rice
1995; Kissman, 1997; fields, roadsides
Bryson et al., 1998;
Simpson & Koyama, 1998;
Simpson & Inglis, 2001
Cyperus erythrorhizos Muhl. Holm et al., 1979; Moody, A rice fields, wet ASI, NA
1989; WSSA, 1989; Bryson areas
et al., 1998; DeFelice &
Bryson, 2004
Cyperus esculentus L. USDA, 1970; Kern, 1974; P crops, fields, AFR, ASI, AUS,
Terry, 1976; Holm et al., irrigated fields, CAR, EUR, IND,
1977; Godfrey & Wooten, pastures, rice NA, PI, SA
1979; Holm et al., 1979; fields, turf,
Kühn, 1982; Lorenzi, 1982; waste places
Moody, 1989; WSSA, 1989;
Gordon-Gray, 1995;
Le Bourgeois & Merlier,
1995; Johnson, 1997;
Kissman, 1997; Bryson
et al., 1998; Simpson &
Koyama, 1998; Kukkonen,
2001; Simpson & Inglis,
2001; DeFelice & Bryson,
2004; Rzedowski & Rzedowski,
2004; Carter, 2005
Cyperus exaltatus Retz. Holm et al., 1979; Koyama, P low wet sites, AFR, AUS, IND
1985; Moody,1989; Johnson,
rice fields
1997; Kukkonen, 2001;
Simpson & Inglis, 2001
Cyperus fasciculatus Elliott Holm et al., 1979 ASI
Cyperus flavescens L. Holm et al., 1979; Soerjani A crops, pastures, AFR, ASI, CAR,
et al., 1987; Gordon-Gray, rice fields, EUR, IND, NA,
1995; Johnson, 1997; roadside ditches, SA
Kukkonen, 2001 seeps, turf,
waste places
Cyperus flavicomus Michx. Godfrey & Wooten, 1979; A crops, pastures, AFR, ASI, NA,
Holm et al., 1979; Johnson, rice fields, turf, SA
1997; Bryson et al., 1998 waste places
Cyperus flavidus Retz.
Lin, 1968; Kern, 1974; Reed,
A/P crops, fallow AFR, ASI, AUS,
1977; Holm et al., 1979; fields, rice fields, IND, PI
Moody,1989; Zhirong et al., wet places
1990; Simpson & Koyama,
1998; Simpson & Inglis, 2001
Cyperus floribundus (Kük.) Carter & Jones, 1997; A/P agricultural fields, NA
R. Carter & S. D. Jones Carter,pers. obs. disturbed sites,
roadsides
Cyperus foliaceus C. B. Clarke Simpson & Inglis, 2001 A rice fields AFR
Cyperus friburgensis Boeckeler Kissman, 1997 P SA
Cyperus fulvus R. Br.Holm et al., 1979 P SA
Cyperus fuscus L. Holm et al., 1979; WSSA, Acrops, moist AFR, ASI, EUR,
1989; Zhirong et al., 1990; fields, rice fields, NA
Kukkonen, 2001; Simpson wet areas
&Inglis, 2001
84 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Species1Source Habit 2Habitat Distribution3
Appendix 2. Continued.

The Significance of Cyperaceae as Weeds 85
Cyperus giganteus Vahl Kissman, 1997 P SA
Cyperus glaber L. Kukkonen, 2001 A moist fields ASI, EUR, IND
Cyperus glomeratus L. Zhirong et al., 1990; A/P rice fields, ASI, EUR, IND
Kukkonen, 2001 wetlands
Cyperus gracilis R. Br. Holm et al., 1979 P AUS, PI
Cyperus gracilinux C. B. Clarke Holm et al., 1979 P AFR
Cyperus grandibulbosus C. B. Clarke Terry, 1976; Simpson & P unspecified AFR
Inglis, 2001
Cyperus hakonensis Franch. & Sav. Holm et al., 1979 ASI
Cyperus haspan L. Ohwi, 1965; Lin, 1968; A/P grasslands, crops, AFR, ASI, AUS,
Kern, 1974; Holm et al., aquatic biotypes, IND, NA, PI, SA
1979; Kühn,1982; Koyama,
rice fields
1985; Akobundu & Agyakwa,
1987; Soerjani et al., 1987;
Moody, 1989; Zhirong et al.,
1990; Johnson, 1997;
Kissman, 1997; Simpson &
Koyama, 1998; Kukkonen,
2001; Simpson & Inglis,
2001; Ravi & Mohanan, 2002
Cyperus hermaphroditus (Jacq.) Standl. Holm et al., 1979; Kissman, P NA, SA
1997
Cyperus hyalinus Vahl WSSA, 1989; Simpson & A gardens, turf AFR, ASI, AUS,
Inglis, 2001 IND, NA
Cyperus imbricatus Retz. Kern, 1974; Reed, 1977; P aquatic biotypes, AFR, ASI, CAR,
Holm et al., 1979; Kühn, crops, rice fields IND, NA, PI, SA
1982; Soerjani et al., 1987;
Moody, 1989; Zhirong et al.,
1990; Kissman, 1997;
Simpson & Inglis, 2001
Cyperus intactus Vahl Gordon-Gray, 1995 A/P disturbed grasslands AFR
Cyperus iria L. Ohwi, 1965; Lin, 1968; A/P aquatic biotypes, AFR, ASI, AUS,
Kern, 1974; Holm et al., crops, rice fields, CAR, EUR, IND,
1977, 1979; Moody,1981; waste places NA, PI, SA
Kühn, 1982; Lorenzi, 1982;
Koyama, 1985; Akobundu
&Agyakwa, 1987; Soerjani
et al., 1987; Moody, 1989;
WSSA, 1989; Zhirong et al.,
1990; Gordon-Gray, 1995;
Johnson, 1997; Kissman,
1997; Bryson et al., 1998;
Simpson & Koyama, 1998;
Kukkonen, 2001; Simpson &
Inglis, 2001; Ravi & Mohanan,
2002; DeFelice & Bryson,
2004; Carter, 2005
Cyperus javanicus Houtt. Holm et al., 1979; Moody, P rice fields ASI, IND, PI
1989
Cyperus laetus J. Presl & C. Presl Holm et al., 1979; Kissman, P SA
1997
Cyperus laevigatus L. Holm et al., 1979; Kühn, Paquatic biotypes, AFR, ASI, AUS,
1982; Moody,1989; rice fields CAR, EUR, IND,
Simpson & Inglis, 2001 NA, PI, SA
Species1Source Habit 2Habitat Distribution3

