JOURNAL
OF THE
ARNOLD ARBORETUM
VOLUME 68 OCTOBER 1987 NUMBER 4
THE GENERA OF CYPERACEAE IN THE
SOUTHEASTERN UNITED STATES'
GORDON C. TUCKER)
CYPERACEAE A. L. de Jussieu, Gen. Pl. 26. 1789, nom. cons.
(SEDGE FAMILY)
Small to large perennial or annual herbs of aquatic or terrestrial habitats.
Roots fibrous; many species rhizomatous or stoloniferous. Plants glabrous or
'Prepared for the CGeneric Flora of the Southeastern United States, a long-term project made possible
by grants from the National Science Foundation and at this writing supported by BSR-8415367
(Norton G. Miller, principal investigator), under which this account was prepared, and BSR-8415769
(Carroll E. Wood, Jr., principal investigator). This treatment, 118th in the series, follows the format
established in the first paper (Jour. Arnold Arh. 39: 296-346. 1958) and continued to the present.
The area covered by the Generic Flora includes North and South Carolina, Georgia, Florida. Ten-
nessee, Alabama, Mississippi, Arkansas, and Louisiana. The descriptions are based primarily on the
plants of this area. with information about extraregional members of a family or genus in brackets.
Those references I did not verify are marked with asterisks.
Preceding the reference for each genus, a paragraph is given listing by author all familial or tribal
references pertinent to that genus.
I have enjoyed working with Norton Miller and Carroll Wood on the Generic Flora Project, and
I thank them for their interest and assistance. Thomas .1. Rosatti has given helpful advice on many
occasions. Stephen A. Spongberg and Elizabeth B. Schmidt improved the final manuscript with their
editorial expertise. Melinda F. Denton helped initiate my interest in the Cyperaceae. and Robert L.
Wilbur ably supervised my graduate studies on C('perus. Thanks are extended to the staffs of the
New York State Library (especially Alta Beach. Senior Librarian), the Missouri Botanical Garden
Library,. the New York Botanical Garden Library, and the Libraries of the Arnold Arboretum and
the Gray Herbarium for providing access to the needed literature. I am grateful to the curators at A.
AC. ALU, CCNL, CONN, )DUKE. i, F, FSUi, GA. iH, KIRi. MASS. MUHW, NASC, NCBS, NCSC, NCU. NEBC, NY,
NYS. PENN. PH. SD, SMiU, UC. UNA, 'US, USCH. WRSL. WVA. and vu who have sent specimens or provided
access to collections and hospitality during my visits. Charles T. Bryson. Patricia L. Forbes. Paul
Goeltghebeur, Encarnacion R. Guaglianone, Robert Kral. Anton A. Reinicek, Alfred E. Schuyler.
Lisa A. Standley. Wm. Wayt Thomas. Marcia 1. Waterway, and Karen L. Wilson have shared
information and discussed nomenclatural matters. Thomas J. Rosatti. William R. Linke. Jr.. Bruce
A. Sorrie. Jason R. Tucker, and Joshua D. Tucker have helped with field work. Vicky Martin Tucker.
my wife. has been supportive (and tolerant) of my weekend collecting trips, week-long research trips.
and numerous evenings given over to sedges: her understanding is deeply appreciated.
The illustrations were prepared by Karen Stoutsenberger under the supervision of Carroll Wood
or Kenneth R. Robertson, from plants collected by Carroll Wood or from specimens in the herbaria
of Harvard I niversity (A, oi).
Contribution number 543 of the Nes York State Science Service.
'Biological Survey, New York State Museum, The State Education Department, Albany, New York
12230.,
,c President and Fellows of Harvard College, 1987.
Journal of the Arnold .rboretum 68: 361-445. October. 1987.


362 JOURNAL OF THE ARNOLD ARBORETUM [voi. 68
scabrellate. Culms single, approximate, or caespitose, trigonous, triquetrous,
or terete, the cortex chlorenchymatous, the central region aerenchymatous or
hollow: cortical bundles with sheaths like those in the leaves. Leaves basal or
both basal and cauline: sheaths closed: blades linear to lanceolate, flat. con-
duplicate, plicate, or involute: stomata paracytic, sometimes surrounded by 1-
4 porrect or arching cuticular papillae: anatomy non-kranz or kranz, if kranz,
the bundle sheaths 2-layered ("C'erius type") or 3-layered ("Fimhristilis type").
Inflorescences spicate or umbelliform [corymbose], sessile, simple, or with
second- and third- [to fifth-]order branching. Spikelets 1- to many-flowered.
basally subtended by a scalelike prophyll. above which may be 1 or more sterile
scales: flowers perfect or imperfect and monoecious (rarely dioecious), each
borne in the axil of a scale ("glume" of some authors), anemophilous (infre-
quently entomophilous): perianth absent or comprising 1 or 2 series of smooth
or barbed bristles, at maturity shorter to several times longer than mature
achene. Stamens (1, 2, or) 3: filaments ribbonlike or capillary: anthers broadly
ellipsoid to linear, basifixed; pollen maturing as cryptotetrads (pseudomonads),
subspheroidal, trinucleate (binucleate?) when shed. Gynoecium tricarpellate
and stigmas 3, or bicarpellate (dorsiventrally or laterally compressed) and stig-
mas 2: styles and stigmas capillary, glabrous or glandular-pubescent: ovules
basal, anatropous, bitegmic, crassinucellar: megagametophyte (embryo sac) of
the Polygonum type. Achene trigonous or lenticular, ovoid, obovoid, or ellip-
soid, smooth. puncticulate, or papillose: endosperm mealy, with starch grains,
protein crystals, and oil droplets, filling most of the achene: embryo small:
embryogeny of the Onagrad (Juncus variation) or Asterad type: germination
epigeal. Base chromosome numbers 5, 6, 7. 8. TYPI GiE Nos: (Cyperus Linnaeus.
A large family ofabout 80 genera and 3500 species, worldwide in distribution.
Seventeen genera occur in our area, including C'arcx L.. with 165 species, the
largest genus of seed plants in the Southeast.
There is general agreement that the Juncaceae are the closest relatives of the
('yperaceae (Thorne: Dahlgren & Rasmussen). Both families have tristichous
phyllotaxy, simultaneous microsporogenesis, post-reductional meiosis, non-
localized (difluse) centromeres, anatropous ovules. and Onagrad embryogeny.
The ('yperaceae are distinguished from the Juncaceae in having conical silica
bodies in the epidermal cells, solitary ovules and basal placentation, pollen-
grain formation in which three of the meiotic products degenerate. nuclear
endosperm, and indehiscent fruits (achenes). North American Cyperaceae lack
a perianth or have one of bristles: North American Juncaceae have expanded
chartaceous tepals. This is useful regionally for distinguishing the two families,
but it cannot he used on a worldwide basis because Orcobolus R. Br. and
several other genera of Southern Hemisphere ('yperaceae also have chartaceous
tepals.
Some authors (e.g.. Fernald, C('ronquist) have treated the Gramineae as the
closest relatives of the ('yperaceae. However, the grasses have apical placen-
tation, orthotropous ovules, distichous phyllotaxy. and open leaf sheaths, and
their affinities are with the Restionaceae and the Flagellariaceae (Thorne: Dahl-
gren & Rasmussen). Also, the grasses are chemically unlike the sedges (Har-


borne, 1971). For example, anthocyanins are common in grasses but unknown
in sedges, while aurones are common in sedges and unknown in grasses (and
in the Juncaceae).
The tribal classification was first elaborated on a worldwide basis by Nees
von Esenbeck and Kunth and has been rather stable since. Some authors
recognized tribes only; some, subfamilies and tribes; and others, subtribes also.
Two subfamilies, both distributed worldwide, are accepted in this treatment:
the Cyperoideae (Scirpoideae Pax, flowers perfect) and the Caricoideae Pax
(flowers imperfect). Included in the Cyperoideae are four tribes, of which the
Scirpeae Dumort. (including Fimbristylideae Raynal; spikelets with I or 2
sterile basal scales, numerous fertile scales spirally arranged, perianth bristles
generally present, embryos well differentiated), the Cypereae (1 or 2 sterile
basal scales, several to many fertile scales distichously arranged, perianth ab-
sent, embryos well differentiated), and the Schoeneae Dumort. (Rhynchospo-
reae Fenzl; spikelets with several sterile basal scales, fertile scales 1 or 2 (to
several), perianth bristles generally present, embryos slightly differentiated) are
represented in our area. No members of tribe Hypolytreae Fenzl (Mapanieae
Koyama) of the tropics grow in North America. Subfamily Caricoideae is
divided into two tribes: the Scleriae Fenzl (achenes naked, borne on a hardened
disk), represented in North America by a single genus, Scleria Berg.; and the
Cariceae Dumort. (achenes enclosed in a perigynium), represented in the South-
east by Cvmophyllus Mack. and Carex (and also in North America by Kobresia
Willd., a circumboreal genus occurring in the northern United States and
Canada, and Uncinia Pers., an austral genus extending north to Jamaica and
Mexico).
Microsporogenesis in sedges differs markedly from that in other angiosperms.
The nucleus of the microsporocyte divides meiotically, but cytokinesis does
not follow immediately. Rather, three nuclei migrate to one end of the pollen
mother cell, where they begin to disintegrate. The fourth nucleus remains in
the center of the cell, where it divides mitotically. One of the resulting daughter
nuclei migrates to the end of the cell, joining the other three disintegrating
products of meiosis. The remaining haploid daughter nucleus divides mitoti-
cally, forming generative and tube nuclei. The generative nucleus divides again
as the exine matures, resulting in the trinucleate pollen grain characteristic of
the family. The four degenerated nuclei often remain visible as dark streaks
near the exine. The wall of the mature pollen grain is thus homologous to the
wall of the pollen mother cell. This pattern of microsporogenesis, presumably
characteristic of the entire family, has been reported in Abildgaardia Vahl,
Bulbostylis Kunth, Carex, Cladium P. Br., Eleocharis R. Br., Fimbristylis Vahl,
Fuirena Rottb., Scirpus L., Scleria, and Rhynchospora Vahl. In the closely
related Juncaceae cytokinesis is delayed in the pollen mother cells until each
daughter nucleus has divided a second time. Thus, the Juncaceae provide a
pattern of microsporogenesis intermediate to that in the Cyperaceae and other
monocots, and emphasizing the relationship of the Cyperaceae and the Jun-
caceae.
Embryology is nearly uniform in the Cyperaceae. Endosperm formation is
nuclear in all genera that have been investigated. Endosperm wall formation
TUCKER, CYPERACEAE
363
1987]


JOURNAL OF THE ARNOLD ARBORETUM
is complete in most genera, but incomplete in Rhynchospora and Scieria. The
mature embryos of the Cyperaceae vary considerably in shape and in the
position of the cotyledon and the radicle. As a rule, each genus has its char-
acteristic type of embryo (Van der Veken). When the achenes mature, the
embryos of tribe Schoeneae are considerably less differentiated than those of
other tribes (Vanhecke).
The sedges are incompletely investigated chemically, although Crperus is
much better known than other genera. Ethereal oils occur in the roots of three
species of C(perus (Hegnauer). Cyanogenesis is evidently uncommon but has
been reported for three species of (Cyperus and for one each of Fimbristylis and
Kyllinga Rottb. (Gibbs). This is surprising because it is widespread in the closely
related Juncaceae. Tannins occur in many sedges, having been reported in
C'perus. Dulichium Pers., Fuirena, and Scirpus (one species each). Alkaloids
are rare; brevicarine, brevicolline, and harman occur in Carex hrevicollis DC.
(Gibbs). Some terpenoids have been reported. Citral, a monoterpenoid, occurs
in species of Kyllinga (Gibbs), and several sesquiterpinoids are known from
species of C(perus (Hegnauer). Quinones are found in both Cvperus and Fi'm-
hristylis (Allan et al.). Leucoanthocyanins are reported from species ofC(arex.,
Cyperus, Du/lichium, Kvllinga, and Scirpus. Anthocyanins are absent from the
family (Harborne; Harborne et al.).
Flavonoids occur in many genera (Kukkonen, 1969; Harborne). Recently,
Harborne and collaborators have done much to expand what is known about
flavonoids in sedges. Among this class of compounds are aurones, which give
a yellowish tint to the inflorescences of many sedges. These are absent from
the Gramineae and the Juncaceae. Flavonols were present in only 15 percent
of 11 genera tested by Harborne. Flavonoid aglycones, especially quercetin and
luteolin, are widespread in the family, as are proanthocyanidins (particularly
in the leaves). Harborne and colleagues (p. 765) concluded that there are "no
dramatic correlations between flavonoid distribution and higher level classi-
fication of the Cyperaceae." However, certain genera or subgenera are distin-
guished chemically from closely related groups (see under (C'perus and Ahild-
gaardia). Flavonoid profiles have been shown to distinguish between related
taxa in Carex and (Cperus (discussed under those genera).
Metcalfe presented much useful information on the anatomy of the Cyper-
aceae, including clear illustrations and insightful comments on the taxonomic
significance of anatomical features. Many of his descriptions were derived from
studies of specimens collected in the Southeast, particularly Florida.
Developmental anatomy and morphology have received some attention (Bar-
nard). The apex of spikelets in all examples studied conforms to the tunica-
corpus pattern. Periclinal division of dermatogen and hypodermal cells gives
rise to tissues that develop into the scales subtending flowers (Scirpus. Cvperus),
the carpels (in all species), the perianth bristles (Scirpus). and the perigynia
(({arex).
The first fossil remains of the Cyperaceae date from the Eocene. Fruits of
Carex, Scleria, and Scirpus are known from the Eocene and Oligocene of
Eurasia and North America; those of Du/ichium and Cladium from the Oli-
[voL. 68


gocene and Pliocene of Europe. Reports of Cyperaceae from pre-Tertiary strata
(i.e., Caricopsis Samylina) are not considered reliable (Daghlian).
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BERGGREN, G. Atlas of seeds and small fruits of northwest European plant species. Part
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TUCKER, CYPERACEAE
365
1987]


JOURNAL OF THE ARNOLD ARBORETUM
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366
[VOL. 68


TUCKER, CYPERACEAE
--, C. A. WILLIAMS, & K. L. WILSON. Flavonoids in leaves and inflorescences of
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species.]
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Zussamenfassung. Aquilo 6: 18-42. 1967.*
1987]