Cyperus lanceolatus Poir. Holm et al., 1979; Lorenzi, P ditches, rice fields, AFR, NA, SA
1982; Akobundu & roadsides
Agyakwa, 1987; Johnson,
1997; Kissman, 1997
Cyperus latifolius Poir. Simpson & Inglis, 2001 P pastures AFR
Cyperus lecontei Torr. ex Steud. WSSA, 1989; Bryson et al., P shorelines, NA
1998 waste places
Cyperus ligularis L. Holm et al., 1979; Kissman, P AFR, ASI, NA,
1997 SA
Cyperus longibracteatus (Cherm.) Kük. Akobundu & Agyakwa,1987 P forests, rice fields AFR
Cyperus longus L. Terry, 1976; Holm et al., P aquatic biotypes, AFR, ASI, EUR,
1979; Moody,1989; crops, rice fields IND
Kukkonen, 2001; Simpson
&Inglis, 2001
Cyperus luzulae (L.) Rottb. ex Retz. Cardenas et al., 1972; Holm Pcultivated fields, SA
et al., 1979; Lorenzi, 1982; rice fields
Moody, 1989; Kissman,1997;
Simpson & Inglis, 2001
Cyperus macrostachyos Lam. Moody, 1989 rice fields IND
Cyperus malaccensis Lam. Holm et al.,1979; Kühn,1982; P aquatic biotypes, ASI, AUS, IND,
Soerjani et al.,1987;Moody, cultivated fields, PI
1989; Simpson & Inglis,2001 rice fields
Cyperus mapanioides C. B. Clarke Simpson & Koyama, 1998; Pcrops, cultivated AFR
Simpson & Inglis, 2001 fields
Cyperus maranguensis K. Schum. Terry, 1976; Holm et al., P cultivated fields AFR
1979; Simpson & Inglis, 2001
Cyperus meyenianus Kunth Kissman, 1997 P SA
Cyperus michelianus (L.) Link. subsp.
Kern,1974; Holm et al.,1979;
Aditches, cultivated AFR, ASI, AUS,
pygmaeus (Rottb.) Asch. & Graebn. Soerjani et al.,1987; Moody,fields, fallow rice EUR, IND, PI
1989; Zhirong et al., 1990; fields, gardens,
Kukkonen, 2001; Simpson & rice fields
Inglis, 2001
Cyperus microiria Steud. Ohwi, 1965; Holm et al., A crops, cultivated ASI, IND, NA
1979; Moody, 1989 fields, rice fields,
wetlands
Cyperus mirus C. B. Clarke Wilson, 1993 P gardens AUS
Cyperus mitis Steud. Moody, 1989 rice fields ASI
Cyperus mutisii (Kunth) Andersson Holm et al., 1979 P NA, SA
Cyperus nipponicus Franch. & Sav.
Reed,1977; Holm et al.,1979
A crops, waste places ASI
Cyperus niveus Retz. Reed, 1977; Moody, 1989 P open forests, rice ASI, IND
fields, wet places
Cyperus novae-hollandiae Boeckeler Holm et al., 1979 AUS
Cyperus nutans Vahl Kern, 1974; Moody, 1989; P crops, rice fields AFR, ASI, IND,
Simpson & Inglis, 2001 PI
Cyperus obtusiflorus Vahl Simpson & Inglis, 2001 P gardens AFR
86 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Species1Source Habit 2Habitat Distribution3
Appendix 2. Continued.

The Significance of Cyperaceae as Weeds 87
Cyperus ochraceus Vahl Bryson et al., 1996, 1998 P ditches, roadsides, CAR, NA, SA
waste places
Cyperus odoratus L. Lin, 1968; Cardenas et al., A/P aquatic biotypes, AFR, ASI, AUS,
1972; Kern, 1974; Holm crops, rice fields, CAR, EUR, IND,
et al., 1979; Kühn, 1982; taro paddies, NA, PI, SA
Lorenzi,1982; Soerjani waste places
et al., 1987; Moody,1989;
WSSA,1989;Wagner et al.,
1990; Kissman,1997;
Bryson et al., 1998; Simpson
&Koyama, 1998; Simpson
&Inglis, 2001; DeFelice &
Bryson, 2004
Cyperus oxylepis Nees ex Steud. Godfrey & Wooten, 1979; P mechanically NA
WSSA, 1989; Bryson et al., disturbed sites,
1996, 1998 waste places,
roadsides
Cyperus pangorei Rottb. Holm et al., 1979; Moody, P rice fields AFR, IND
1989
Cyperus papyrus L. Holm et al., 1979; Simpson P aquatic AFR, NA
&Inglis, 2001
Cyperus pilosus Vahl Lin, 1968; Kern, 1974; Reed, P crops, rice fields, ASI, IND, NA, PI
1977; Soerjani et al., 1987; waste places
Moody, 1989; Zhirong et al.,
1990
Cyperus platystylis R. Br. Kern, 1974; Soerjani et al., P rice fields ASI, AUS, IND,
1987; Moody,1989 PI
Cyperus pohlii (Nees) Steud. Kissman, 1997 P SA
Cyperus polystachyos Rottb. Lin,1968; Kern,1974; Reed, A/P aquatic biotypes, AFR, ASI, AUS,
1977; Holm et al., 1979; crops, fallow rice EUR, IND, NA,
Kühn, 1982; Soerjani et al., fields, grasslands, PI, SA
1987; Moody,1989; Kissman,
waste places,
1997; Kukkonen, 2001; wet places
Simpson & Inglis, 2001;
Ravi & Mohanan, 2002
Cyperus polystachyos var.texensis WSSA, 1989; Bryson et al., A/P crops, grasslands, NA
(Torr.) Fernald 1998 lawns, rice fields,
roadsides, waste
places, wet places
Cyperus procerus Rottb. Kern, 1974; Holm et al., P rice fields ASI, AUS, IND,
1979; Soerjani et al., 1987; PI
Moody, 1989
Cyperus prolifer Lam. WSSA, 1989; Carter et al., P aquatic AFR, NA
1996; Bryson et al., 1998
Cyperus prolixus Kunth Kissman, 1997 P aquatic NA, SA
Cyperus pseudosomaliensis Kük. Simpson & Inglis, 2001 P gardens AFR
Cyperus pseudovegetus Steud. WSSA, 1989; Bryson et al., P field borders, NA, SA
1998; Ramos et al., 2004 pastures, wet soil
Cyperus pulcherrimus Willd. ex Kunth Kern, 1974; Holm et al., P rice fields ASI, IND, PI
1979; Soerjani et al., 1987;
Moody, 1989
Species1Source Habit 2Habitat Distribution3

Cyperus pumilus L.
Kern, 1974; Holm et al.,1979;
A disturbed sandy AFR, ASI, AUS,
Soerjani et al.,1987; Moody, soils, fallow fields, CAR, IND, NA,
1989; Kukkonen, 2001; Ravi rice fields, waste PI
&Mohanan, 2002 lands
Cyperus puncticulatus Vahl Ravi & Mohanan, 2002 A rice fields ASI, IND
Cyperus pustulatus Vahl Johnson, 1997; Simpson A aquatic, rice fields AFR
&Inglis, 2001
Cyperus radians Nees & Meyen Holm et al., 1979; Kühn, P aquatic biotypes, ASI, PI
1982; Moody, 1989; rice fields,
Simpson & Inglis, 2001 waste places
Cyperus reduncus Hochst. ex Boeckeler Holm et al., 1979; Simpson A AFR
&Inglis, 2001
Cyperus reflexus Vahl Kissman, 1997 P pastures, NA, SA
waste places
Cyperus remotispicatus S. S. Hooper Simpson & Inglis, 2001 rice fields AFR
Cyperus retroflexus Buckley Carter et al., 1987; P sandy waste places NA
Carter & Bryson,1991a, b;
Carter & Jones, 1997
Cyperus retrorsus Chapm. WSSA, 1989; Bryson et al., P grasslands, turf, NA
1998 waste places
Cyperus rigidifolius Steud. Terry,1976; Holm et al., P crops, disturbed AFR
1979; Gordon-Gray, 1995; grassland, gardens,
Simpson & Inglis, 2001 lawns
Cyperus rotundus L. Elliott, 1821; Lin, 1968; Pcrops, gardens, AFR, ASI, AUS,
USDA,1970; Cardenas et al., field crops, CAR, EUR, IND,
1972; Kern,1974; Terry,1976; grasslands, lawns, NA, PI, SA
Holm et al., 1977; Godfrey pastures, rice fields,
&Wooten,
1979; Holm et al.,
roadsides, taro,
1979; Moody,
1981, 1989; turf, waste places
Kühn,1982; Lorenzi, 1982;
Koyama,1985; Soerjani et al.,
1987; WSSA, 1989; Zhirong
et al., 1990; Wilson,1993;
Hughes,1995; Le Bourgeois
&Merlier,1995; J
ohnson,1997;
Kissman,1997; Waterhouse,
1997; Bryson et al., 1998; Simpson
&Koyama, 1998; Kukkonen,
2001; Simpson & Inglis, 2001;
Ravi & Mohanan, 2002; DeFelice
&Bryson, 2004; Ramos et al.,
2004; Rzedowski & Rzedowski,
2004; Carter, 2005
Cyperus rubicundus Vahl Moody, 1989 rice fields IND
Cyperus sanguinolentus Vahl Lin,1968; Kern,1974; Reed, A/P aquatic biotypes, AFR, ASI, AUS,
1977; Holm et al.,1979; crops, grasslands, IND, NA, PI
Kühn, 1982; Soerjani et al., rice fields, wet
1987; Moody,1989; Zhirong places
et al.,1990; Carter& Bryson,
2000b; Kukkonen, 2001;
Simpson & Inglis, 2001
Cyperus schweinfurthianus Boeckeler Holm et al., 1979 PAFR
88 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Species1Source Habit 2Habitat Distribution3
Appendix 2. Continued.

The Significance of Cyperaceae as Weeds 89
Cyperus seemannianus Boeckeler Holm et al., 1979 PI
Cyperus serotinus Rottb. Reed, 1977; Holm et al., P aquatic biotypes, AFR, ASI, EUR,
1979; Kühn, 1982; Moody, crops, rice fields, IND, NA
1989; Zhirong et al., 1990; wet places
Kukkonen, 2001; Simpson
&Inglis, 2001
Cyperus seslerioides Kunth Holm et al., 1979 P NA
Cyperus soyauxii Boeckeler Simpson & Inglis, 2001 P cultivated fields AFR
Cyperus sphacelatus Rottb. Kern, 1974; Reed, 1977; A/P aquatic biotypes, AFR, ASI, AUS,
Holm et al., 1979; Kühn, crops, grasslands, IND, NA, PI, SA
1982; Soerjani et al.,1987; rice fields, waste
Moody, 1989; Carter et al., places
1996; Johnson, 1997;
Kissman, 1997; Bryson et al.,
1998; Simpson & Koyama,
1998; Simpson & Inglis,2001
Cyperus squarrosus L. Holm et al., 1979; Kühn, A aquatic biotypes, AFR, ASI, AUS,
1982; Moody, 1989; crops, forests, EUR, IND, NA,
Le Bourgeois & Merlier, gardens, grass- SA
1995; Kukkonen, 2001; lands, rice fields,
Simpson & Inglis, 2001 waste places
Cyperus stenophyllus J. V. Suringar Moody, 1989 rice fields PI
Cyperus stoloniferus Retz. Moody, 1989 P rice fields ASI
Cyperus strigosus L. Holm et al., 1979; Moody, P crops, pastures, ASI, EUR, NA,PI
1989; WSSA, 1989; Bryson roadsides, wet areas
et al., 1998; Simpson & Inglis,
2001; DeFelice & Bryson, 2004
Cyperus substramineus Kük. Kern, 1974; Moody, 1989; A/P rice fields ASI, IND
Ravi & Mohanan, 2002
Cyperus sulcinux C. B. Clarke Kern, 1974; Moody, 1989 A fields, rice fields, ASI, AUS, IND,
roadsides PI
Cyperus surinamensis Rottb. WSSA,1989; Kissman, A/P SA
1997; Bryson et al.,1998
Cyperus tegetiformis Roxb.
Holm et al.,1979; Moody,1989
rice fields AFR, ASI, IND
Cyperus tegetum Roxb.
Holm et al.,1979; Moody,1989
rice fields ASI, IND
Cyperus tenellus L. f. Moore & Edgar, 1970; gardens, irrigation AFR, AUS, PI
Simpson & Inglis, 2001 ditches
Cyperus tenuiculmis Boeckeler Kern, 1974; Holm et al., P fallow fields, AFR, ASI, AUS,
1979; Soerjani et al.,1987; gardens, grass- PI
Moody, 1989; Simpson & lands, rice fields
Inglis, 2001
Cyperus tenuis Sw. Holm et al.,1979; Johnson, P rice fields SA
1997; Simpson & Inglis, 2001
Cyperus tenuispica Steud.
Kern,1974; Holm et al.,1979;
A/P cultivated fields, AFR, ASI, IND,
Koyama,1985; Soerjani et al., rice fields, wet PI
1987; Moody,1989; Simpson places
&Koyama,1998; Kukkonen,
2001; Simpson & Inglis, 2001;
Ravi & Mohanan, 2002
Cyperus trialatus (Boeckeler) J. Kern Kern, 1974; Holm et al., P rice fields, ASI, IND, PI
1979; Moody, 1989 roadsides
Species1Source Habit 2Habitat Distribution3

Cyperus truncatus C. A. Mey. ex Turcz. Holm et al., 1979 ASI
Cyperus uncinatus Poir. Holm et al., 1979; Moody, rice fields PI
1989; Simpson & Inglis, 2001
Cyperus ustulatus A. Rich. Moore & Edgar, 1970; Terry, pastures PI
1976; Simpson & Inglis, 2001
Cyperus virens Michx. WSSA, 1989; Kissman, P NA, SA
1997; Bryson et al., 1998
Cyperus vorsteri K. L. Wilson Gordon-Gray, 1995 P aggressive weed in AFR
parks and gardens
Cyperus zollingeri Steud. Kern,1974; Holm et al.,1979; A crops, rice fields, AFR, ASI, AUS,
Moody, 1989; Simpson & roadsides PI
Inglis, 2001
Eleocharis acicularis (L.) Roem. & Schult. Ohwi,1965; Lin,1968; Kern, P aquatic biotypes, AFR, ASI, AUS,
1974; Holm et al.,1979,1997; crops, grasslands, EUR, IND, NA,
Kühn, 1982; Soerjani et al., rice fields PI, SA
1987; Moody, 1989; WSSA,
1989; Simpson & Inglis, 2001
Eleocharis acuta R. Br. Holm et al., 1979 P AUS
Eleocharis acutangula (Roxb.) Schult. Lin, 1968; Kern, 1974; Reed, P aquatic biotypes, AFR, ASI, AUS,
1977; Holm et al.,1979; Kühn, crops, rice fields IND, NA, PI, SA
1982; Koyama,1985; Soerjani
et al.,1987; Moody, 1989;
Simpson & Koyama, 1998;
Simpson & Inglis, 2001
Eleocharis albida Torr. Carter, 2005 P disturbed CAR, NA
saltmarsh, ditches
Eleocharis atropurpurea (Retz.) Kunth Kern, 1974; Holm et al., A aquatic biotypes, AFR, ASI, AUS,
1979; Kühn, 1982; Soerjani crops, rice fields EUR, IND, NA,
et al., 1987; Moody, 1989; PI, SA
Kukkonen, 2001; Simpson
&Inglis, 2001; Ravi &
Mohanan, 2002
Eleocharis attenuata (Franch. & Sav.) Holm et al.,1979; Moody, P/A? rice fields ASI
Palla 1989
Eleocharis baldwinii (Torr.) Chapm. WSSA, 1989 A wet places NA
Eleocharis cellulosa Torr. Holm et al.,1979; WSSA,1989 P brackish wet places CA, NA, SA
Eleocharis complanata Boeckeler Johnson, 1997 Price fields AFR
Eleocharis congesta D. Don Lin,1968; Kern,1974; Holm A/P rice fields ASI, IND
et al.,1979; Koyama,1985;
Soerjani et al., 1987; Moody,
1989; Kukkonen, 2001;
Simpson & Inglis, 2001
Eleocharis dulcis Trin. ex Hensch. Lin,1968; Reed,1977; Holm P aquatic biotypes, AFR, ASI, AUS,
et al., 1979, 1997; Kühn, crops, fallow CAR, IND, NA,
1982; Soerjani et al., 1987; fields, rice fields PI
Moody, 1989; Simpson & Inglis,
2001; Ravi & Mohanan, 2002
Eleocharis elegans (Kunth) Roem. Kühn, 1982; Lorenzi, 1982; P aquatic biotypes, CAR, NA, PI, SA
&Schult. Kissman, 1997; Simpson & crops, rice fields
Inglis, 2001
90 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Species1Source Habit 2Habitat Distribution3
Appendix 2. Continued.