JOURNAL OF THE ARNOLD ARBORETUM
Flavonoid chemistry of the Cyperaceae: a preliminary survey. Mitt. Bot. Staats-
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Special features of the inflorescence structure in the family Cyperaceae. Ann.
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KuiNTH, C. S. Enumeratio plantarum. Vol. 2, Cyperographia synoptica. . . iii t 591
pp. Stuttgart and Tuibingen. 1837. [Worldwide monograph.]
Li MAOI T, E., & J. DI AI.SNF. Traite general de botanique descriptive et analytique.
viii � 746 pp. Paris. 1868. [Cyperaceae. 600-604: synopsis by tribes (Cypereae.
Scirpeae. Hypolytreae, Schoeneae (Rhynchosporeae), Scleriae, Cariceae); represen-
tative illustrations; summary of materia mednica.
LIRMAN, J. C., & J. RA .NAl. La teneur en isotopes stables du carbone chez les Cyper-
acees: sa valeur taxonomique. Compt. Rend. Acad. Sci. Paris, D. 275: 1391-1394.
1972. [List of C3 and C. genera and subgenera; determined by U" carbon values.]
LiOYIv, N. P. H., & H. W. WooL notsi. Comparative aspects of photosynthesis, pho-
torespiration, and transpiration in four species of Cyperaceae from the relict flora
of Teesdale, northern England. New Phytol. 83: 1-7. 1979. [Carex. capillaris. C.
ericetorum. Eriophorum latifliuimn, and Kohresia simpliuscula.]
Lovicii, J. H. The flower and the bee. xvii i 286 pp. New York. 1918. [Flies and beetles
feed on sedge pollen.]
MARTIN, A. C. The comparative internal morphology of seeds. Am. Midl. Nat. 36: 513-
660. 1946. [Cyperaceae. 534.]
MATTFELD, J. Zur Morphologic und Systematik der Cyperaceae. Proc. 6th Int. Bot.
Congr. 1: 330-332. 1935.
MCAIEE, W. 1.. Wildfowl food plants: their value. propagation, and management, ix -
141 pp. Ames, Iowa. 1939. [Cyperaceae. 35-45.]
MEusEUs. A. I). J. Interpretive floral morphology of the Cyperaceae on the basis of the
anthoid concept. Acta Bot. Neerl. 24: 291-304. 1975. [Defends theory that simple
"bisexual" flowers of Cyperaceae are synanthial in origin.]
Mi A'.LEE, C. R. Anatomy of the monocotyledons. Vol. 5. Cyperaceae. 597 pp. London.
1977.
MORA, L. . Beitrage Zur Entwicklungsgeschichte und vergleichenden Morphologic der
Cyperaceen. Heitr. Biol. Pflanzen 35: 253-341. 1960.
N APxR, D. M. Cyperaceae of East Africa. I. E. Afr. Nat. Hist. Soc. Natl. Mus. Jour.
24: 1-18. 1964a. [Floristic account; Caricoideae.] II. Ibid. 23-45. 1964b. [Scleriae,
Schoeneae.] III. Ibid. 25: 1-27. 1965. [Scirpeae, Cypereae (Lipocarpha).] IV. Ibid.
26: 1-17. 1966. [Cypereae (Cy perus).]
Nri:s VON ESiENiHE(K, C. G. Debersicht der Cyperaceengattungen. Linnaea 9: 273-306.
1834. [Synopsis of tribes, genera, and subgenera.]
NoBI. , R. E., & P. K. MUtRPHY. Short term effects of prolonged backwater flooding on
understory vegetation. Castanea 40: 228-238. 1975. [Tensas Parish, Louisiana: pro-
longed spring flooding of Mississippi River reduced populations of Cy'perus and
C arex species.]
OtnD EN, E. C. Anatomical patterns of some aquatic vascular plants of New York. New
York State Mus. Bull. 424. v + 133 pp. 1974. [Many southeastern species: Rhyn-
chospora capitellata (Michx.) Vahl misidentified as R. glomerata (L.) Vahl.]
O'Nru tL, H. T. The sedges of the Yucatan Peninsula. Carnegie Inst. Wash. Publ. 522:
249-322. 1940. [Keys, detailed descriptions.]
PACnH, E. M. Food-plant catalogue of the aphids of the world, including the Phyllox-
eridae. Maine Agr. Exper. Sta. Bull. 393. 431 pp. 1935. [Cyperaceae, 66-69; 23
genera of aphids recorded from species of Care., C'perui., Eriophorum, "itmbristylis.
and Scirpus.]
368
[VOL. 68


TUCKER, CYPERACEAE
PLOWMAN, A. B. The comparative anatomy and phylogeny of the Cyperaceae. Ann.
Bot. 20: 1-33. 1906.
RADFORD, A. E., H. E. AHLES, & C. R. BELL. Manual of the vascular flora of the Carolinas.
lxi + 1183 pp. Chapel Hill, North Carolina. 1968. [Cyperaceae, 168-255.]
RAYNAL, J. Repartition et evolution des modes de photosynthese chez les Cyperacces.
Compt. Rend. Acad. Sci. Paris 275: 2231-2234. 1972.
. Notes cyperologiques: 19. Contribution A la classification de la sous-famille des
Cyperoideae. Adansonia 13: 145-171. 1973.
Notes cyperologiques: 33. Melanges nomenclaturaux (2). Ibid. 17: 273-280.
1978. [Validation of tribe Fimbristylideae.]
RIKLI, M. Beitrage zur vergleichenden Anatomie der Cyperaceen mit besonderer Beriick-
sichtigung der inneren Parenchymscheide. Jahrb. Wiss. Bot. 27: 485-580. 1895.
[Kranz anatomy in Cyperus, Fimbristylis, and Bulbostylis.]
SAVILE, D. B. O. A study of the species of Cintractia on Carex, Kobresia, and Scirpus
in northern America. Canad. Jour. Bot. 30: 410-435. 1952. [Rusts.]
SCHULZE-MOTEL, W. Entwicklungsgeschichte und vergleichend-morphologische Unter-
suchungen im Bliitenbereich der Cyperaceae. Bot. Jahrb. 78: 129-170. 1959.
Cyperales. Pp. 602-607 in H. MELCHIOR, Engler's Syllabus der Pflanzenfamilien.
ed. 12. Berlin. 1964.
SMITH, R. J., JR., W. T. FLINCHUM, & D. E. SEAMAN. Weed control in U. S. rice
production. U. S. Dep. Agr. Agr. Handb. 497. iv + 78 pp. 1977. [Photographs,
descriptions, discussions of Rhynchospora corniculata, 53, 54; Scirpus mucronatus.
S. fluviatilis, S. acutus, 54, 55; Fimbristylis autumnalis, F. miliacea, 59, 60; Ele-
ocharis spp., 69, 70; Cyperus spp., 70-72.]
STACE, C. A., ed. Hybridization and the flora of the British Isles. xiii + 626 pp. London
and New York. 1975. [Scirpus, 510-512; Eleocharis, 512, 513; Schoenus, 513;
Carex, 513-540.]
STANDLEY, P. C. The Cyperaceae of Central America. Fieldiana Bot. 8: 239-292. 1931.
[Synoptic account.]
TEERI, J. A., L. G. STOWE, & D. A. LIVINGSTONE. The distribution of C, species of
Cyperaceae in North America in relation to climate. Oecologia 47: 30-310. 1980.
[Floristic study; percentage of C4 species decreases with latitude; cf. TUCKER, 1986b,
under Cyperus.]
THORNE, R. F. A phylogenetic classification of the Angiospermae. Evol. 9: 35-106.
1976.
TIETZ, H. M. An index to the described life histories, early stages, and hosts of the
Macrolepidoptera of the continental United States and Canada. 2 vols. vi + 1041
pp. Sarasota, Florida. 1972. [Host references published through 1950; Carex, 852;
Cyperus, 875; Eleocharis, 881; Scirpus, 991, 992.]
TORREY, J. Monograph of the North American Cyperaceae. Ann. Lyc. Nat. Hist. New
York 5: 181-448. 1836. [Classic; a thorough study.]
VANHECKE, L. Embryography of some genera of the Cladiinae and the Gahniinae (Cy-
peraceae) with additional notes on their fruit anatomy. Bull. Jard. Bot. Natl. Belg.
44: 367-400. 1974. [Embryos and fruits of Cladium and Schoenus; illustrations.]
VEKEN, P. VAN DER. Contribution A l'embryographie systematique des Cyperaceae-
Cyperoideae. Bull. Jard. Bot. Natl. Belg. 35: 285-354. 1965. [Descriptions of most
of our genera (remainder treated by VANHECKE) with taxonomic comments; illus-
trations.]
WINFREY, H. J., & G. L. SAMSEL. Preliminary effects on algal succession resulting from
nutrient enrichment of two central Virginia ponds with different trophic states.
Castanea 38: 140-152. 1973. [Bulbostylis capillaris, Carex spp., Cyperus strigosus,
Scirpus americanus, S. cvperinus.]
369
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JOURNAL OF THE ARNOLD ARBORETUM
KEY 10 THE GENERA OF CYPERA(EAE IN THE
SOUTHEASTERN UNITED STATES
General characters: perennial (occasionally annual), of/ten rhizomatous herbs of diverse,
usually wet, often disturbed habitats; rhizomes frequently present: leaves linear, the sheaths
usually closed: inflorescences simple or variously branched, lateral or crowded at the apices
of the culms; flowers perfect or imperfect (the plants very rarely dioecious), borne in the
a.\ils of scales or in perigynia, perianth bristles present or absent; fruit an achne; embryo
small; endosperm abundant.
A. Flowers perfect (stalninate or carpellate flowers infrequently formed at base or apex
of spikelets).
B. Scales of the spikelets spirally arranged.
C. Achenes without obconical or pyramidal apical tubercle, but sometimes with
persistent swollen style base much less than half as wide as the achene.
I). Achenes subtended by 15-20 cinnamon or whitish, silky bristles 5-10
times as long as the achenes. .. . . . . . . . . .. . 2. Eriophorum.
D. Achenes subtended by bristles at most 3 times as long as the achenes, or
with bristles lacking.
E. Inner whorl ofperianth bristles with expanded spongy petaloid blades.
........ .... . . ....... . 3 . F uirena .
E. Perianth bristles absent or lacking expanded blades.
F. Bulbous base of style persistent on mature achenes............
............................ . 6 . B ulbostvylis.
F. Base of style not persistent.
(i. Styles tim briate. .................... . 5. Fim bristvlis.
Ci. Styles smooth.
H. Spikelets maturing a single achene; bristles absent .....
......... . 14. (ladium .
H. Spikelets maturing several to many achenes: bristles usu-
ally present.
I. Spikes and spikelets borne on rays. rarely sessile:
achenes and scales appressed to rachilla... 1. Scirpus.
1. Spikelets sessile: achenes and scales borne at right an-
gles to rachis .. . . . . . . . . .. 10. Lipocarpha.
C. Achenes with pyramidal or obconical apical tubercle 10: to nearly as broad as
the achene.
J. Leaf blades absent;: inflorescences unbranched, a single spikelet termi-
nating the culm . ..... . . . ..... ........... 4. Eleocharis.
J. Leaf blades present; inflorescence of several to many spikelets, some usu-
ally borne on branches. . . . . . . . . . . . . . 1. Rhvnchospora.
B. Scales of the spikelets distichously arranged.
K. Perianth bristles absent.
L. Plants bulbous-thickened basally: style base sclerilied, persistent: spikelets
1(-3). ......... ......... .................. ... . . 7. ,Abildgaardia.
L. Plants not bulbous-thickened basally: style base soft, deciduous; spikelets
numerous.
M. Inflorescences branched: spikelets 1- to many-flowered: rachilla elon-
gate: scales broadly rounded. ....................... 8. ( tperus.
M. Inflorescences unbranched (spikes sessile); spikelets 1- (infrequently
2-)flowered; rachilla not or barely elongate: scales conduplicate, con-
spicuously keeled . . . ... .......... . 9. K //llinga.
K. Perianth bristles present.
N. Leaves cauline: inflorescences several, axillar. .. 12. Dulichiu.
N. Leaves basal: inflorescence solitary, terminal. .......... 13. Schoenus.
370
[VOL. 68


TUCKER, CYPERACEAE
A. Flowers strictly imperfect.
0. Achenes naked, often borne on a discoid hypogynium. .......... 15. Scieria.
0. Achenes enclosed in perigynia.
P. Spikes single, white: leaf blades broadly lanceolate, the apices broadly round-
ed, the midvein not distinguishable from other veins. .. 16. Cvmophyllus.
P. Spikes I to several, greenish, yellowish green, or light brown; leaf blades
linear, the apices acute, the midvein much larger and more conspicuous than
other veins . .............. ... ...................... . 17. Carex.
Subfamily CYPEROIDEAE
Tribe SCIRPEAE Kunth ex Dumortier, Fl. Belg. 143. 1827.
1. Scirpus Linnaeus, Sp. Pl. 1: 47. 1753; Gen. Pl. ed. 5. 26. 1754.
Small to medium-sized perennials or annuals of shallow fresh or tidal waters,
disturbed moist soils, moist [mesic to dry-mesic] woodlands, marshes, open
mountaintops, and grassy balds. Roots fibrous; perennial species with rhizomes
short, branched, producing loose to dense tussocks of culms; annual species
without rhizomes, forming dense clumps of culms. Culms trigonous (with
planar, concave, or slightly convex surfaces) or terete, smooth throughout or
scabrellate distally. Leaves all basal or scattered along the culm; sheaths closed,
smooth or sometimes with conspicuous cross veins, greenish white, reddish
brown, or blackish; blades flat, conduplicate, or subterete, 1/2 to nearly as long
as the culm, stiffor arching (limp when growing underwater); stomata paracytic;
chlorenchyma not radiate: longitudinal air chambers often present. Involucral
leaves (1 or) 2-10, the blades resembling cauline ones but sheaths generally
much shorter, approximate at the summit of the culm or rather widely spaced
over the upper '/ of it, horizontal to ascendent. or the longest nearly vertical
and simulating a continuation of the culm. Inflorescences composed of primary
and secondary (sometimes tertiary) rays, in many species reduced to glomer-
ulate clusters or heads, in some to a cluster of several more or less sessile
spikelets or a single sessile spikelet: prophylls of the rays tubular, obtuse to
acute apically, smooth but usually conspicuously costate; primary rays smooth,
or scabrellate distally or throughout, terete. stiff or flexuous, secondary (and
sometimes tertiary) rays similar to primary ones, but shorter and usually more
slender. Spikelets ovoid to linear-oblong. Scales (3 to) 20 to about 100, spirally
arranged and closely imbricate, with 2 lowermost sterile and others fertile, all
deciduous at maturity, ovate to oblong, with 1-9 subtle to conspicuous nerves
and sometimes a conspicuous midrib, the apex obtuse to acute, entire or mu-
cronulate to strongly cuspidate, the awn straight to strongly excurved. Flowers
perfect, protogynous. Perianth bristles 3-6(-8) or lacking, smooth or retrorsely
scabrellate, straight, highly curled, or crinkled at maturity, from '/3 to 4 times
as long as the mature achene, deciduous or remaining attached to the mature
achene. Stamens (2 or) 3: filaments slender, about equaling the subtending
scales; anthers broadly ellipsoid to narrowly linear, the apices of the connectives
in some species prolonged as subulate appendages up to /4 the length of the
anther, sometimes tipped with crystalline prickles; pollen uniaperturate, sub-
spheroidal in polar view and triangular to obovoid in equatorial view, psilate,
1987]