The Significance of Cyperaceae as Weeds 91
Eleocharis equisetina J. Presl & C. Presl Holm et al., 1979 P ASI, AUS, PI
“Eleocharis erecta Schumac.” Holm et al., 1979 AFR
Eleocharis filiculmis Kunth Reed, 1977; Kühn, 1982; P aquatic, wet places AFR, NA, SA
Lorenzi, 1982
Eleocharis flavescens (Poir.) Holm et al., 1979; Walters, A/P rice fields CAR, EUR, NA,
Urb. var. flavescens 1980 SA
Eleocharis flavescens var. olivacea Walters, 1980 A/P rice fields EUR, NA
(Torr.) Gleason
Eleocharis geniculata (L.) Roem. Cardenas et al., 1972; A aquatic biotypes, AFR, ASI, AUS,
&Schult. Kern, 1974; Holm et al., crops, grasslands, EUR, IND, NA,
1979; Kühn, 1982; Soerjani moist areas, rice PI, SA
et al., 1987; Moody, 1989; fields, taro paddies
Wagner et al., 1990;
Kissman, 1997; Waterhouse,
1997; Kukkonen, 2001;
Simpson & Inglis, 2001;
Ravi & Mohanan, 2002
Eleocharis interstincta (Vahl) Roem. Holm et al., 1979 P NA, SA
&Schult.
Eleocharis kuroguwai Ohwi Holm et al., 1979 P ASI
Eleocharis macbarronii K. L. Wilson Wilson, 1993 P rice fields AUS
Eleocharis macrostachya Britton P NA
Eleocharis mamillata H. Lindb. Holm et al., 1979 P ASI
Eleocharis montana (Kunth) Roem. Holm et al., 1979 P disturbed sites, CAR, NA, SA
&Schult. wet ditches
Eleocharis montevidensis Kunth Carter, 2005 P crops, wet ditches NA, SA
Eleocharis multicaulis Sm. Holm et al., 1979 EUR
Eleocharis mutata (L.) Roem. & Schult. Holm et al., 1979; P rice fields SA
Simpson & Inglis, 2001
Eleocharis obtusa (Willd.) Schult. Holm et al., 1979; Walters, A(P) crops, rice fields, EUR, NA,
1980; Moody,1989; WSSA, wet places PI (Hawaii)
1989; Carter, 2005
Eleocharis ochrostachys Steud. Soerjani et al., 1987; P rice fields ASI
Moody,1989
Eleocharis ovata (Roth) Roem. & Schult. Holm et al., 1979 A(P) crops, wet places ASI, EUR, NA
Eleocharis palustris (L.) Roem. & Schult. Holm et al., 1979,1997; P aquatic biotypes, AFR, ASI, CAR,
Kühn, 1982; Moody,1989; rice fields EUR, IND, NA,
WSSA, 1989; Simpson & SA
Inglis, 2001
Eleocharis parodii Barros Wilson, 1993 P rice fields AUS
Eleocharis parvula (Roem. & Schult.) Holm et al., 1979; WSSA, P AFR, ASI, EUR,
Link ex Bluff, Nees & Schauer 1989; Carter, 2005 NA, SA
Eleocharis pellucida J. Presl & C. Presl Reed, 1977; Kühn, 1982; A/P aquatic biotypes, ASI, IND, PI
Zhirong et al., 1990 crops
Eleocharis philippinensis Svenson Kern, 1974; Soerjani et al., P rice fields ASI, AUS, PI
1987; Moody, 1989; Simpson
&Koyama, 1998; Simpson
&Inglis, 2001
Species1Source Habit 2Habitat Distribution3

Eleocharis plantaginoidea W. F. Wight Holm et al., 1979 IND
Eleocharis quadrangulata (Michx.) Elliott, 1821; Holm et al., P pond shores, NA
Roem. & Schult. 1979; WSSA, 1989; rice fields
Carter, 2005
Eleocharis quinqueflora (Hartmann) Moody, 1989 P rice fields IND
O. Schwarz
Eleocharis radicans (Poir.) Kunth Wagner et al., 1990 P taro paddies CAR, NA, PI
(Hawaii), SA
Eleocharis retroflexa (Poir.) Urb. Kern, 1974; Holm et al., A rice fields ASI, AUS, IND,
1979; Soerjani et al.,1987; SA
Moody, 1989; Simpson &
Koyama, 1998; Simpson
&Inglis, 2001; Ravi &
Mohanan, 2002
Eleocharis rostellata Torr. WSSA, 1989 P wet places CAR, NA
Eleocharis sellowiana Kunth Kissman, 1997 SA
Eleocharis sphacelata R. Br. Holm et al., 1979 P AUS
Eleocharis spiralis R. Br. Moody, 1989 P rice fields ASI, IND
Eleocharis subtilis Boeckeler Holm et al., 1979 CAR
Eleocharis tetraquetra Nees Holm et al., 1979; Moody, P rice fields ASI, IND
1989; Zhirong et al., 1990
Eleocharis tuberosa Schult. Holm et al., 1979 ASI
Eleocharis valleculosa Zhirong et al., 1990 P rice fields, ASI
Ohwi f. setosa (Ohwi) Kitag. wetlands
Eleocharis variegata (Poir.) C. Presl Holm et al., 1979; ASI
Moody,1989
Eleocharis vivipara Link WSSA, 1989 P wet places NA
Eleocharis wichurae Boeckeler Reed, 1977 P? wet places ASI
Eleocharis wolfii A. Gray Holm et al.,1979; P rice fields, ASI, NA
Moody,1989 wet places
Eleocharis yokoscensis Zhirong et al., 1990 Price fields, ASI
(Franch. & Sav.) Ts. Tang & F. T. Wang wetlands
Fimbristylis acuminata Vahl Kern, 1974; Holm et al., P aquatic biotypes, ASI, AUS, IND,
1979; Kühn, 1982; Soerjani crops, rice fields PI
et al., 1987; Moody,1989;
Simpson & Inglis, 2001;
Ravi & Mohanan, 2002
Fimbristylis aestivalis (Retz.) Vahl Lin, 1968; Kern, 1974; Reed, A aquatic biotypes, ASI, AUS, IND,
1977; Holm et al., 1979; Kühn, crops, rice fields, PI, SA
1982; Koyama, 1985; Soerjani taropaddies
et al., 1987; Moody,1989;
Zhirong et al., 1990; Wagner
et al.,1990; Simpson & Koyama,
1998; Simpson & Inglis, 2001;
Ravi & Mohanan, 2002
Fimbristylis albicans Nees Moody,1989 rice fields IND
92 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Species1Source Habit 2Habitat Distribution3
Appendix 2. Continued.