JOURNAL OF THE ARNOLD ARBORETUM
/
Sf
h II
/ ~ ~7~$�~, (
372
[vol.. 68


TUCKER, CYPERACEAE
bi- or trinucleate. Styles capillary; stigmas 2 or 3, about equaling the style in
length. Achenes lenticular to trigonous, equilateral in transverse section, or
slightly to strongly dorsiventrally flattened, the base stipitate or cuneate, the
apex apiculate, beaked, or entire, the surface essentially smooth, finely pitted,
reticulate, or rugulose. Embryos ellipsoid, turbinate, or fungiform, the radicle
lateral or basal. Base chromosome numbers 5, 7. LECTOTYPE SPECIES: Scirpus
sylvaticus L.; see Hitchcock & Green, Prop. Brit. Bot. 118. 1929. (Latin name
for a bulrush, probably Scirpus Tahernaemontani Gmelin.)-BULRUSH, REED,
CLUB-RUSH, WOOL-GRASS, THREE-SQUARE.
Scirpus, the third-largest genus of the Cyperaceae, with about 300 species
worldwide, is best represented in temperate regions. North America (including
Mexico), with about 80 species, is the center of diversity. Only about 15 species
occur in the West Indies and Central America, and about 30 in all of South
America, most of these in Argentina and Chile. Twelve species occur in Europe,
and perhaps 50 in Africa. It is difficult to estimate the number of species in all
of Asia; 24 grow in the Soviet Union, and ten in Malesia. A recent synopsis
included 44 in Australia (Wilson).
Studies in Scirpus have been hampered by lack of a worldwide treatment
(such as those prepared for several other large genera of the family, i.e., Carex,
Cyperus, Eleocharis, and Rhynchospora). Some botanists (e.g., Wilson, Koy-
ama) have recognized each of the sections at the generic level. Most American
authors (Fernald, Schuyler), however, have recognized the genus in a broad
sense: this traditional circumscription is accepted here. Although several re-
searchers have lamented the "diverse" nature of the genus, most of the kinds
of variation that are represented in Scirpus are also present in Cyperus, which
FIGURE 1. Scirpus sect. JUNCO-SCIRPUS. a-h, S. Tabernaemontani (S. validus): a, un-
derwater rhizome collected late in season, apex at right (note remains of shoot of current
season and developing shoots of next year's growth), x '/2; b. apex of culm with inflo-
rescence, x 1: c. single spikelet with lower flowers past anthesis (filaments visible), upper
ones with anthers visible and styles exserted, x 12; d, flower and subtending scale
removed from spikelet, view of adaxial surface, stigmas exserted, anthers still included
(note barbed bristles), x 20: e, flower, showing different maturation of stamens, the
stamen to right with elongate filament and anther ready to dehisce, the middle stamen
with anther fallen, the stamen to left before elongation of filament, x 12: f, spikelet at
stage later than that in "c." immature achenes below and flowers with receptive stigmas
above, x 6: g. mature achene with persistent bristles, x 12; h, same, in vertical section,
fruit wall hatched, endosperm stippled, embryo unshaded (seed coat too thin to show),
x 12. i, j, S. cespitosus: i, spikelet, lower flowers with persistent filaments, upper ones
with exserted stigmas, involucral bracts greatly reduced, scalelike, spikelet solitary, x 6:
j, achene with smooth bristles. x 12. k. 1, S. koilolepis: k. solitary spikelet, subtended
by scalelike involucre, scales keeled, x 6: 1. mature. trigonous. bristleless achene, x 12.
m-o, S. Erismaniae: m, basal flower in axil of leaf. x 6; n, achene from basal flower, x
12; o, achene from cauline spikelet, x 12. p, S. cvperinus: achene with elongate bristles,
x 12.
373
1987]


JOURNAL OF THE ARNOLD ARBORETUM
has traditionally been maintained as one genus. Moreover, there has yet to
appear a thorough study of Scirpus that presents compelling arguments for
recognizing Schoenoplectus (Reichenb.) Palla, Trichophorum Pers., Baeothryon
A. Dietr., and other segregate genera. Many useful papers on the taxonomy of
single species or groups of species have been written by several authors, most
notably Schuyler.
The achenes of species of Scirpus are probably dispersed after being eaten
by waterfowl (McAtee) (wild ducks in the case of S. paludosus Nelson). Most
are digested, but those that survive have 94 percent germination, compared
with two or three percent for those treated with acid or alkali, and nine percent
after fermentation treatment (Low). Light is required for germination (Isely).
Achenes of many species, particularly S. ctperinus (L.) Kunth, are probably
dispersed by the wind, although their long, contorted perianth bristles likely
also cause them to cling to fur or feathers.
Some 30 species of Scirpus. representing six sections, occur in our area.
Following is a brief account of these.
Species with leafy stems are classified in three sections. In all of these, leaves
are borne along the length of the culm. while in plants of other sections they
are basal. Schuyler (1961, 1962. 1963. 1964, 1966, 1967a, 1967b, 1967c.
1971b) has studied the species with leafy stems and has provided most of the
available information on morphological variation, cytology, hybridization, and
distribution.
Section SCiRPts (sect. Taphrogeton (Reichenb.) Ascherson; plants leafy
stemmed: spikelets in dense heads: achenes ellipsoid, with perianth bristles
straight, about as long as the achenes) includes the type species, the Eurasian
Scirpus sy/vaticus L., n = 31, 32. The section is represented by seven species
in our area, which fall into three groups. The first includes the North American
relatives ofS. sylvaticus, among which the only representative occurring in the
Southeast is S. evpansus Fern., n ..= 32. This bulrush grows mostly in the
Northeast, but it ranges south in the Appalachians to northern Georgia and
northern Alabama. A second eastern North American species, S. microcarpus
Presl (S. ruhrotinctus Fern.), n = 33, occurs southward to the uplands of West
Virginia and also in western North America and eastern Asia.
The second group (leaves tristichous. spikelets in glomerules, plants typically
viviparous, bristles straight) includes what was treated as Scirpus atrovirens
Willd. by Fernald (1950). Schuyler (1967a. 1967b, 1967c) demonstrated that
there are four species in this group that can be distinguished morphologically
and separated geographically and phenologically. Scirpus georgianus Harper,
n = 25, 26, 27, is the most common in our area (specimens examined from
every state). It lacks a perianth and leaf cross veins (these are present in the
more northern S. utrovirens Willd., n = 28, which is occasional in our area
from the Ridge and Valley Province westward). Scirpus Iattorianus Makino.
a = 28, is a northeastern species known in our area from only six collections
from the uplands of North Carolina, Tennessee. and Alabama. Scirpus /laci-
difolius (Fern.) Schuyler, n = 27, is endemic to river bottoms in eastern Virginia
and northeastern North Carolina.
The third group of sect. SCIRPIS (leaves distichous, spikelets in glomerules,
374
[voI. 68


TUCKER, CYPERACEAE
plants not viviparous, bristles contorted) is represented by a single species in
North America, Scirpus polyphyllus Vahl, n = 29, which is known from all the
Southeastern States.
Section ANDROCOMA (Nees) Bentham (plants leafy stemmed; perianth bristles
smooth, approximately as long as the subtending scales) is represented in our
area by three species. Scirpus pendulinus Muhl. (S. lineatus auct.. non Michx.),
n = 20, has the greatest range of the three, occurring from Maine to Minnesota
south to the Gulf Coast. Scirpus lineatus Michx. (S. fontinalis Harper), n =
18, is found along the Coastal Plain from Virginia to Florida; S. divaricatus
Ell., n = 14, has a similar range but is found westward to Louisiana.
Section TRICHOPHORUM (Pers.) Darl., the wool grasses (plants leafy stemmed;
perianth bristles contorted, several times longer than the achenes), comprises
several species of cold-temperate regions. At maturity the elongate, crinkled
bristles give the spikelets and the inflorescences a woolly appearance. Extensive
hybridization in this group has resulted in a nomenclatural mire of species,
varieties, and forms. Schuyler (1962, 1967a) has carefully documented infra-
specific variation, cytology, and hybridization: he concluded that only a single
species, Scirpus cyperinus (L.) Kunth (including S. rubricosus and S. eriophorum
Michx.), n = 33. should be recognized in the Southeast. Three others that
hybridize with S. cvperinus, S. pedicellatus Fern., n = 34; S. Longii Fern.,' n =
33; and S. atrocinctus Fern., n = 34, occur in the Northeast.
Section OXYCARYUM (Nees) Beetle (plants rhizomatous; heads of spikelets
ovoid, pedunculate: scales acute. excurved) is represented in the Southeast by
a single species. Scirpus cubensis Poeppig & Kunth. In our area the species
occurs from southern Florida to Louisiana in brackish or freshwater marshes.
The affinities of this section are unclear, and no chromosome counts are avail-
able.
Section BOLBOSCHOENUS (Ascherson) Beetle (plants tall; spikelets large, few;
scales awned, pubescent) is represented in our area by two species of freshwater
or tidal wetlands. Scirpus robustus Pursh grows in tidal marshes and estuaries
from eastern Canada to Texas. A second species, S. cylindricus (Torr.) Britton,
occurs in marshes from Delaware to Georgia. It was confused with S. robustus
and S. etuberculatus until it was restudied by Schuyler (1975). The third species.
S. etuberculatus (Steudel) Kuntze. grows in brackish waters and is known near
the coast from Delaware to Louisiana. It is morphologically transitional to the
next section (Fernald, 1950).
Section JUNCO-SCIRPUS Syme' (sect. Ptero/epis Beurl., sect. Schoenoplectus
(Reichenb.) Bentham) (plants tall; culms often leafless; involucral leaves 1 or
2, more or less erect; achenes sessile, beaked, with bristles persistent) is rep-
resented in the Southeast by seven species. Scirpus pungens Vahl (S. americanus
'Reported from North Carolina bh ( appel and Radlbrd and colleagues. I was unable to locale anm
specimens to substantiate this. \ccording to Schu lcr (19%2 pers. comm.). records of .S. o.W from
south of New Jersey were the result of misidentifications of S. ciperinu.
4ScTrpit sect. JUNC( -sCrIRPIS S me in Sowcrbi . Engl. Hot. cd. 3. 10: 62 1870. l.ErCTOTi P- SPF( I-S
(here designated): S. Iacuosris 1. Syme included three species in this section. S. lacustris, S. triqutler
L.. and S. pungens Vahl: S. /acu'itr is the only one with terete culms suggesting those of plants of
the genus Juncus L., a feature emphasized by the sectional name.
1987]


JOURNAL OF THE ARNOLD ARBORETUM
auct., non Pursh), n = 39, of sunny wetlands, is widespread in temperate North
America and occurs in all the southeastern states. It is closely related to S.
americanus Pursh (S. Olnevi Gray), n = 39, a taller, thicker-stemmed species
of tidal, alkaline, or saline marshes from Massachusetts to Florida and west to
southern California. The two species occasionally hybridize in brackish upper
edges of tidal marshes, but in general they are isolated ecologically. A recently
described species, S. deltarum Schuyler, n = 39, occurs in the Mississippi Delta
region, the Mobile Bay area, and disjunctly in the prairie marshes of eastern
Kansas and Missouri. A fourth species, S. subterminalis Torrey, n = 37, is
widespread in eastern North America but is known in the Southeast from only
a few collections from the Coastal Plain and Piedmont of North and South
Carolina. Two growth forms exist: submersed, in which the leaves are filiform
and flaccid, and terrestrial or stranded, in which they are conduplicate and stiff
(Schuyler. 1972b). The highly reduced inflorescence consists of a single spikelet
subtended by one erect involucral bract. The species has an unusual photo-
synthetic metabolism: the tissues of the stem, leaf, and rhizome accumulate
malic acid at night, providing a reservoir of fixed carbon for photosynthetic
reactions during daylight (Beer & Wetzel). Such physiology is similar to that
of terrestrial plants having crassulacean-acid metabolism.
The remaining three species of sect. JUN(CO-SCIRPus were once segregated as
sect. Pterolepis (Fernald, 1950). These reportedly differ in having plumose
bristles and pedunculate clusters of spikelets. However, on a worldwide basis
several extraregional species are intermediate with respect to these two char-
acters: Koyama (1963) therefore concluded that the two sections should be
merged. Scirpus Tahernaemontani Gmelin (S. validus Vahl), n = 21, grows in
freshwater marshes nearly throughout the United States and southern Canada
and in much of the Old World: it is common throughout the Southeast. Scirpus
acutus Bigelow, n = 19. a species of the Midwest and Great Plains, is represented
in our area by a few collections from North Carolina and Tennessee. Dabbs
studied these two species in Saskatchewan and found that they were morpho-
logically distinct. Hybrids were occasionally found, but these were sterile and
spread only by rhizomes. A western species, S. ca/ilornicus (C. Meyer) Steudel,
n = 34, is known from a few places in Louisiana, Mississippi, and South
Carolina. Other North American species of Scirpu. lack its plumose perianth
bristles. Scirpus heterochaetus Chase, n = 19, might be found in the north-
western part of our area: it is a species of quiet calcareous waters of the St.
Lawrence and upper Mississippi drainages.
Section BAEOTnRYON Dumort.' (plants caespitose, often forming tussocks:
leaves basal: inflorescences ofa single terminal spikelet: involucral bract greatly
reduced, resembling a fertile scale of the spikelet) is represented by four species
in northeastern North America. Only one of these, the circumboreal Scirpus
ccspitosus L., reaches our area, growing in the grassy balds of the high mountains
of North Carolina. Georgia, and Tennessee. The southeastern populations are
disjunct from the nearest occurrences of the species in the northeastern United
', ipr/O sect. BAOIIIRYoN I)umort. 1. Ielg. 143. 1827. Fernald (1947, l950) and other authors
hale attributed the sectional name to Endlicher (Gen. P. 1 18. 1 S136).
376
[VOL. 68


TUCKER, CYPERACEAE
States (in the Adirondack Mountains of New York) by some 1200 km. A
widespread but easily overlooked species of the northeastern and midwestern
United States, S. verecundus Fern., has not yet been collected in our area but
might occur in the uplands of North Carolina, Tennessee, or Arkansas. It is
perhaps the most mesic species pf the genus in North America, inhabiting dry
woodlands and basic ledges, in contrast to the aquatic habitats of most species
of Scirpus.
Section ISOLEPIS (R. Br.) Griseb. (plants annual: inflorescences unbranched:
spikelets sessile, few) is represented in our area by live species. Scirpus koilolepis
(Steudel) Gleason probably occurs in all the states in our area, as well as in
the Midwest and the Great Plains. The remaining species are much less frequent
and are local in range. Scirpus Erismaniac Schuyler, n = 5, is recorded from
Georgia, western Florida, and Alabama. This species produces basal spikelets
on very short culms (see FIGURE 1), as do several African species of this section
(Haines). Scirpus molestus M. C. Johnston, described from Texas, also occurs
in southern Louisiana. The remaining species have perianth bristles (in most
collections) and have been distinguished by some authors (e.g., Fernald, 1950)
as sect. Actaeogeton (Reichenb.) Beetle. Scirpus Hallii Gray, n = 11, known
from widespread localities in the eastern United States, has been collected in
Georgia; and S. Purshianus Fern.. n = 19, a primarily northeastern species. is
known in the Southeast from North and South Carolina. Tennessee, and Geor-
gia.
REFERENCES:
Under family references see BADEN et al.: BARNARD: BARROS (1935); BENTHAM: BLASER
(1941 a, 1941 c): BURKHALTER: CLARKF (1908, 1909); EITEN (1976a): EviYs & ROBERTSON:
FERNALD: GOOD et al.: HAINES: HARRIS & MARSHALL; HOLTTUM: G. E. HUTCHINSON:
J. HUTCHINSON; J. H. KERN; KRAL; KUKKONEN (1969); LEMAOUT & DECAISNE; LERMAN
& RAYNAL; MCATEE; MEEUSE; METCALFE; MORA; O'NEILL; RADFORD et al.; RAYNAL
(1972, 1973); RIKLI; SAVILE: SCHULZE-MOTEL (1959, 1964); STANDLEY; TEERI et al.; TIETZ:
TORREY: and WINFREY & SAMSEI .
BEER, S., & R. G. WETZEL. Photosynthetic carbon metabolism in the submerged aquatic
angiosperm Scirpus sutecrninalis. Pl. Sci. Lett. 21: 199-207. 1981.
BEETLE, A. A. Studies of the genus Scirpus L.. V. Notes on the section Actaeogeton.
Am. Jour. Bot. 29: 653-656. 1942.
. A key to the North American species of the genus Scirpus based on the achene
characters. Am. Midi. Nat. 29: 533-538. 1943.
Studies in the genus Scirpus VII. Conspectus of sections represented in the
Americas. Am. Jour. Bot. 31: 261-265. 1944. [See review by FERNALD (1947).]
. Sedge boats in the Andes. Jour. N. Y. Bot. Gard. 46: 1-4. 1945. [Boats in Lake
Titicaca made from culms of S. Tatora Steudel.]
. Cyperaceae: Scirpeae. Scirpus. N. Am. Fl. 18: 479-504. 1947. [Keys, descrip-
tions.]
. Annotated list of original descriptions of Scirpus. Am. Midi. Nat. 41: 453-493.
1949. [Worldwide.]
. Bulrushes (Scirpus) and their multiple uses. Econ. Bot. 4: 132-138. 1950. [Sum-
mary of economic importance.]
CAPPEL, E. D. The genus Scirpus in North Carolina. Jour. Elisha Mitchell Sci. Soc. 70:
85-91. 1954. [Keys, descriptions, distributions.]
377
1987]


JOURNAL OF THE ARNOLD ARBORETUM
DAI)xi, D. L. A study of Scirpus acutus and Scirpus validus in the Saskatchewan River
delta. (anad. Jour. Bot. 49: 143-153. 1971.
F RN LI0). M. L. Studies of the North American species of Scirpus. Rhodora 45: 279-
296. 1943.
. Unverified bibliography of .cirpu.s. Ibid. 49: 49-52. 1947. [Review of BET II
(1944); most infrageneric names used by Beetle were not published at the ranks
indicated.,
IH x\NsiTER. R. Recovery, productivity. and phosphorus content of selected marsh plants
after repeated cuttings. 81 pp. Unpubl. M.S. Thesis, Univ. Wisconsin, Oshkosh.
1975.* [S;. acutus and S. validus recovered well even when harvested as frequently
as every two weeks; .S. fluriatilih, however, decreased in number and size of shoots
under this treatment: information from (iooi) c a!.]
Is 1 v, D. A study of the conditions that affect the germination of Scirpus seed. Cornell
S!niv. Agr. Exper. Sta. Mem. 257. 28 pp. 1952. [Light enhances germination in many
species.]
KoV x.M, T. Taxonomic study of the genus Scirpus l.inne. Jour. Fac. Sci. FUniv. Tokyo
Hot. 7: 271-366. 1958. [Broad generic concept including Lriophorum, Fuirena.]
--- The genus Scirpux Linn. Some North American aphylloid species. Canad. Jour.
Bot. 40: 913. 1962.
- The genus Scirpio Linn. C(ritical species of the section Ptirolepiv. Ibid. 41: 1108-
1131. 1963.
- & . (C. STroNI. The genus oSiirpus in the Hawaiian Islands. Bot. Mag. Tokyo
73: 288-294. 1960.
L irxirc.A, B. W., & A. E. S( it i I R. Scirpui trqululctr established along tidal portions
of the Columbia River. Bartonia 50: 23, 24. 1984.
Low, J. Germination tests of some aquatic plants important as duck foods. 27 pp.
Utnpubl. B.S. Thesis, Utah State Univ., Logan. 1937.* [Summarized by G. E.
Hti r iiiNSON.]
I-vi . K. A. Moderne oppfatning av slekta .Sirpus L. Blttia 29: 141-147. 1971.*
RAYMOND, M. Additional notes on some Southeast Asian Scirpus. Nat. Canad. 84: 111-
150. 1957.
R xxNLS J. Notes cyperologiques 26. Le genre .Schoe'toplectus. 11. L'amphicarpie et le
sect. Supoi. Adansonia, II. 16: 119-155. 1976. [Formation of basal spikelets in
certain African species.]
SAVn i, I). B.O. Some rusts of Sc'irpus and allied genera. Canad. Jour. Bot. 50: 2579-
2596. 1972.
S( ii i-R, A. F. Evidence for the hybrid origin of Sc. irpus Peckit. Rhodora 63: 237-
243. 1961. [Sterile hybrid of NS. atrovircn and S. atroi'nctu.s or S. pedicellatus.]
. Sporadic culm formation in .Sci rpus Long'i. Bartonia 32: 1-5. /1 nnmiobered
pi. 1962. [Report from North Carolina (Fi RNALi), 1943) based on misidentification
ofS. cvpcrinuis: see also S i t!tiit R & SI .xz.]
Notes on live species of .S'cipus in eastern North America. ibid. 33: 1-6. 1963.
[Comments on taxonomy of S. uancisr'acetiu, S. atroirc'i, . dSivaricatus. S.
oifminalis. and .S'. lin'atul.s chromosome counts for each.]
A biosistematic study of the SN'pu/i cyperiinu. complex. Proc. Acad. Nat. Sci.
Phila. 115: 283-31 1. 1964. [Hybridi/ation of l cyperinus and related species.]
. The taxonomic delineation of Scirpus lineatuls and Sciupus pendulus. Not. Nat.
390: 1-3. 1966. [With nomenclatural comments.]
A taxonomic revision of the North American leafy species of Scirpus. Proc.
Acad. Nat. Sci. Phila. 119: 295-323. 1967a. [Keys, descriptions, chromosome num-
bers for 18 species.]
.Sci'rpus Hattorianus in North America. Not. Nat. 398: 1-5. 1967b. [Common
northeastern species previously confused with S. atre/irens and .S. ,icorgianms: south-
ern range limit in North Carolina mountains.]
[voL. 68


TUCKER, CYPERACEAE
A new status for an eastern North American Scirpus. Rhodora 69: 198-202.
1967c. [S. flaccidifolius distinguished from S. atrovirens.]
Three new species of Scirpus (Cyperaceae) in the southern United States. Not.
Nat. 423: 1-12. 1969. [S. Bergsonii, S. Erismaniae, and S. Wilkensii, all from Gulf
Coastal Plain, related to S. Hallii; chromosome counts and specimen citations.]
A new North American aquatic bulrush (Cyperaceae: Scirpus). Ibid. 427: 1-3.
1970. [S. deltarum from Mississippi, Louisiana, Alabama, and Missouri, related to
S. pungens; illustrations.]
. Some relationships in Scirpeae bearing on the delineation of genera. Mitt. Bot.
Staatssam. Miinchen 10: 577-585. 1971 a.
. Scanning electron microscopy of achene epidermis in species of Scirpus (Cy-
peraceae). Proc. Acad. Nat. Sci. Phila. 123: 29-52. 1971b. [Survey of epidermal
features of Scirpus and some species of Eriophorum with comments on taxonomy;
clear, informative photographs.]
. Chromosome numbers of Scirpus Purshianus and S. Smithii. Rhodora 74: 398-
406. 1972a. [Distribution of named forms of each species; S. Purshianus, n = 19;
S. Smithii, n = 20.]
Morphological and anatomical differences in leaf blades of three North Amer-
ican aquatic bulrushes (Cyperaceae: Scirpus). Bartonia 41: 57-60. 1972b. [S. etu-
berculatus, S. subterminalis, and S. Torrevi; illustrations; these closely related species
differ greatly in anatomy of leaf blades.]
. Scirpus cylindricus: an ecologically restricted eastern North American tuberous
bulrush. Ibid. 43: 29-37. 1974. [Illustrations, specimen citations.]
Chromosome numbers of some eastern North American species of Scirpus.
Ibid. 44: 27-31. 1975.
-- & J. L. STASZ. Influence of fire on reproduction of Scirpus Longii. Bartonia 51:
105-107. 1985. [Fire stimulates culm formation and flowering.]
SEIDEL, K. Macrophytes and water purification. Pp. 109-122 in J. TouRBIER & R.
PIERSON, eds., Biological control of water pollution. New York. 1976. [S. lacustris
and other marsh plants used to treat wastewater in artificial marshes in northern
Europe.]
-- & R. Kic KUTH. Biological treatment of phenol-containing wastewater with bul-
rush (Scirpus lacustris L.). Wasserwirtschaft-Wassertechnik 17: 209, 210. 1967.*
[Summarized by Goon et al.]
SERODES, J. B., J. DESCHENES, & J.-P. TOURDIE. Temps de submersion des marais a
scirpe (Scirpus americanus) de l'estuaire du Saint-Laurent. Nat. Canad. 112: 119-
129. 1985.
SMITH, S. G. Natural hybridization in the Scirpus iacustris complex in the north central
United States. Pp. 175-200 in J. G. GiUN(KEL,. ed., Current topics in plant science.
New York. 1969.
Ecology of the Scirpus lacustris complex in North America. Polsk. Arch. Hy-
drobiol. 20: 215, 216. 1973.
SOENDERGAARD, M., & R. G. WETZEL. Photorespiration and internal recycling of CO,
in the submersed angiosperm Scirpus subterminalis. Canad. Jour. Bot. 58: 591-598.
1980.
STEINMANN, F., & R. BRANDLE. Carbohydrate and protein metabolism in the rhizomes
of the bulrush (Schoenoph'ctus lacustris (L.) Palla) in relation to natural development
of the whole plant. Aquatic Bot. 19: 53-64. 1984. [S. lacustris.]
WESTHOFF, V., & M. F. MORZER BRUIJNS. De groeiplaats van Scirpus americanus Pers.
op het Groene Strand bij West-Terschelling. (English summary.) Acta Bot. Neerl.
5: 344-354. 1956. [Optimum habitat for S. pungens at upper edge of tidal marsh,
where the salinity was less than 9 g chlorine/liter; disturbance reduced competition
to the benefit of this species.]
WIISON, K. L. A synopsis of the genus Scirpus sens. lat. (Cyperaceae) in Australia.
379
1987]


JOURNAL OF THE ARNOLD ARBORETUM
Telopea 2: 153-172. 1981. [Keys, descriptions, discussions for 43 species; subgenera
recognized as genera.]
2. Eriophorum Linnaeus. Sp. P1. 1: 52. 1753; Gen. P1. 27. 1754.
Small to medium-sized, single-stemmed or loosely caespitose [densely caes-
pitose or tussock-forming] perennials of bogs, swamps, and pocosins. Roots
fibrous: rhizomes short, horizontal to oblique. Culms terete or nearly so, gla-
brous. Leaves basal and cauline: sheaths glabrous. ligules lacking; blades flat
[conduplicate], the midrib conspicuous, the margins scabrellate, especially dis-
tally; chlorenchyma not radiate: air chambers present. Inflorescences of I to
several sessile or pedunculate spikelets; bracts 1-6, closely spaced at the summit
of the culm, oblique or slightly reflexed [ascendent to erect], sheaths very short,
blades leaflike: rays short [elongate and drooping or absent]. Spikelets oblong-
ovoid: empty basal scales 3-5[-15]. Scales 50-150. oblong-ellipsoid, acute to
obtuse, 1- to 5-nerved, deciduous after the achenes mature. Flowers perfect.
Perianth bristles [6 to] 12 to ca. 50, about equaling the scales at anthesis but
elongating greatly as the achenes mature. Stamens 1 [or 2 or 3]; filaments
flattened; anthers linear [ellipsoid], the apices of the connectives not prolonged.
Styles capillary, glabrous; stigmas 3, about as long as the style. Achenes tri-
gonous. slightly compressed dorsiventrally, oblong-ellipsoid (widest in distal
half), the apex obtuse, apiculate, the base sessile, the surface smooth, glossy.
Embryos more or less turbinate [obconical or ellipsoid], the radicle sublateral.
Base chromosome number 29. TYPE SPECIES: E. vainaltu L.; see Britton &
Brown, Illus. Fl. No. U. S. Canada, ed. 2. 1: 322. 1913. (Name from Greek.
crios, cotton or wool, and phoros, bearing, in reference to the cottony mature
inflorescence.)--COTTON-GRASS. BOG-(C OTTON.
A genus of about 12 species of boreal regions. About eight species are cir-
cumpolar, occurring in both northern Eurasia and northern North America.
There is relatively little endemism. Only Eriophorum virginicum L. occurs in
the Southeast; it ranges from Newfoundland to Minnesota southward and is
known in our area from a few scattered collections made in the mountain bogs
of North ('arolina and Tennessee and the Coastal Plain swamps of North
Carolina, South Carolina, and Georgia (southern limit in the Okefenokee
Swamp). No species of the genus is reported from Missouri or Kentucky, and
only E.. virinic*num occurs in Virginia and West Virginia.
A few workers (e.g., Koyama) have treated the cotton grasses as constituting
Scirpus sect. 'aginuati (Andersson) Koyama, but most have kept Eriophorumn
separate from Scirpus. The two genera are readily distinguished by the number
and the length of the perianth bristles. lriophorinum is divided into two sections
(Goncharov ci a!.). each with about six species: sect. ERIOPHORtUM (sect. 1 -
iginati Andersson) contains those species in which the inflorescence is a single
sessile spike, while sect. PHYv ANTHELA Andersson comprises those (including
E. virpinicunm) in which the inflorescence consists of several pedunculate spikes.
The genus is almost uniform cytologically; ten of the 12 species have been
counted as n- 29. Two are .n 27, and in the case of Eriophorum anguustidlmun
L., n 29 and n = 35 have been reported.
380
[VOL. 68