The Significance of Cyperaceae as Weeds 93
Fimbristylis alboviridis C. B. Clarke Soerjani et al., 1987; rice fields ASI, IND
Moody, 1989
Fimbristylis anisoclada Ohwi Kern, 1974; Moody, 1989 P rice fields ASI, IND, PI
Fimbristylis annua (All.) Roem. & Schult. Kral, 1971; Cardenas et al., A rice fields ASI, EUR, NA,
1972; Godfrey & Wooten, PI, SA
1979; Holm et al., 1979;
Moody, 1989; WSSA, 1989;
DeFelice & Bryson, 2004;
Carter, 2005
Fimbristylis argentea (Rottb.) Vahl Moody, 1989; Ravi & P rice fields ASI, IND
Mohanan, 2002
Fimbristylis autumnalis (L.) Holm et al., 1979; Lorenzi, A aquatic biotypes, AFR, ASI, IND,
Roem. & Schult. 1982; Kühn, 1982; WSSA, crops, rice fields NA, SA
1989; Kissman, 1997;
Simpson & Inglis, 2001
Fimbristylis bisumbellata (Forssk.) Kern, 1974; Holm et al., A cultivated fields, AFR, AUS, EUR,
Bubani 1979; DeFilipps, 1980b; rice fields IND, PI
Koyama, 1985; Soerjani
et al., 1987; Moody, 1989;
Simpson & Koyama,
1998; Kukkonen, 2001
Fimbristylis caesia Miq. Kern, 1974; Moody, 1989 A? edges of rice fields, PI
roadsides
Fimbristylis caroliniana (Lam.) Fernald Kral, 1971 P disturbed soil, CAR, NA
waste lands
Fimbristylis castanea (Michx.) Vahl Kral, 1971 P disturbed soil, CAR, NA
waste lands
Fimbristylis cinnamometorum
(Vahl) Kunth Moody, 1989 A/P rice fields IND
Fimbristylis complanata (Retz.) Link Kern, 1974; Holm et al., P aquatic biotypes, ASI, IND, PI
1979; Moody, 1989; Simpson crops, rice fields
&Koyama, 1998; Simpson
&Inglis, 2001
Fimbristylis cymosa R. Br. Holm et al., 1979; Moody, P rice fields, sweet ASI, PI
1989; Waterhouse, 1997 potato, taro
Fimbristylis decipiens Kral Kral, 1971; Godfrey & A disturbed soil, NA
Wooten, 1979 waste lands
Fimbristylis dichotoma (L.) Vahl Lin, 1968; Kral, 1971; Kern, A/P aquatic biotypes, AFR, ASI, AUS,
1974; Holm et al.,1977,
crops, grasslands, EUR, IND, NA,
1979;
Godfrey & Wooten, lawns, rice fields, PI, SA
1979; Kühn, 1982; Lorenzi, sugarcane and tea
1982; Koyama, 1985; plantations, waste
Soerjani et al., 1987; Moody,places, wetlands
1989; WSSA, 1989; Zhirong
et al., 1990; Gordon-Gray,
1995; Kissman, 1997;
Waterhouse, 1997; Simpson
&Koyama, 1998; Kukkonen,
2001; Simpson & Inglis,
2001; Ravi & Mohanan,
2002; Carter, 2005
Fimbristylis dipsacea (Rottb.) C. B. Clarke Kern, 1974; Moody, 1989; A rice fields AFR, ASI, PI
Ravi & Mohanan, 2002
Fimbristylis dura (Zoll. & Moritz) Merr. Kern, 1974; Moody, 1989 P rice fields ASI, IND, PI
Species1Source Habit 2Habitat Distribution3

Fimbristylis eragrostis (Nees) Hance Moody, 1989 P rice fields ASI
Fimbristylis falcata (Vahl) Kunth Moody, 1989 P rice fields IND
Fimbristylis ferruginea (L.) Vahl Reed,1977; Holm et al.,1979; A/P aquatic biotypes, ASI, AUS, EUR,
Kühn, 1982; Akobundu & rice fields, IND, PI, SA
Agyakwa, 1987; Moody, waste places
1989; Johnson, 1997;
Simpson & Inglis, 2001
Fimbristylis globulosa (Retz.) Kunth Kern, 1974; Holm et al., P aquatic biotypes, ASI, IND, PI
1979; Kühn, 1982; Soerjani crops, rice fields
et al., 1987; Moody, 1989;
Simpson & Koyama, 1998;
Simpson & Inglis, 2001
Fimbristylis griffithii Boeckeler Soerjani et al., 1987; A rice fields ASI
Moody, 1989
Fimbristylis hispidula (Vahl) Kunth Kühn, 1982 A/P aquatic biotypes, ASI, NA, PI, SA
crops, grasslands
Fimbristylis koidzumiana Ohwi Lin, 1968; Reed, 1977; A crops, wet places ASI
Holm et al., 1979
Fimbristylis littoralis Gaudich. Kern, 1974; Moody, 1981, A/P rice fields pantropical
1989; Akobundu &
Agyakwa, 1987; Johnson,
1997
Fimbristylis merrillii J. Kern Kern, 1974; Moody, 1989 A rice fields ASI, AUS, PI
Fimbristylis miliacea (L.) Vahl Ohwi, 1965; Lin, 1968; A/B aquatic biotypes, AFR, ASI, AUS,
Kral, 1971; Kern, 1974; crops, rice fields, CAR, IND, NA,
Reed, 1977; Holm et al., wet places PI, SA
1977, 1979; Kühn, 1982;
Lorenzi, 1982; Koyama,
1985; Soerjani et al., 1987;
Moody,1989; WSSA,1989;
Zhirong et al., 1990;
Kissman,1997; Waterhouse,
1997; Simpson & Koyama,
1998; Kukkonen, 2001;
Simpson & Inglis, 2001;
Ravi & Mohanan, 2002;
DeFelice & Bryson, 2004;
Carter, 2005
Fimbristylis nutans (Retz.) Vahl Moody, 1989; Simpson & P rice fields ASI, AUS, IND
Koyama, 1998; Simpson
&Inglis, 2001
Fimbristylis pauciflora R. Br. Moody, 1989; Wilson,1993; P/A? rice fields ASI, AUS, IND,
Simpson & Inglis, 2001 PI
Fimbristylis polytrichoides (Retz.) R. Br. Moody, 1989 P rice fields AFR, AUS, IND
Fimbristylis quinquangularis (Vahl) Holm et al.,1979; Koyama, A/B rice fields AFR, ASI, IND
Kunth 1985; Moody,1989;
Kukkonen, 2001
Fimbristylis schoenoides (Retz.) Vahl Kern, 1974; Holm et al., A/P rice fields ASI, AUS, IND,
1979; Koyama, 1985; NA
Soerjani et al., 1987;
Moody, 1989; Simpson
&Koyama, 1998;
Simpson & Inglis,
2001; Carter,2005
94 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Species1Source Habit 2Habitat Distribution3
Appendix 2. Continued.