Hybridization is known among both the Eurasian and the North American
species. Although it is generally not difficult to distinguish Eriophorum virgini-
cum from the other members of the genus, there are species pairs that appear
to intergrade-for example, E. angustifolium and E. viridicarinatum (Engelm.)
Fern. It is surprising that the genus has not received more systematic study,
considering its broad distribution.
The circumboreal Eriophorum alpinum L., n = 29, was placed in Scirpus
(as S. hudsonianus) by Fernald. Following a survey of epidermal features of
achenes of Scirpus and Eriophorum, Schuyler concluded that the species be-
longs in Eriophorum. Its chromosome number also supports this placement.
In the Arctic, species of Eriophorum are dominant and sometimes form a
vegetation type known as "tussock tundra." The plants provide an important
forage for deer and caribou in North America and for sheep, ponies, and
reindeer in northern Europe and Asia. In the United States the plants are seldom
dominant (except in alpine grasslands in limited montane areas). However,
they sometimes form a conspicuous element of fen and bog vegetation because
of their showy fruiting heads.
Wein summarized ecological information about Eriophorum vaginatum, a
circumboreal tussock-forming species. Species of Eriophorum occurring in the
eastern United States are rhizomatous or rather loosely caespitose. There is
much information on the autecology and physiological ecology of the genus,
although nearly all is derived from studies of E. vaginatum.
Despite its abundance in arctic regions, Eriophorum has conspicuously few
insect herbivores. Larvae of the cottongrass moth, Celaena haworthi Curtis,
tunnel in the culms of E. vaginatum in Europe, but no macrolepidopteran
species is reported to feed on Eriophorum species in North America (Tietz).
The aphid Rhopalosiphum eriophori (Walker) is reported on E. angustifolium
and E. vaginatum. The larvae of the beetle Plateumaris discolor (Panzer) live
in anaerobic conditions among the roots of E. vaginatum in Europe, obtaining
needed oxygen by tapping into the intercellular air spaces in the cortex of the
roots.
REFERENCES:
Under family references see BENTHAM; BiERO(GRN: Bi ASER (1941 a. 1941c); GONCHAR-
ov et al.: Hot T !M; LE MAOuIT & DECAISNE: LI RMAN & RAYNAL; LLOYD & WOOLHOUSE;
METCALFE; PATCH; RAYNAL (1972, 1973); TIETZ; and TORREY.
Under Scirpus see KOYAMA (1958) and S iHUYLER (1971b).
FAEGRI, K. Zur Hybridbildung in der Gattung Eriophorum. Verh. Inst. Ruibel Zurich
33: 50-58. 1958. [Hybridization of several European species; many illustrations.]
FERNALD, M. L. The North American species of Elriophorum. Rhodora 7: 81-92, 129-
136. 1905. [Eight species.]
FETCHER, N., & G. R. SHAVER. Growth and tillering patterns within tussocks of Erio-
phorum vaginatum. Holarct. Ecol. 5: 180-186. 1982.
& . Life histories of tillers of Elriophorun vaginatum in relation to tundra
disturbance. Jour. Ecol. 71: 131-147. 1983.
GOODMAN, G. T., & D. F. PERKINS. The role of mineral nutrients in Eriophorum
communities. Il1. Growth response to added inorganic elements in two E. vaginatum
381
TUCKER, CYPERACEAE
1987]


JOURNAL OF THE ARNOLD ARBORETUM
communities. Jour. Ecol. 56: 667-683. 1968; IV. Potassium supply as a limiting
factor in an E. r'uinatum community. Ibid. 685-696. 1968.
Hvi), H. A., & K. F. ADAMS. An atlas of airborne pollen grains. xvi 1 12 pp. London
and New York. 1958. [E. angiuslifbium Honck., 28, 29.]
RAYMOND, M. Two new Eriophorum hybrids from northeastern North America. Sv.
Bot. Tidskr. 45: 523. 1951. [E ' Pylaieanum (E. spissum x tE. russeolum), E. Porsildli
(E. ('hamissonis I E. spissum).]
What is Eriophoruon Chatissonis C. A. Meyer? Ibid. 48: 65. 1954.
RonIR rSON. K. P., & H. W. Wool Hiiousi. Studies of the seasonal course of carbon uptake
of Erioplhorum vaginatum in a moorland habitat. 11. The seasonal course of pho-
tosynthesis. Jour. Ecol. 73: 685-700. 1985.
WHN, R. W. Biological flora of the British Isles: Eriophnrum vaginatum L. Jour. Ecol.
61: 601-615. 1973.
-- & L. C. BIiss. Changes in arctic cottongrass tussock tundra communities. Arct.
Alp. Res. 6: 261-274. 1974.
3. Fuirena Rottboell, Descr. Icon. 70. 1772.
Rhizomatous perennials or caespitose annuals of sunny, wet, often disturbed
soils. Rhizomes horizontal, covered with persistent lanceolate scales, producing
cormlike axillary offshoots from which new culms arise. Culms erect or slightly
inclined, unbranched, terete, hollow. Leaves with sheaths tubular, costate, pu-
bescent, barely reaching to decidedly separated from the base of the next sheath,
the ligules hyaline, hispid (or glabrous) apically: basal leaves bladeless, cauline
leaves with blades lanceolate to linear, flat or slightly conduplicate [crescen-
tiform], pubescent (blades absent or reduced to an awned apex of the sheath
in I species); stomata paracytic: chlorenchyma not radiate. Inflorescences of 1
to several sessile or pedunculate glomerules in the axils of the upper leaves;
rays lacking or 1-4, smooth or hispidulous. Spikelets 1-6, ovoid to oblong.
Scales 30-60(-100 or more), ovate to oblong, widest at or above the middle,
hispid adaxially, less often glabrous or glabrescent, 3- to 9-nerved, the 3 central
nerves prolonged into a cuspidate, straight, or excurved apex '/ as long as to
nearly equaling the length of the body of the scale, the 3 basal scales sterile,
longer, narrower, and more conspicuously awned than the fertile ones. Flowers
perfect, protogynous. Perianth biseriate [uniseriate or absent], outer whorl (se-
pals) of 3 smooth or retrorsely scabrellate bristles, /4 to nearly as long as the
achene; inner whorl (petals) of bristles bearing expanded, entire [fimbriate],
hyaline to somewhat spongy blades with obtuse, acute, aristate, or emarginate
apices. Stamens 3 (infrequently 1, 2, or 6): filaments ribbonlike, about as long
as the subtending scale; anthers linear to ellipsoid; pollen grains uniaperturate,
obovoid to subspheroidal, psilate, trinucleate. Styles linear, frequently hispid:
stigmas 3. linear, about as long as the styles, pubescent. Achenes trigonous with
conspicuous ridged angles, ellipsoid, the apex acute but not apiculate, the base
stipitate (usually conspicuously so), the faces flat to slightly concave, delicately
striate or smooth [cancellate], glossy. Embryo fungiform. Base chromosome
number 23. (Including I 'aginaria Persoon.) LiEcroIYPE SPECIEi: t. unmbelata
Rottb.; see Britton & Brown, Illus. Fl. No. U. S. Canada, ed. 2. 1: 337. 1913.
(Named for Joergen Fuiren, 1581-1628, Danish physician.)
382
[VOL. 68


A warm-temperate and tropical genus of about 30 species. Seven occur in
the Southeast; these are well known through Kral's recent revision. An addi-
tional three occur in the southwestern United States. Fuirena repens Boeck. is
endemic to Mexico, while five primarily South American species extend north-
ward into Central America, Mexico, and the West Indies. About 12 species
occur in South America, and about as many in Africa. Only F. umbellata is
recorded in Europe, and it is limited to the southern part of the continent. Five
species occur in southern Asia, but none is recorded from the Soviet Union.
Most of our species are distributed from Texas to Florida along the Gulf
Coastal Plain and northward on the Atlantic Coastal Plain. Fuirena scirpoidea
Michx. and F. longa Chapman occur only as far north as southern Georgia,
F. breviseta Cov. as far as eastern Virginia, F. squarrosa Torrey north to Long
Island, and F. pumila to Cape Cod. The last species is disjunct in southern
Michigan and northern Indiana. Two others in our area, F. Bushii Kral and
F. simplex Vahl, are southern Great Plains species that occur eastward to
Louisiana, Arkansas, and Missouri.
All of the southeastern species have haploid chromosome numbers of 23.
The only exception is Fuirena simplex, for which n = 15 has been reported
from Texas populations, in addition to n = 23 from southeastern representa-
tives (Kral).
Plants of Fuirena have no reported economic significance in North America.
although F. glomerata Lam. and F. umbellata have been reported as important
weeds in Borneo, India. Taiwan, and Malaysia (Holm et al.).
REFERENCES:
Under family references see BEALI BENTHAM: BLASER (1940. 1941 a); CLARKE (1908.
1909); COOK; EYLES & ROBERTSON; FASSETI; GODFREY & WOOTEN: HESLA e' al.; HOLM
et al.: HOLTTUM; HUANG: J. HUTCHINSON: J. H. KERN; KUNTH: LE MAOUT & DECAISNE:
METCALFE: NAPPER (1965): NEES VON ESENBEWCK; O'NEILL; SC HULZE-MOTEL (1959. 1964);
STANDLEY; TORREY; and VAN DER VEKEN.
Under Scirpus see KOYAMA (1958).
BUSH, B. F. The North American species of Fuirena. Rep. Missouri Bot. Gard. 16: 87-
99. 1905. [Eight species: keys, descriptions, specimen citations.]
COVILLE, F. V. Revision of the United States species of the genus Fuirena. Bull. Torrey
Bot. Club 27: 1-14. 1890. [Four species.]
FORBES, P. L. Studies in Cyperaceae of southern Africa: VI. A new combination in
Fuirena with notes on the species. Jour. S. Afr. Bot. 35: 83-98. 1969. [F. hirsuta
(Berg.) Forbes; good illustrations of inflorescences, perianth parts, achenes.]
Scanning electron microscopy of the leaf blade epidermis of Fuirena Rottb.
(Cyperaceae). Proc. Electron Microscop. Soc. S. Afr. 3: 27, 28. 1973. [Adaxial epi-
dermis and substomatal chambers showing interspecific differences.]
Studies in Cyperaceae in southern Africa: 11. A new species of Fuirena Rottb.
S. Afr. Jour. Bot. 3: 359-362. 1984. [F. tennis, from eastern Cape Province and
Lesotho; illustrations.]
_ & C. M. LALKHAN. A preliminary study of silicon distribution in the leaf blade
epidermis of Fuirena coerulescens (Cyperaceae). Proc. Electron Microscop. Soc. S.
Afr. 13: 79, 80. 1983.
GOVINDARAJALU, E. The systematic anatomy of South Indian Cyperaceae: Fuirena Rottb.
383
TUCKER, CYPERACEAE
1987]


JOURNAL OF THE ARNOLD ARBORETUM
Hot. Jour. Linn. Soc. 62: 27-40. 1969. [' uncinata Kunth. / Ii allichiana Kunth,
F. puheve'ns Kunth, F. ciliaris (L.) Roxb.. and F. umbellata Rottb.: cross sections
of leaves and culms: species distinguishable by surface features of leaf blades: key.]
Hot M. T. Studies in the Cyperaceae. V. luirena squarrosa Michx. and F. scirpoihdea
Vahl. Am. Jour. Sci. 154: 13-26. 1897. [Morphological and anatomical study of
two species of the Southeast: illustrations.]
KRAI , R. A synopsis of Fuirena (Cyperaceae) for the Americas north of South America.
Sida 7: 309-354. 1978. [Keys, descriptions, illustrations, chromosome counts.]
4. Eleocharis R. Brown, Prodr. 224. 1810.
Small to medium-sized, loosely to densely caespitose or single-stemmed,
rhizomatous or stoloniferous. submersed, emergent, or littoral perennials (rare-
ly annuals) of marshes, ditches, and pond and river shores. Roots fibrous:
rhizomes (lacking in some species) slender. horizontal, covered with appressed
ovate to lanceolate scales. Culms terete or ellipsoid (less often trigonous, quad-
rangular, or flattened), solid or hollow (sometimes with thin transverse par-
enchymatous septa), smooth, with numerous paracytic stomata (submersed
lower portions of culms with few or no stomata): in submersed species sec-
ondary branches present, very closely spaced and seemingly verticillate. Leaves
1-4: sheaths closely fitting the base of the culm, the summit firm or scarious
(sometimes apiculate): blades lacking. Inflorescences single spikelets terminat-
ing the culms. Spikelets slenderly cylindrical to ovoid, slightly less than to
about 3 times thicker than the summit of the culm. Scales (2-)20-100, oblong.
lanceolate, obovate, or orbiculate. hyaline, firm, or coriaceous, strongly to
weakly nerved or nerveless, deciduous or persistent. Flowers perfect. Perianth
bristles (3-)6(-12) or absent, extrorsely or retrorsely barbed or smooth, per-
sistent on the base of the mature achene or falling from it. Stamens 3: filaments
hyaline, about equaling to shorter than the subtending scale: anthers ellipsoid
to linear: pollen grains 1- [to 4-]apertlurate. obovoid to subspheroidal. psilate
(scabrellate), trinucleate. Styles with swollen, bulbous base: stigmas 2 or 3,
capillary. Achenes lenticular or trigonous, ovoid, obovoid, or ellipsoid, the base
broadly rounded, the apex capped by a small to large, pyramidal, conical, or
swollen tubercle. the surface smooth or variously reticulate, dull, frequently
glossy, or iridescent. Embryos turbinate to fungiform. Base chromosome num-
ber 5. TYPE SPE( IES: E. palustris (L.) Roemer & Schultes (Scirpus palustris L.):
see Britton & Brown, Illus. Fl. No. U. S. Canada, ed. 2. 1: 310. 1913. (Name
from Greek, helos, marsh, and charis, grace, from the paludal habitat of most
species.)-SPIKE-RUSH, I)OG'S-HAIR GRASS.
A genus of about 250 species, worldwide in distribution. Eleocharis is evi-
dently closely related to Scirpus but is distinguished by its leafless culms and
its single, erect, terminal spikelets. Although the apical tubercles of the achenes
of E/leocharis are similar to those of some species of Fimbristlis Vahl, sug-
gesting that Eleocharis is most closely related to that genus (Svenson, 1929),
recent evidence supports a closer relationship between Scirpus and E/cocharis.
Both of these genera have non-kranz anatomy, while inmbhrisitlis has kranz
anatomy (Metcalfe). The embryos of l:/eocharis (turbinate to fungiform, radicle
basal, coleoptile lateral) are similar to those of species in Scirpus sect.
384
[voL. 68


TUCKER, CYPERACEAE
BOLBOSCHOENUS (Van der Veken), rather than to those ofFimbristylis (turbinate
to fungiform, radicle lateral, coleoptile basal).
Some 40 species occur in the Southeast, and many of these have rather wide
ranges. Holarctic, neotropical, and pantropic groups are represented in our
area. Svenson's (1929, 1957) division of the genus into seven series has received
wide acceptance, and our species are presented here according to his classifi-
cation. The two largest are ser. ELEOCHARIS (ser. Palustriformes Svenson) and
ser. TENUISSIMAE Svenson, having 13 and ten species in our area, respectively.
Plants of ser. ELEOCHARIS are characterized by slender culms and a stolon-
iferous habit; there are both tristigmatic and distigmatic species. Our repre-
sentatives are mostly northeastern species that occur southward only as far as
Virginia, Tennessee, or Arkansas. However, Eleocharis falax Weatherby, 2n =
42, and E. arenicola Torrey, 2n = 20, both of the Coastal Plain, are found in
most of the Southeastern States. Eleocharis montevidensis Kunth, 2n = 10, 20,
is a neotropical species that has been found north to the Carolinas and Cali-
fornia; it is sometimes treated as conspecific with E. arenicola.
Plants of ser. TENUISSIMAE are loosely caespitose and have slender, wiry
culms. Our species are mostly restricted to the Coastal Plain. In the Southeast
the neotropical Eleocharis nana Kunth has been found only in southern Florida,
while E. nodulosa (Roth) Schultes occurs along the Gulf Coast from Florida
to Louisiana. The most widely distributed of our species, E. tuberculosa (Michx.)
Roemer & Schultes, 2n = 30, is found throughout the Southeast northward to
Nova Scotia. It is distinctive in having perhaps the largest tubercle in any
species of the genus-as large as the body of the mature achene.
Plants of ser. MUTATAE Svenson are the tallest in the genus; three of our
species regularly reach 1 m. The plants are characterized by spikelets that are
barely wider than the apices of the subtending culm and that have persistent
scales. The plants are unusual ecologically because they grow in ponds or pools
with a stable water level. Most other species of the genus grow where receding
water levels leave the plants exposed in summer. Species of ser. MUTATAE have
very high chromosome numbers. Briggs has made counts for the Australian
Eleocharis equisetina Presl, 2n = 172, and E. sphacelaia R. Br., 2n = 94-100,
140, 180, 188. Six species of the series occur in our area: E. equisetoides (Ell.)
Torrey and E. quadrangulata (Michx.) Roemer & Schultes are reported
throughout the Southeast and range north to southern New England; E. cellulosa
Torrey, E. interstincta (Vahl) Roemer & Schultes, and E. elongata Chapman
are restricted to the Coastal Plain; and E. Robbinsii Torrey, a species mainly
of the Northeast, ranges south to Virginia and northern Florida along the
Coastal Plain. Tubers of E. dulcis (Burman f.) Trin. ex Henschel, n = ca. 100,
provide the familiar water chestnut of Oriental cuisine. The juice of the tubers
is strongly antibiotic (Hegnauer). The species is closely related to the eastern
North American E. equisetoides, and the pair serve as an example of the eastern
Asian-eastern North American pattern of disjunction (Wood).
Species of ser. PAUCIFLORAE Svenson are tiny plants with few-flowered spike-
lets. Eleocharis parvula (Roemer & Schultes) Link, 2n = 8, 10, E. rostellata
Torrey, and E. melanocarpa Torrey occur in the Southeast, and all have broad
ranges in our area.
385
1987]


386 JOURNAL OF THE ARNOLD ARBORETUM [vot. 68
i ,
_ ,t ,ilom I, -< "
Fl-it RE 2. Il rcharis. a-k. 1:. ccllulosua: a. habit ofstoloniferous plants. /4; b, cross
section ofculm. showing air spaces (black) with cross partitions (stippled-cellular detail
too small to be shown). x 10; c, detail of culm with apex of bladeless sheath, x 2; d,
spike of flowers in carpellate phase (flowers protogs nous), styles protruding (note flowers
with either 2 or 3 stigmas), x 2: e, abaxial side of flower, stigmas receptive, filaments


TUCKER, CYPERACEAE
Plants of ser. ACICULARES Svenson are also small. Three species occur in our
area, and their contrasting distribution patterns are notable. The northern
Eleocharis Wolfii Gray is found southward to Tennessee and Louisiana, while
the neotropical E. radicans (Poiret) Kunth ranges northward to Virginia and
Oklahoma. However, E. acicularis (L.) Roemer & Schultes, 2n = 20, 30-38,
50-58, a widespread north-temperate species, is reported from throughout
eastern North America. Both emergent and submersed growth forms of E.
acicularis have been described. Submersed plants have three large lacunae per
culm, while emergent plants have about ten small ones. These forms are ge-
netically identical and fully interconvertible, as is demonstrated by reciprocal
transplants (Rothrock & Wagner). The plants are able to grow in acidic runoff
from Appalachian coal mines and flourish in streams with pH as low as 2.8.
This is odd and suggests some overlooked variability in the species, because
in northern Europe it nearly always occurs in basic waters (Iversen).
The plants of ser. OVATAE Svenson have broadly ellipsoid to ovoid spikelets.
Three species are in our area: Eleocharis obtusa (Willd.) Schultes, 2n = 10, in
every Southeastern State, is one of the commonest spike-rushes in eastern
North America; the closely related E. Engelmannii Steudel, 2n = 10, occurs
from Georgia and Missouri south to the Gulf Coast; and E. lanceolata Fern.
is a southwestern species that just extends into our area in Arkansas and
Louisiana.
Plants of ser. MACU LOSAE Svenson are characterized by dark purple to black,
biconvex achenes. Some species grow submersed, while others are found in
littoral habitats. There are four species in our area: Eleocharis caribbaea (Rottb.)
Blake is pantropic (northward to South Carolina and Texas); E. olivacea Torrey,
2n = 20, is endemic to the Coastal Plain from Virginia to Florida; E. atro-
purpurea (Retz.) Kunth is widely but sporadically distributed in the Southeast
(but otherwise is found throughout temperate and tropical regions of both the
Old and New Worlds); and E. llavescens (Poiret) Urban is neotropical, growing
north along the Coastal Plain to Delaware.
Plants of ser. WEBSTERIA (S. H. Wright) G. Tucker' are submersed, flaccid,
6Ek/ocharis ser. WEIISTERIA. comb. nov.. based on 11t 'eh irla S. H. Wright. Bull. Torrey Bot. Club
14: 135. 1887.
of stamens not yet elongated. 4 of 6 perianth bristles visible, x 10: f, apex of spikelet,
carpellate phase past, lower flowers with protruding stamens, x 5; g, flower in staminate
phase with apex of subtending scale, x 10; h. abaxial view of flower in staminate phase
(note 4 of 6 perianth bristles, ovary with enlarged stylar base), x 10; i, adaxial view of
mature achene with persistent stylar base and smooth perianth bristles, achene lenticular,
x 12; j, mature achene, abaxial view, x 12; k, longitudinal cross section of achene
(tubercle, pericarp, seed coat, and basal embryo unshaded, endosperm stippled), x 12.
1-n, E. obtusa: 1, spikelet with mature achenes (hidden by subtending scales, few stigmas
visible at upper left), x 5: m, abaxial side of mature achene crowned by tubercle (persistent
style base) and with perianth bristles. x 12: n, detail of perianth bristle to show retrorse
barbs, x 25. o, p, E. tuberculosa: o, abaxial side of mature achene (trigonous in cross
section) with tubercle, x 12; p. detail of perianth bristle, x 25. q, E. atropurpurea: abaxial
side of mature achene (lenticular in cross section) with tubercle, perianth bristles absent,
x 25.
387
1987]


JOURNAL OF THE ARNOLD ARBORETUM
slender-branched plants of shallow, still waters. There are one or perhaps two
pantropic species (Eiten, 1976b). Eleocharis conlervoides is an uncommon
plant of cypress swamps and lakes in Florida. southern Alabama, southern
Georgia, and Louisiana. It is also known from widely scattered localities in the
neotropics and in tropical Africa and Sri Lanka. The species has been variously
placed in Rhynchospora Vahl (Bentham; Kiikenthal, 1948). Scirpus, and the
monotypic tI ebsteria. The slender, leafless culms are similar to those of other
species of Eleocharis. Additional submersed one-flowered species of Eleocharis
occur in Brazil and Africa (Nelmes). At anthesis, the one-flowered spikelets of
E. conaervoides are exserted just above the water surface. The achenes lack the
differentiated tubercle of most species of 'Eeocharis, but the embryos are typical
of the genus (Van der Veken).
Rikli reported that the inner parenchymatous layer was absent from the
bundle sheaths in many species of Elcocharis, a feature on which he based the
segregate genus C'hlorocharis. Metcalfe could not confirm this in any species
including those investigated by Rikli but suggested that further study might be
profitable.
Chromosomes of Eleocharis have been extensively studied. Cytologically.
the genus is the best known in the Cyperaceae. Most species have the diffuse
centric condition typical of the family; some have holocentric chromosomes
(Battaglia). Although aneuploidy has been frequent in most other genera of
Cyperaceae, polyploidy has been important in the evolution of this genus.
Several species have tetraploid (and sometimes hexaploid) races or subspecies.
Strandhede (1965, 1966) studied about 1100 European populations of species
of ser. ELEOCHARIS (ser. Palustrifornies) and reported that chromosome break-
age and refusion were common. Most species had several cytotypes, and various
kinds of multivalents were frequent at meiosis. Heterovalents formed in mei-
osis. and aberrant but apparently viable gametes were often observed. Similar
reports of chromosomal variability have been made for North American species.
Karyotypic rearrangements have been noted in Elecocharis flavescens (Poiret)
Lam., which had 30 chromosomes in various combinations of univalents,
bivalents, tetravalents, and ring complexes (Schuyler, 1977).
When the sample size is large, chromosome number can be correlated with
morphology within species and between species pairs. For example, the Eu-
ropean Elcocharis unigl/umis (Link) Schultes consists of two subspecies that
differ in ecology and in features of the spikelet scales. Subspecies unig/umnis
has n = 46, while subsp. Sterm'ri Strandhede has n = 74-82. Apparently. the
latter taxon was derived from the former by tetraploidy followed by fusion of
some of the chromosomes, but fusion of different chromosomes in different
populations has also resulted in mixoploidy. In some cases affinities between
species can be confirmed cytologically. For example, E. Engelmannii Steudel
and E. obtusa are both n = 5 and have very similar karyotypes.
Species with different chromosome numbers are known to hybridize in the
wild. Some hybrids (e.g.. Eleocharis manmillata x E. palustris subsp. palustris)
:lh c( pri nr OthfrvIde (PoIiret) (i. Tucker. cnmb. no\.. based on S c'ir onnervuicn s Poiret in
ILam. I ncwl. Meth. Hlot. 6: 755. I804.
388
[VOL. 68


TUCKER, CYPERACEAE
have greatly reduced fertility, while others (e.g., E. palustris subsp. palustris x
subsp. vulgaris) have fertility comparable to that of the parent species.
Several species are important weeds, especially of rice fields.
REFERENCES:
Under family references see BARROS (1928); BA TAGLIA; BEAL; BENTHAM; BERGGREN;
BLASER (1940, 1941a); BREWBAKER: CI'ARKE (1908, 1909); EITEN (1976a, 1976b); EYLES
& ROBERTSON; FASSETT: GODFREY & WOOTEN: GONCHAROV el at.; GOOD et al.; HA-
KANSSON (1954. 1958): HARBORNE 1e al.: HARRIS & MARSHALl; HEGNAUER; HESLA et
al.; HEUSSER; HOLM et al.; HOLTTUM: HOTCHKISS: HUANG; G. E. HUTCHINSON,
J. HUTCHINSON; F. D. KERN; J. H. KERN: KUNTH: LE MAOUT & DECAISNE; MCATEE;
METCALFE; NAPPER (1965): NEES VON ESENBECK; OGDEN; O'NEILL; PATCH: RIKLI;
SCHULZE-MOTEL (1959, 1964): SMITH et al.: STACE: STANDLEY; TIETZ; TORREY; and VAN
DER VEKEN.
BERNARDINI, J. V. Studies of the kinetochore of Eleocharis macrostachra Britt. Proc.
Minnesota Acad. Sci. 27: 104-114. 1959.
BOYD, C. E., & D. H. VICKERS. Relationships between production, nutrient accumu-
lation, and chlorophyll synthesis in an Eleocharis quadrangulata population. Canad.
Jour. Bot. 49: 883-888. 197 1.
BRIGGS, B. G. Chromosome numbers in some Australian species of Eleocharis (Cy-
peraceac). Contr. Natl. Herb. New South Wales 4: 130-136. 1970. [Summary of all
reported chromosome numbers arranged in series: E. acicularis (L.) Roemer &
Schultes, 2n = 20.]
BRONNER, G. Aquarium plants. (English translation by G. VEVERS.) [vii] - 94 pp.
Princeton and New York. 1966. [E. acicularis, E. vivipara Link, 23, 24, propagated
by division and runners.]
EVANS, P. S. Intercalary growth in the aerial shoot of Eleocharis acula R. Br. I. Structure
of the growing zone. Ann. Bot. 79: 205-217. 1965.
HARMS, L. J. Cytotaxonomic studies in Eleocharis subser. Palustres: central United
States taxa. Am. Jour. Bot. 55: 966-974. 1968.
Cytotaxonomy of the Eleocharis tennis complex. Ibid. 59: 483-487. 1972.
HORN AF RANTZIEN, H. Certain aquatic plants collected by Dr. J. T. Baldwin Jr. in
Liberia and the Gold Coast. Bot. Not. 1951: 384-398. 1952. [Subg. RHEOCHARIS
Horn described and illustrated.]
IVERSEN, J. Studien liber die pH-Verhaltnisse dinische Gewaisser und ihren Einfluss auf
die Hydrophyten-Vegetation. Bot. Tidsskr. 40: 277-326. 1929. [Distribution of E.
acicularis f. submerva: 95 percent of its occurrences in Denmark are in neutral,
alkaline, or variable waters, only 5 percent in acidic waters: cf. ROITlRO( K & WAGNER.]
KOKENTHAL, G. Vorarbeiten zu einer Monographie der Rhynchosporoideae. Rhyn-
chospora. Bot. Jahrb. 74: 375-509. 1949: Ibid. 75: 90-195. 1950: Ibid. 75: 273-
314. 1951.
LEWIS, K. R., & B. JOHN. Hybridisation in a wild population of Eleocharis palustris.
Chromosoma 12: 433-468. 1961.
NELMES, E. Submersed species of Eleocharis with 1-flowered spikelets. Kew Bull. 1952:
289, 290. 1952. [E. Naumanniana Bock. and E. Callei Hutchinson & Dalz. of
Africa.]
POGAN, E. Studies in Eleocharis R. Br. I. Chromosome numbers of E. palustris (L.) R.
et S. and E. uniglumis (Link) Schult. Acta Biol. C('racov. Bot. 15: 69-76. 1972. [E.
palustris subsp. palustris, 2n = 16: E. palustris subsp. vulgaris Walters, 2n = 38,
39, 40; E. uniglumis, 2n = 46.]
REJMANEK, M., & J. VELAZQUEZ. Communities of emerged fishpond shores and bottoms.
Pp. 206-211 in D. DYKYJOVA & J. KVET, eds., Pond littoral ecosystems. (Ecological
1987]