The Significance of Cyperaceae as Weeds 95
Fimbristylis sericea R. Br. Moody, 1989 rice fields ASI, AUS, IND
Fimbristylis squarrosa Vahl Holm et al., 1979; Moody, A rice fields AFR, ASI, EUR,
1989; Kukkonen, 2001 IND, SA
Fimbristylis stauntoni Debeaux & Franch. Zhirong et al., 1990 A rice fields, ASI
wetlands
Fimbristylis stolonifera C. B. Clarke Moody, 1989 rice fields ASI
Fimbristylis subbispicata Nees & Meyen Holm et al., 1979; rice fields ASI, IND
Moody, 1989
Fimbristylis tenera Roem. & Schult. Holm et al., 1979; A/P rice fields IND
Moody, 1989
Fimbristylis tetragona R. Br. Kern, 1974; Holm et al., P rice fields ASI, AUS, IND,
1979; Koyama, 1985; PI
Moody,1989
Fimbristylis thonningiana Boeckeler Holm et al., 1979 AFR
Fimbristylis tomentosa Vahl Kern, 1974; Soerjani et al., A rice fields AFR, ASI, AUS,
1987; Moody, 1989; NA, PI
Carter,2005
Fimbristylis tristachya R. Br. Holm et al., 1979; rice fields AFR, ASI
Moody, 1989
Fimbristylis turkestanica (Regel) Kukkonen, 2001 P fields, gardens AFR, ASI, EUR,
B. Fedtsch. IND
Fimbristylis verrucifera (Maxim.) Makino Reed, 1977 A wet places ASI
Fuirena breviseta (Coville) Coville WSSA, 1989 P NA
Fuirena ciliaris (L.) Roxb. Ohwi, 1965; Kern, 1974; A aquatic biotypes, AFR, ASI, AUS,
Holm et al., 1979; Kühn, crops, grasslands, IND, PI
1982; Akobundu & Agyakwa, rice fields
1987; Soerjani et al., 1987;
Moody, 1989; Simpson &
Koyama, 1998; Simpson
&Inglis, 2001; Ravi &
Mohanan, 2002
Fuirena pumila (Torr.) Spreng. WSSA, 1989 A NA
Fuirena scirpoidea Michx. WSSA, 1989 P NA
Fuirena simplex Vahl WSSA, 1989 A/P NA
Fuirena squarrosa Michx. WSSA, 1989 P NA
Fuirena stricta Steud. subsp. chlorocarpa Johnson, 1997; P rice fields AFR
(Ridl.) Lye Simpson & Inglis, 2001
Fuirena umbellata Rottb. Kern, 1974; Holm et al., P aquatic biotypes, AFR, ASI, AUS,
1979; Kühn, 1982; ditches, grasslands, IND, PI, SA
Akobundu & Agyakwa, rice fields
1987; Soerjani et al.,
1987; Moody, 1989;
Johnson, 1997; Ravi &
Mohanan, 2002
Isolepis carinata Hook. & Arn. ex Torr. Godfrey & Wooten, 1979; A crops, grasslands, NA
WSSA, 1989 waste places
Kyllinga aurata Nees Holm et al., 1979 P AFR
Species1Source Habit 2Habitat Distribution3

Kyllinga brevifolia Rottb. Lin, 1968; Cardenas et al., P crops, disturbed AFR, ASI, AUS,
1972; Kern, 1974; Godfrey sites, fallow fields, CAR, EUR, IND,
&Wooten, 1979; Kühn, gardens, grasslands, NA, PI, SA
1982; Lorenzi, 1982; pastures, rice fields,
Koyama, 1985; Soerjani et al., turf, waste places
1987; WSSA, 1989; Zhirong
et al., 1990; Wagner et al.,
1990; Wilson,1993; Gordon-
Gray, 1995; Holm et al.,
1997; Kissman, 1997;
Waterhouse, 1997; Bryson
et al., 1998; Kukkonen, 2001;
Simpson & Inglis, 2001;
Ravi & Mohanan, 2002;
Carter, 2005
Kyllinga bulbosa P.Beauv. Akobundu & Agyakwa, P rice fields AFR
1987
Kyllinga colorata (L.) Druce Zhirong et al., 1990 P ASI
Kyllinga erecta Schumach. Terry, 1976; Holm et al., P crops, cultivated AFR, ASI, IND,
1979; Kühn, 1982; lands, grasslands, PI
Akobundu & Agyakwa, rice fields, waste
1987; Gordon-Gray, 1995; places
Johnson, 1997; Simpson
&Inglis, 2001
Kyllinga gracillima Miq. WSSA, 1989; Bryson et al., P crops, grasslands, ASI, NA
1998; Simpson & Inglis, 2001 turf, waste places
Kyllinga melanosperma Nees Moody, 1989 P rice fields IND, PI
Kyllinga nemoralis (J. R. Forst. & Holm et al., 1979; Kühn, Pcrops, gardens, AFR, ASI, AUS,
G. Forst.) Dandy ex Hutch. & Dalziel 1982; Soerjani et al., 1987; grasslands, lawns, IND, PI, SA
Moody, 1989; WSSA,1989; pastures,
Wagner et al., 1990; plantations,
Waterhouse, 1997; Kukkonen, rice fields,
2001; Simpson & Inglis, roadsides, turf,
2001; Ravi & Mohanan, 2002 waste places
Kyllinga odorata Vahl Terry, 1976; Godfrey & P damp sandy AFR, ASI, AUS,
Wooten, 1979; Holm et al., ground, disturbed NA, PI, SA
1979; Lorenzi, 1982; Moody, grassland, gardens,
1989; Wilson, 1993; pastures, lawns,
Gordon-Gray, 1995; Kissman, rice fields, turf,
1997; Bryson et al., 1998; waste places
Simpson & Inglis, 2001;
Carter, 2005
Kyllinga polyphylla Willd. ex Kunth Holm et al., 1979; Moody, P agricultural land, AFR, ASI, PI
1989; Waterhouse, 1997; crops, pastures,
Simpson & Inglis, 2001 rice fields, roadsides,
turf, waste places
Kyllinga pumila Michx. Holm et al.,1979; Akobundu A crops, pastures, AFR, CAR, NA,
&Agyakwa, 1987; WSSA, turf, rice fields, SA
1989; Le Bourgeois & Merlier, waste places
1995; Johnson, 1997; Bryson
et al., 1998; Simpson & Inglis,
2001
96 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Species1Source Habit 2Habitat Distribution3
Appendix 2. Continued.