JOURNAL OF THE ARNOLD ARBORETUM
studies 28.) New York. 1978. [Maximum standing crop of E. acicularis is 469-644
gnm .]
RoruRO( K, P. E., & R. H. WAGNER. The autecology of an acid tolerant sedge. Eleocharis
acicularis (L.) R. & S. Castanea 40: 279-290. 1976.
SC HYULER, A. E. Chromosome observations on some eastern North American Eleocharis
(C'yperaceae). Brittonia 29: 129-133. 1977.
& W. R. FERREN, JR. A new intertidal form of Eleocharis olivacea (Cyperaceae).
Bartonia 43: 46-48. 1975.
Si sc iAn, F. K., J. M. BERNARD. & K. FIA.A. Above- and belowground standing crop
partitioning of biomass by Elocharis rostellata in the Byron-Bergen Swamp, Ge-
nesee County, New York. Am. Midi. Nat. 114: 70-76. 1985.
SriANDHnie, S. O. Chromosome studies in Eleocharis subser. Palustres. III. Obser-
vations on western European taxa. Op. Bot. 9(2): 1-86. 1965.
L leocharis subser. Palu.irs in North America, taxonomical comments and
chromosome numbers. Bot. Not. 120: 355-368. 1967.
SViNSON, H. K. Monographic studies in the genus EIcocharis. Rhodora 31: 121-135,
152-163, 167-191, 199-219, 224-242. 1929. [The basic monograph, worldwide:
keys. descriptions, distribution maps, synonymies.]
Monographic studies in the genus Ehleocharis-II. Ibid. 34: 193-203, 215-227.
pl. 221. 1932; II111. Ibid. 35: 377-389. pls. 320, 321. 1935; IV. Ibid. 39: 210-273.
p/s. 460-465. 1937: V. Ibid. 41: 1-77, 93-1 10. p/s. .537-547.
The group of Eleocharis palustris in North America. Rhodora 49: 61-67. 1947.
. locharis. N. Am. Fl. 18: 509-540. 1957. [Keys, descriptions.]
WALTERS, S. M. Eleocharis. In: Biological flora of the British Isles. Jour. Ecol. 37: 192-
206. 1949.
On the vegetative morphology of Eleocharis R. Br. New Phytol. 49: 1-7. 1950.
WARD, D. B.. & E. M. H. LiGii. Contributions to the flora of Florida-8, E/cocharis
(Cyperaceae). Castanea 40: 16-36. 1976.
Wool), C. E., JR. Morphology and phylogeography: the classical approach to the study
of disjunctions. Ann. Missouri Bot. Gard. 59: 107-124. 1972. [Eastern North Amer-
ican-Eastern Asian disjunctions: Du/lichiumn arundinaccum, extant and fossil distri-
bution, 118.]
WRiciH, S. 11. A new genus in Cyperaceac. Bull. Torrey Hot. Club 14: 1887. [ WelCbsera.]
5. Fimbristylis Vahl, Enum. P1. 2: 285. 1805.
Small to medium-sized annuals or perennials of'disturbed, open. wet habitats.
Roots fibrous; rhizomes regularly present in some species. Culms slender, terete
or nearly so, glabrous. Leaves all basal; sheaths smooth or pubescent, with
ligule present or not, glabrous or ciliate: blades linear to filiform, flat, condu-
plicate, or involute, glabrous or pubescent, the margins glabrous or scabrellate;
chlorenchyma radiate; bundle sheaths 3-layered ("Fimbristylis type"). Inflo-
rescences terminal, branched (rarely sessile, capitate); bracts 1-6, erect to oblique,
the sheaths greatly reduced to essentially absent, the blades leaflike; primary
rays absent or 1-10, glabrous or scabrellate, secondary rays regularly produced
in some species. Spikelets single or in clusters of 2-5, ovoid to lanceolate. Scales
5-100, ovate to oblong, obtuse or acute, blunt or mucronate [aristate], glabrous
or puberulent abaxially, 1- to 5-nerved medially. nerveless laterally, deciduous
at maturity. Flowers perfect. Perianth lacking. Stamens (1, 2, or) 3; filaments
about as long as the subtending scales, flattened; anthers oblong, the apices of
the connectives sometimes prolonged; pollen grains uniaperturate, obovoid,
390
[voL. 68


TUCKER, CYPERACEAE
subspheroidal, or spheroidal. scabrate, trinucleate. Styles slender, terete
throughout or trigonous basally, usually fimbriate distally, deciduous from the
mature achene; stigmas 2 (or 3), about as long as the style, glabrous. Achenes
lenticular or trigonous, ovoid, oblong, or obovoid, the apex broadly rounded
to subacute, apiculate or not, the base cuneate or stipitate, the surface smooth,
warty, or reticulate with isodiametric or horizontally arranged rectangular cells,
these cells concave or with a central papilla. Embryos turbinate, radicle lateral,
coleoptile basal. Base chromosome number 5. TYPE SPECIES: F. dichotoma (L.)
Vahl, typ. cons. (Name from Latin timbria, fringe, and stylus, style, referring
to the fringed style of most species.)
A genus of about 200 species, mainly pantropic but also well represented in
warm-temperate regions. Most of the species grow in disturbed wet habitats,
especially roadsides and croplands. The center of diversity is southeastern Asia
(Goetghebeur & Coudijzer). Thirteen species are recorded from the United
States. Twelve of these occur in the Southeast, while Fimbristylis thermalis S.
Watson is endemic to California, Arizona, and Nevada (Kral). Kral's thorough
monograph includes illustrations and chromosome counts for all species in
North America.
Fimbristylis is closely related to Bul/ostylis and Abildgaardia. Chromosome
numbers in the three genera are based on five (Gordon-Gray, Kral), and their
kranz anatomy is similar (three-layered bundle sheaths). Such anatomy is not
reported in any other genera of the Cyperaceae (Metcalfe: Raynal, 1972). The
three genera have been distinguished from the remainder of the Scirpeae as
tribe Abildgaardiae Lye (Fimbristylideae Raynal).
Koyama (1961) treated Bulbostylis as a subgenus of Fimbristylis, while Kral
recognized three genera, Bulbostiylis, Abildoaardia, and Fimbristylis. Additional
information supports Kral's belief. Gordon-Gray made a careful study of the
southern African representatives of the three genera. Abildgaardia can be dis-
tinguished from Bulbostylis and Fimbristylis by its distichous spikelet scales.
Bulbostylis and Fimbristylis are separated by a suite of characters. The embryos
are consistently different (in Fimbristylis the radicle is lateral, the coleoptile
basal; in Bulbostylis, the radicle is basal and the coleoptile lateral), although
there is no single morphological character that separates the two genera. The
styles of Fimbristylis are usually fimbriate (occasionally entire) and are decid-
uous, while those of Bulbostylis are always entire and have a persistent base.
The spikelet scales of Fimbristylis are generally glabrous, while those of Bul-
bostylis are generally puberulent. The ligules of Fimbristylis are glabrous, while
those of Bulhostylis are hispid. Species of Fimbristylis always lack intrapro-
phyllar buds at the base of the inflorescence rays, while such buds are frequently
present in Bulbostylis (Guaglianone). Species of the two genera differ in surface
ornamentation of the achenes. Goetghebeur & Coudijzer examined about 100
species from throughout the world and found that the epidermal cells of Fim-
bristylis are horizontally elongate (infrequently isodiametric) and in vertical
bands, but those of Bulbostylis are vertically elongate in horizontal bands. The
two genera also differ in habit and habitat: Fimbristylis species are mostly
perennials of moist soils, while Bulbostylis species are generally annuals of dry
sandy soils.
1987]


392 JOURNAL OF THE ARNOLD ARBORETUM [VOL. 68
Svenson recognized two sections in Finihristylis: Kral did not comment on
the infrageneric classification. Plants of sect. FIMBRISTiLIS (sect. Dichelostvlis
Bentham) have two stigmas, lenticular achenes, and styles commonly fringed
apically. This section includes eleven of the fourteen species of the southeastern
United States. Most of our species are somewhat weedy plants of disturbed
wet habitats: f'imhristylis tomentosa Vahl, n = 5: t1" dichotoma (L.) Vahl, n =
10. 15: F'. decipicns Kral. n = 10: F. annua (All.) Roemer & Schultes, n = 15:
F1. 'ahlii (Lam.) Link. n 10: F. puberula (Michx.) Vahl, n 10, 20: and !.
pcerpusilla Harper, n = 5. In general these are widely distributed in the Southeast.
Fimhristvlis perpusill/a, endemic to southeastern North America, is a notable
exception. Kral knew of only two localities in southwestern Georgia for this
tiny annual. Recently, the species has been reported in Horry County. South
Carolina (tLeonard) and in eastern Maryland (Schuyler. pers. comm.). The four
remaining southeastern species of sect. FIMnRISi tis. F. caroliniana (Lam.)
Fern. (n = 10. 20, 30). F. schoenoides (Retz.) Vahl (i = 5), F. spathacea Roth
(n r 24). and F. castanea (Michx.) Vahl (n 10). are tall plants of tidal marshes.
Plants of sect. TRICHELOSTYLIS Bentham have three stigmas, lenticular achenes,
and entire styles. In our area this section is represented by Fihmbristylis aurum-
nalis (L.) Roemer & Schultes, F. compl/alnaa (Retz.) Link, and F. miliacea (L.)
Vahl, all n = 5.
'imahristtr/is autttunadi and t1 iniliacea are detrimental weeds in rice fields
in the Southeast and California (Smith ct al.), as well as in Asia and Africa
(Holm ce al.). Fimnhristv/is toicntosa is rapidly becoming a common weed in
rice fields from South Carolina to Texas (Kral).
REFERENCES:
Under lamily references see BAive N c l.; BARROS (1945): Bi \i; BEN i HAM; BREWBAKER:
BlRoWN; (AROIIN 'et al.: (Cl RK (1 9()8, 1909): FssI i Is (iOI)i RiNE & WOOTiEN: GONCHA-
ROV 't al.: HARiORNF: HARBORNI ct al.: Hoi M ct al.: Hoit iti M: HUANo: J. H�I'II INSON:
.1. H. KIRN: KOYAMA (1961): KiKKONI N (1969): Ki N IIi Li MaOI & D: .AISNi; LFR-
MAN & RA\NAI ; MT( Afir: NAPIiR (1965): Ni.i s vON l.SEiNiBK(K: O'NeILIi; PATCH: RAYNAI
(1972, 1973. 1978): RIK I: S< 1t ii -Moii EI (1959, 1964): SMUiI i et al.: STANDI Fv: TEiRi
t a(! TORRI EY: and V \N Di VI KIN.
(101 iHirnHFtR, P., & .1. Co Iti iI/-R. Studies in ( 'peraccae 3. Flimbhristi/s and .bhild-
uaardia in ( entral Africa. Bull. .lard. Bot. Nail. Belg. 54: 65-89. 1984. [SEM pho-
tographs of achenes.]
(ioRi)N-(iRAy, K. 1). F/ nibrist./ and Hui!/ .st'li.s: generic limits as seen by a student
of southern African species. Mitt. Bot. Staalssam. Muinchen 10: 549-574. 1971.
GUAGLIANONI. E. R. tUn nuevo earacler, util en la distincion generica entre Fimbristylis
Vahl s Bulhostvi's Kunth ((Cyperaceae). D)arwxiniana 16: 40-48. 1970.
1101 Mt. T. Studies in the Cyperaceae. X. 'i7hubrstiv/is Vahl: an anatomical treatise of
North American species. Am. Jour. Sci. 157: 435-450. 1899.
KRi. R. A treatment of .lhild,,aardia, Biulhnvstylis, and Fiinhrisitylis (Cyperaceae) for
North America. Sida 4: 57-227. 1971.
ION ARI, S. W. FI'imrihnstvhi pcrp)iiilla Harper in South Carolina. Castanea 46: 235.
236. 1981.
SviNSON, H. K. l"inibri.v/wi is. iihUMbsi.is, and Ahblduaardia (C(peraceae: Scirpeae). N.
Am. l. 18: 540-556. 1957.


WARD, D. B. Contributions to a flora of Florida, 4. Fimbristylis (Cyperaceae). Castanea
33: 123-134. 1968a.
Supplemental note to Fimbristylis of Florida. Ibid. 350. 1968b.
6. Bulbostylis Kunth ex C. B. Clarke in Hooker f. Fl. Brit. India 6: 651. 1983,
nora. cons.
Small to medium-sized, tufted (solitary-stemmed) perennials or annuals of
open or disturbed, dry or wet habitats. Roots fibrous; rhizomes lacking [present].
Culms slender, terete, glabrous. Leaves all basal; sheaths expanded basally or
not, with ligule fimbriate or ciliate apically; blades filiform or narrowly linear,
shorter than to slightly exceeding the culm, conduplicate or involute, often
pubescent on one or both surfaces, the margins and midvein scabrellate or
smooth; chlorenchyma radiate: bundle sheaths 3-layered ("Finmbristylis type").
Inflorescences terminal, capitate or branched: bracts 1-4, erect to oblique,
shorter than to exceeding the length of the rays: primary rays lacking or 1-6,
erect or spreading, subterete, glabrous or scabrellate, secondary rays absent.
Spikelets solitary or in small clusters, ovoid to oblong or lanceolate. Scales 2-
50, ovate to oblong, mucronulate, mucronate, or aristate, glabrous or scabrel-
late, or puberulent abaxially, 3- to 7-nerved, deciduous at maturity, the 1-4
lowest ones sterile. Flowers perfect. Perianth lacking. Stamens (1, 2, or) 3;
filaments slender, hyaline, about as long as the subtending scales; anthers ob-
long, the apices of the connectives prolonged as tiny subulate tips; pollen grains
uniaperturate, subspheroidal or obovoid, psilate or scabrate, trinucleate. Styles
papillate, the bulbous basal portion persistent on the mature achene; stigmas
3, slender, glabrous, equaling to exceeding the style in length. Achenes trigonous
(rarely biconvex), ovoid to oblong or ellipsoid, the apex obtuse to acute, crowned
by the persistent bulbous style base, the base cuneate to stipitate, the surface
smooth or reticulate with vertically elongate, rectangular (rarely isodiametric)
cells, these cells smooth or sometimes with a single central papilla. Embryos
turbinate, radicle basal, coleoptile lateral. Base chromosome number 5. TYPE
SPECIES: B. capillaris (L.) C. B. Clarke, tip. cons. (Name from Latin bulbus,
bulbous, and stylus, style, referring to the characteristic bulbous style base.)
A genus of about 120 species, mostly pantropic but with some in the warm-
temperate regions. The genus is related to Abildgaardia and Fimbristylis. (A
discussion of the distinguishing features of these genera appears under Fi'm-
bristylis.) Bulbostylis was first distinguished from Fimbristylis as the genus
Stenophyllus Raf. (Neogenyton, 4. 1828). Although the generic name Bulbo-
stylis Kunth was published in synonymy (Kunth) and validated by Clarke (q.v.),
it has been conserved over Stenophyllus. Kral's illustrated monograph (in-
cluding chromosome numbers) is the basic reference for the North American
species.
Bulbostylis is represented in the United States by eight species, live of which
occur in the Southeast. Bulbostylis barbata (Rottb.) C. B. Clarke. n = 5, B.
capillaris (L.) C. B. Clarke. n = 36, and B. ciliati/olia (Ell.) Fern., n = 30, have
each been reported from all or nearly all the southeastern states. Bulbostylis
stenophylla (Ell.) C. B. Clarke and B. I'arei (Torrey) C. B. Clarke, both n =
TUCKER, CYPERACEAE
393
1987]