The Significance of Cyperaceae as Weeds 97
Kyllinga squamulata Thonn. ex Vahl Terry, 1976; Akobundu & A cultivated lands, AFR, ASI, CAR,
Agyakwa,1987;WSSA,1989; turf, waste places IND, NA
Le Bourgeois & Merlier,
1995; Bryson et al., 1998;
Simpson & Inglis, 2001;
Carter, 2005
Kyllinga triceps Rottb. Kern, 1974; Moody, 1989; P sandy lawn AFR, ASI, AUS,
Le Bourgeois & Merlier, IND, PI
1995; Simpson & Inglis, 2001
Lepidosperma chinense Nees & Meyen Kern, 1974 P rice fields ASI, PI
Lepironia articulata (Retz.) Domin Moody, 1989 P rice fields ASI
Lipocarpha chinensis (Osbeck) J. Kern Lin, 1968; Kern,1974; Reed, A/P aquatic biotypes, AFR, ASI, AUS,
1977; Holm et al., 1979; crops, grasslands, IND
Kühn, 1982; Koyama, 1985; rice fields, waste
Soerjani et al., 1987; Johnson, wet places
1997; Simpson & Inglis, 2001
Lipocarpha maculata (Michx.) Torr. Carter, pers. obs. A disturbed wet sites, NA, SA
ditches roadsides
Lipocarpha microcephala (R. Br.) Kunth Kern, 1974; Holm et al., A rice fields, ASI, AUS, IND
1979 sugarcane fields
Lipocarpha squarrosa (L.) Goetgh. Kern, 1974; Moody 1989; A/P? crops, cultivated AFR, ASI, AUS,
Kukkonen, 2001; Simpson land, rice fields, IND
&Inglis, 2001; Ravi & wet fields
Mohanan, 2002
Mapania cuspidata (Miq.) Uittien Moody,1989 P rice fields ASI
Oxycaryum cubense (Poepp. & Kunth) Holm et al., 1979; P aquatic, AFR, CAR, NA,
Palla Simpson & Inglis, 2001 floating mats SA
Rhynchospora aurea Vahl Kissman, 1997 P SA
Rhynchospora caduca Elliott Wagner et al., 1990; Ppastures NA, PI (Hawaii)
Wagner & Herbst, 1995
Rhynchospora cephalotes (L.) Vahl Kissman, 1997 P SA
Rhynchospora colorata (L.) H. Pfeiff. WSSA, 1989 P wet places NA
Rhynchospora corniculata (Lam.) WSSA, 1989; DeFelice Pditches, wet places NA
A. Gray &Bryson, 2004
Rhynchospora corymbosa (L.) Britton Kern, 1974; Holm et al., Paquatic biotypes, AFR, ASI, AUS,
1979; Kühn, 1982; Lorenzi, crops, rice fields, IND, PI, SA
1982; Koyama, 1985; waste places,
Akobundu & Agyakwa, 1987; wet places
Soerjani et al.,1987; Moody,
1989; Johnson, 1997;
Simpson & Inglis, 2001;
Ravi & Mohanan, 2002
Rhynchospora fascicularis (Michx.) Vahl Godfrey & Wooten, 1979 P pastures, roadsides CAR, NA, SA
Rhynchospora globularis (Chapm.) Godfrey & Wooten, 1979; Pdisturbed areas, CAR, NA,
Small WSSA, 1989 roadsides PI (Hawaii), SA
Rhynchospora glomerata (L.) Vahl Godfrey & Wooten, 1979 Ppastures, roadsides NA
Species1Source Habit 2Habitat Distribution3

Rhynchospora holoschoenoides (Rich.) Simpson & Inglis, 2001 P rice fields AFR, SA
Herter
Rhynchospora inexpansa (Michx.) Vahl Godfrey & Wooten, 1979 P pastures, roadsides NA
Rhynchospora latifolia (Baldwin) WSSA, 1989 P wet places NA
W. W. Thomas
Rhynchospora longisetis R. Br. Moody, 1989 A rice fields ASI
Rhynchospora microcarpa Baldwin
Godfrey & Wooten, 1979 P pastures, roadsides CAR, NA
ex A.Gray
Rhynchospora nervosa (Vahl) Cardenas et al., 1972; P crops, grasslands, CAR, NA, SA
Boeckeler Reed, 1977; Kühn, 1982; low elevations,
Lorenzi, 1982; Kissman, wet places
1997; Simpson & Inglis,
2001
Rhynchospora radicans H. Pfeiff. Strong & Wagner, 1997 A/P cultivated lands CAR, PI (Hawaii),
subsp. microcephala (Berteroex Spreng.) SA
W. W. Thomas
Rhynchospora rubra (Lour.) Makino Holm et al., 1979; Moody,A/P rice fields ASI
1989
Rhynchospora submarginata Kük. Kern, 1974; Moody, 1989 A rice fields ASI, AUS, IND,
PI
Rhynchospora tenuis Link Holm et al., 1979 SA
Rhynchospora wightiana (Nees) Steud. Kern, 1974; Moody, 1989 A fallow rice fields, ASI, IND, PI
rice fields
Schoenoplectus acutus (Muhl. ex J. M. USDA, 1970; Holm et al., P aquatic biotypes ASI, NA
Bigelow) Á. Löve & D. Löve 1979; Kühn, 1982; Moody,
1989; WSSA, 1989;
DeFelice & Bryson, 2004
Schoenoplectus americanus (Pers.) Holm et al., 1979; WSSA, P NA
Volkart ex Schinz & R. Keller 1989
Schoenoplectus articulatus (L.) Palla Kern, 1974; Holm et al., A/P rice fields, swampy AFR, ASI, AUS
1979; Soerjani et al., 1987; fallow fields
Moody,1989; Simpson &
Koyama, 1998; Simpson
&Inglis, 2001; Ravi &
Mohanan, 2002
Schoenoplectus californicus Holm et al., 1979; WSSA, Pwet places NA, PI, SA
(C. A. Mey.) Soják 1989; Kissman, 1997
Schoenoplectus corymbosus Moody, 1989 P rice fields IND
(Roth ex Roem. & Schult.) J. Raynal
Schoenoplectus erectus (Poir.) Holm et al., 1979; Kühn, A aquatic biotypes, AFR, ASI, AUS,
Palla ex J. Raynal 1982 wet places EUR, IND, NA,
PI, SA
Schoenoplectus grossus (L. f.) Palla Kern, 1974; Holm et al., P ditches, rice fields ASI, AUS, PI
1979, 1997; Soerjani et al.,
1987; Moody, 1989
98 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Species1Source Habit 2Habitat Distribution3
Appendix 2. Continued.

The Significance of Cyperaceae as Weeds 99
Schoenoplectus juncoides (Roxb.) Palla Lin, 1968; Kern, 1974; A aquatic biotypes, AFR, ASI, EUR,
Holm et al., 1979; DeFilipps, crops, rice fields IND, PI
1980a; Koyama, 1985;
Soerjani et al., 1987; Moody,
1989; Zhirong et al., 1990;
Simpson & Koyama, 1998;
Kukkonen, 2001; Simpson
&Inglis, 2001
Schoenoplectus lacustris (L.) Palla Holm et al., 1979; Moody, P aquatic biotypes AFR, ASI, EUR,
1989; Simpson & Inglis, 2001 PI
Schoenoplectus lacustris (L.) Palla ×Kukkonen, 2001 P rice fields ASI, IND
S. triqueter (L.) Palla
Schoenoplectus lateriflorus (J. F. Gmel.) Kern, 1974; Soerjani et al., A rice fields ASI, AUS, IND
Lye 1987; Moody, 1989;
Kukkonen, 2001
Schoenoplectus litoralis (Schrad.) Palla Holm et al., 1979; Kühn, Paquatic biotypes AFR, ASI, AUS,
1982; Simpson & Inglis, 2001 EUR, IND, PI
Schoenoplectus mucronatus (L.) Palla Kern, 1974; Reed, 1977; P aquatic biotypes, AFR, ASI, AUS,
Holm et al., 1979, 1997; ditches, rice EUR, IND, NA,
Kühn, 1982; Soerjani et al., fields, wet places PI
1987; Moody, 1989; WSSA,
1989; Simpson & Koyama,
1998; Kukkonen, 2001;
Simpson & Inglis, 2001;
DeFelice & Bryson, 2004
Schoenoplectus pungens (Vahl) Palla Moore & Edgar, 1970; P aquatic, AUS, CAR, EUR,
Simpson & Inglis, 2001 unspecified NA, PI, SA
Schoenoplectus roylei (Nees) Moody,1989; Kukkonen, Aditches, rice AFR, IND
Ovcz. & Czukav.2001; Simpson & Inglis, 2001 fields
Schoenoplectus senegalensis Akobundu & Agyakwa, A rice fields AFR
(Hochst. ex Steud.) Palla ex J. Raynal 1987; Johnson, 1997
Schoenoplectus supinus (L.) Palla Moody, 1989; Simpson & A rice fields ASI, AUS, IND
Koyama, 1998; Kukkonen,
2001; Simpson & Inglis,
2001; Ravi & Mohanan, 2002
Schoenoplectus tabernaemontani Reed, 1977; WSSA, 1989; P aquatic, brackish AFR, ASI, AUS,
(C. C. Gmel.) Palla Zhirong et al., 1990 water EUR, IND, NA
Schoenoplectus triqueter (L.) Palla Reed, 1977; Holm et al., P aquatic biotypes, AFR, ASI, EUR,
1979; Kühn, 1982; Moody, rice fields IND, PI
1989; Zhirong et al., 1990;
Kukkonen, 2001; Simpson
&Inglis, 2001
Schoenoplectus wallichii (Nees) T.Koyama Lin, 1968; Kern, 1974; P rice fields, ASI, IND
Reed, 1977; Holm et al., wet places
1979; Moody, 1989
Scirpodendron ghaeri (Gaertn.) Merr. Moody, 1989 P rice fields ASI
Scirpus atrovirens Willd. Holm et al., 1979; P roadsides, NA
WSSA, 1989 wet places
Scirpus cyperinus (L.) Kunth Holm et al., 1979; WSSA, P roadsides, NA
1989; Carter, 2005 wet places
Species1Source Habit 2Habitat Distribution3

Scirpus giganteus Kunth Kissman, 1997 P SA
Scirpus holoschoenus L. Reed, 1977; Holm et al., P crops, waste AFR, ASI, EUR,
1979; Kühn, 1982 places IND
Scirpus michelianus L. Moody, 1989; Zhirong A farmland, field ASI, IND
et al., 1990 borders, rice fields
Scirpus pendulus Muhl. Holm et al., 1979 P roadsides, AUS, NA
wet places
Scirpus sylvaticus L. Holm et al., 1979 P EUR
Scirpus triangulatus Roxb. Holm et al., 1979 P AUS
Scleria bancana Miq. Holm et al.,1979; Moody, rice fields ASI
1989
Scleria biflora Roxb. Kern, 1974; Koyama, A rice fields, ASI, IND, PI
1985; Moody, 1989 roadsides,
tea plantations
Scleria boivinii Steud. Holm et al., 1979 AFR
Scleria bracteata Cav. Holm et al., 1979; Kissman, SA
1997
Scleria canescens Boeckeler Holm et al., 1979 NA, SA
Scleria caricina (R. Br.) Benth. Simpson & Koyama, rice fields PI
1998; Simpson & Inglis, 2001
Scleria depressa (C. B. Clarke) Nelmes Johnson, 1997 Price fields AFR
Scleria lacustris C. Wright Tobe et al.,1998;Wunderlin, A aquatic waste AFR, CAR, NA,
1998; Jacono, 2001 places, wet places SA
Scleria laevis Retz. Kern, 1974; Holm et al., P fallow rice fields ASI, AUS, IND,
1979; Moody, 1989; Ravi PI
&Mohanan, 2002
Scleria lithosperma (L.) Sw. Holm et al., 1979; Kühn, P aquatic biotypes, AFR, ASI, IND,
1982; Moody,1989; crops, rice fields, PI, SA
Simpson & Inglis, 2001 waste places,
wet places
Scleria melaleuca Rchb. ex Schltdl. Cardenas et al., 1972; P CAR, SA
&Cham. Holm et al., 1979; Lorenzi,
1982; Moody,1989
Scleria myriocarpa Steud. Holm et al., 1979 SA
Scleria naumanniana Boeckeler Akobundu & Agyakwa, P forest clearings, AFR
1987 wet areas
Scleria novae-hollandiae Boeckeler Kern, 1974; Moody, 1989 A fallow rice fields, AUS, IND, PI
rice fields
Scleria oblata S. T. Blake Holm et al., 1979; Moody, P rice fields ASI
1989
Scleria poaeformis Retz. Holm et al., 1979; Moody,P rice fields ASI, AUS, IND
1989
Scleria polycarpa Boeckeler Holm et al., 1979 PPI
Scleria purpurascens Steud. Holm et al., 1979; Moody,Price fields ASI
1989
100 Sedges: Uses, Diversity, and Systematics of the Cyperaceae
Species1Source Habit 2Habitat Distribution3
Appendix 2. Continued.

The Significance of Cyperaceae as Weeds 101
Scleria rugosa R. Br. Kern, 1974; Koyama,1985; A rice fields ASI, AUS, IND,
Moody, 1989 PI
Scleria scindens Nees Reed, 1977 P CAR
Scleria scrobiculata Nees & Meyen Holm et al., 1979; Moody, P rice fields PI
1989
Scleria sumatrensis Retz. Holm et al., 1979; Kühn, A aquatic biotypes, ASI, AUS, IND,
1982; Moody,1989; crops, forests, PI
Simpson & Inglis, 2001 rice fields
Scleria tessellata Willd. var. sphaerocarpa Kühn, 1982; Moody,1989; A aquatic biotypes, AFR, ASI, AUS,
E. A. Rob. Le Bourgeois & Merlier, grasslands, rice IND, PI, SA
1995; Simpson & Inglis, 2001 fields, wet places
Scleria verrucosa Willd. Akobundu & Agyakwa, P wet areas AFR
1987
1Plant nomenclature follows Flora of North America,volume 23; plant names were also verified through the Missouri Botanical
Garden w3TROPICOS VAST database (rev. 1.5) (http://mobot.mobot.org/W3T/Search/vast.html) and the International Plant Names
Index (http://www.ipni.org/index.html). A more inclusive list of names cited in the references is available from the authors.
2A = annual; B = biennial; P = perennial; supplemental data from Kükenthal (1935–1936), Kern (1974), Holm et al. (1977, 1997),
Haines and Lye (1983), Koyama (1985), Soerjani et al. (1987), Wilson (1993), Gordon-Gray (1995), Lye (1995), Simpson and Inglis
(2001), Kukkonen (2001), and Flora of North America,volume 23.
3AFR = Africa including Madagascar; ASI = Asia; AUS = Australia; CAR = Caribbean Islands; EUR = Europe; IND = Indian subcontinent
including Sri Lanka; NA = North America; PI = Pacific Islands; SA = South America.
Species1Source Habit 2Habitat Distribution3