JOURNAL OF THE ARNOLD ARBORETUM
15, are more restricted in range than the three preceding species. Both occur
along the Coastal Plain from Florida to North Carolina. Three more species,
B. lFi(nckii (Steudel) C. B. Clarke. n =- 10 . juncoides (Vahl) Kukenthal, n =
60, and B. Schaf/neri (Boeck.) C. B. Clarke, occur in the Southwest. About 15
species occur in Mexico, Central America. and the West Indies, with perhaps
20 in all of South America. The center of diversity for the genus is tropical
Africa, where 30-40 species are reported.
The southeastern species of B1ulostylis are generally found in open. dry,
sandy places, such as pine flatwoods, sand hills, palmetto scrub, roadsides, and
shores. They are annuals or short-lived perennials. The neotropical B. paradoxa
(Sprengel) Lindm., a long-lived perennial that flowers in response to fires (Kral).
occurs in pinelands and savannas in Cuba and from Mexico to northern South
America.
Plants with basal clusters ofspikelets are occasionally encountered in several
species of tHulhotylis (e.g.. 1. Capillaris and B. F'unckii). Formation of such
spikelets may be the result of drought, but no studies have been made to
document this supposition. In some species achenes produced by the basal
spikelets are 1 '/-2 times larger than those produced by typical elongate culms.
Such amphicarpy has also been reported in certain African species (Haines).
lhlostlvlis harhuta is a weed of old fields and sandy croplands in the south-
eastern Coastal Plain. Three species (including B. harbata) are reported as
significant weeds in tropical Africa and Asia (Holm ct al.).
Rf i RI N( is:
U nder lamily references see ItAR{OS (1945); BLAI lt N IIAM: BRoWN: ( CAROl IN e at.:
(C \RKi (1908. 1909): (ioDrRi & \WOiiN;: (GON(HAROV Ct ul.: HAINsES; IHARIORNi.;
FHlRilORNI etal('/.: HO s M t ill.: Oia l i 11 i.: liH N(,.. l 1 I 'Hi[NS(i N; ,J. H. KERN: KiUKKONFN
(I 969): KtINTH: IT M i\ i & D) ) ISN : Li RM\'N & RA'NAx : MN i( AL hi: NAPPFR (1965):
NitS \VON FSFNII ('K: O'Ni I : RN\ sA (1972. 1973. 1978): RIKl I: S( i izF-MorFI (1959,
1964): lii\N rv: TIi RI ei at.: TORRI: V\N IOR VIKiN: and WINRFVt & -\MSisIL.
tlnder /1Ir tihvi7/ts see GoRI)oN-(GRA. KRAI , and Sv iNS(N.
C(ii nI itiBhi R, P. Studies in (xpcraccac 4. Newv species and a new combination in
( entral African li/lho.sti7Iis. Bull. Jard. Bot. Nail. Belg. 54: 91-104. 1984.
(GoviNiARAJAi i, E. The systematic anatomy of south Indian ( peraceae: BHushoi'ii.
Kunth. Jour. Linn. Soc. Bot. 59: 289-304. 1966.
I v>, K. A. The generic concept of B/ulovtiivi Kunth ex (C. B. ( 1. Mitt. Bot. Staatssam.
Miinchen 10: 539-547. 1971.
7. Abildgaardia Vahl, Enum. P1. 2: 296. 1805.
Small, single-stemmed or tufted, bulbous-based, glabrous perennials of trop-
ical and subtropical grasslands. Roots fibrous: rhizomes lacking. Culms sub-
terete, smooth. Leaves about / as long as the culms: sheaths expanded. their
overlapping bases forming the bulblike base of the plant, ligules lacking: blades
linear-tiliform, slightly involute, thickened at margins, scabrellate distally;
chlorenchyma radiate: bundle sheaths 3-layered ("1'imbristvlis type"). Inflo-
rescences simple cymes of 1-3[-6] sessile or pedunculate spikelets: bracts sol-
394
[VOL. 68


TUCKER, CYPERACEAE
itary, filiform. Spikelets broadly lanceolate, slightly compressed, the scales
distichous or essentially so. Scales 3-15, ovate, acute, mucronate, 3- to
5-nerved medially, nerveless laterally, deciduous as the achenes mature. Flow-
ers perfect (although frequently the distal flowers of a spikelet staminate only).
Perianth lacking. Stamens (1, 2, or) 3: filaments flattened; anthers linear, the
apices of the connectives not prolonged; pollen grains uniaperturate, obovoid
to subspheroidal, scabrate, trinucleate. Style trigonous basally, slender and
capillary distally, deciduous from the mature achene; stigmas 3, linear, about
as long as the style, glabrous. Achenes rounded-trigonous, ovoid, the apex
broadly rounded, apiculate, the base abruptly contracted to a stipe, the surface
pebbled. Embryo turbinate, radicle basal. Base chromosome number 10. (Named
for P. S. Abildgaard, an eighteenth-century Danish botanist.) TYPE SPECIES: A.
ovata (Burman f.) Kral (Carex ovata Burman f.; A. monostachya (L.) Vahl);
see Britton & Millspaugh, Bahama Fl. 52. 1920.
A pantropic genus of about 15 species, distinguished from Bulbostylis and
Fimbristylis, with which it has been united, by its distichous spikelet scales
and its deciduous style bases. Chemical data support the recognition of Abild-
gaardia. The four Australian species produce the flavones luteolin and tricin,
whereas the 15 species of Fimbristvlis and Bulbostylis examined had only tricin
(Harborne et al.).
Abildgaardia is represented in the New World by two species. Abildgaardia
mexicana (Palla) Kral, n = 10, is endemic to grasslands of the Mexican High
Plateau. The southeastern representative, A. ovata, n = 10, occurs in Florida,
the West Indies, and the lowlands of Central and South America. Abildgaardia
ovata is found in grasslands over limestone in southern Florida (Dade and
Monroe counties) and in the vicinity of Tampa (Citrus County; Kral).
Species of Abildgaardia have no reported economic significance. None has
been noted as a weed.
REFERENCES:
Under family references see BARROS (1945): BENTHAM: CLARKE (1908): GODFREY &
WOOTEN; HARBORNE: HARHORNE et al.: HUANG: KUNTH: LERMAN & RAYNAL: METCALFE:
NAPPER (1965): NEES VON ESENBIECK: O'NEILL: SCHUIZE-MOTEL (1959, 1964): and VAN
DER VEKEN.
Under Fimbristylis see KRAL and SVENSON.
LYE, K. A. Studies in African Cyperaceae VIII. The taxonomic position of Ahildgaardia
Vahl and Nemum Hamilton. Bot. Not. 126: 325-329. 1973.
Tribe CYPEREAE
8. Cyperus Linnaeus, Sp. Pl. 1: 44. 1753: Gen. P1. ed. 5. 27. 1754.
Tufted or rhizomatous, perennial or less often annual herbs of disturbed wet
to dry soils, marshes, ditches, shallow swamps, and shores in full sun or light
shade. Roots fibrous: rhizomes or stolons sometimes present, horizontal to
oblique. Culms trigonous (sometimes with winged angles) or terete, smooth or
scabrellate. Leaves all basal: sheaths glabrous, sometimes with conspicuous
1987]


JOURNAL OF THE ARNOLD ARBORETUM
cross veins, especially in emergent plants, ligule present or lacking; blades linear
to lanceolate, flat, conduplicate, plicate, filiform, crescentiform, or involute,
the margins and midvein usually scabrellate: stomata paracytic, sometimes
surrounded by 1-4 papillae; chlorenchyma radiate or not (if radiate the bundle
sheaths 2-layered-"(Cperus type"). Inflorescences terminal, diffusely branched,
spicate, or capitate: bracts (1-)3-6(-22), the sheaths very short, the blades
leaflike, closely spaced and appearing verticillate at the apex of the culm, usually
ascendent but in some species erect (the inflorescence thus appearing lateral),
horizontal, or reflexed. forming a conspicuous involucre: rays glabrous (rarely
scabrellate or hispidulous), unequal in length, produced singly from the axils
of the inflorescence bracts: spikes digitate, glomerulate, or spicate; rachis smooth,
rarely scabrellate. Spikelets (1-)5-30(-150), cylindrical to compressed, ovate,
lanceolate, or linear, the scales distichous: rachilla deciduous or persistent,
internodes winged or wingless, spongy and thickened in a few species. Scales
(1 or) 2-20(-80), oblong, elliptic, or ovate, obtuse, acute, mucronulate, or
cuspidate, 3- to 1 1-nerved, deciduous or persistent, the 2 lowermost (bract and
prophyll) sterile. Flowers perfect [imperfect, the plants dioecious]. Perianth
lacking. Stamens (1, 2, or) 3: filaments ribbonlike, usually as long as the sub-
tending scales: anthers ovoid, ellipsoid, or linear, the apices of the connectives
sometimes prolonged as small, reddish, entire or scabrellate appendages: pollen
grains obovoid, subspheroidal, rectangular, or triangular. (I- or) 4-aperturate,
psilate, trinucleate. Styles slender, the base sometimes persistent as an apiculus
or beak on the mature achene: stigmas capillary, shorter than, equaling, or
exceeding the style in length, glabrous [glandular]. Achenes trigonous or len-
ticular, ovoid, ellipsoid, or narrowly oblong, obtuse or acute, apiculate or not,
stipitate, substipitate. or sessile, smooth, puncticulate. or reticulate. Embryos
broadly to narrowly ellipsoid. Base chromosome number 8. (Incl. Pvtcreus
Beauv., Mariscus Vahl, Juncellus (Griseb.) C. B. Clarke, A.corellus Palla, Re-
mirea Aublet, Torulinium C. B. Clarke.) LECTOTYPE SPECIES: C. esculentus L.:
see Britton & Brown, Illus. Fl. No. U.S. Canada, ed. 2. 1: 297. 1913. (Name
from Greek kupeiros, ancient name for C. longus L.)-FFIAT-SEDGE, UMBREL-
LA-SEDIGE, SEDGE-GRASS, GALINGALE (Britain).
A very large genus of about 650 species widely distributed throughout the
tropical and warm- and cool-temperate regions of the world. It is the second
largest genus of the Cyperaceae: only Ca'ex L. is larger. Cy'iperus is morpho-
logically coherent and is readily recognized by the distichous arrangement of
scales on the spikelets. Six subgenera have been recognized: subg. C'YPERtUS,
subg. PYCNOSTACtHYS C. B. Clarke,- subg. PYCRE'S (Beauv.) Gray." subg.
JIJNCEIL US (Griseb.) Kiikenthal, subg. TORILINNitM (Desv.) Kukenthal, and
subg. FIMBRi(YPERUS K. A. Lye. These are circumscribed by features of the
achenes. spikelets, and vegetative anatomy. Most recent workers have followed
'( ip7 n subg. Pu'V NOSIa iH C. B. ( larkc in Hooker 1 1. BItil. India 6: 597. 1893. I.i .i it'i
SPcIies (here designated): C. Jitln \ s Vahl. Synonym: ( irprntu subhg. Protc l entrus K A e. Nordic
Jour Bot. 1: 54. 1981. YPt Si{Ctes: ( (hilfruirnis [..
" 17'r subhg. P'( RIs (B au . ) r( . Man. ot.ed. 1.517. 848 1hiscom imation is consistentl\.
libut erroneously, attributed to ('. I. Clarke. lour, I inn. Soe. Boit 21: 33. 1884.
396
[vol.. 68


TUCKER, CYPERACEAE
Kiukenthal and Fernald, who treated the genus in the broad sense. Others
(Koyama, 1962b; Vorster; Raynal, 1972, 1973) have followed Clarke (1908)
and recognized the subgenera as genera. Subgenera PYCREUS and JUNCELLUS
differ from the others in having the derived conditions of lenticular (vs. trig-
onous) achenes and bifid (vs. trifid) styles (Blaser, 1941a; Raynal, 1972). In
subg. PYCREUS the achenes are laterally compressed, while in subg. JUNCELLUS
the compression is dorsiventral, suggesting that the bicarpellate condition
evolved twice. Several other genera of the family (e.g., Carex and Bulbostylis)
are divided into subgenera on the basis of carpel number.
Subgenus TORULINIUM differs from all other subgenera in having the rachilla
articulate at the base of each scale (i.e., an abscission layer forms) (vs. contin-
uous or articulate only at the base of the spikelet). Thus, the mature spikelet
of plants of subg. TORULINIUM breaks up into one-fruited segments, each con-
sisting of an internode of the rachilla, a scale, and an achene.
Subgenera JUNCELLUS, PYCREUS, and TORULINIUM are readily distinguished
from each other and from the remaining subgenera. However, the subgeneric
classification of the remaining species of the genus has been a matter of long
debate. Traditionally, the species here recognized as constituting subgenera
PYCNOSTACHYS and CYPERUS (Lye. 1981) have been circumscribed differently
as subgenera Mariscus and CYPERUS. Clarke (1908) and Kuikenthal (1935-
1936) defined subg. CYPERUS as differing from subg. Mariscus in having the
spikelet rachilla firmly attached to the rachis, while the scales are deciduous,
falling from the rachilla as the achenes mature. In species of subg. Mariscus,
the scales remain firmly attached to the rachilla even after the spikelet has
fallen from the rachis. O'Neill (1942) listed some twenty species (e.g., Cyperus
strigosus L., a common species throughout the United States) having charac-
teristics of both subgenera-both the rachillas and the scales are more or less
deciduous. Kiikenthal placed such intermediate species in his concept of subg.
Mariscus, but they are clearly transitional between subg. CYPERUS and subg.
Mariscus. Also, as O'Neill (1942) observed, C. rotundus L. and C. esculentus
L., both of which have always been placed in subg. CYPERUS, have persistent
scales, a feature attributed solely to subg. Mariscus by Kiikenthal. Federowicz
surveyed the epidermal features of leaves and achenes of both subgenera and
found no consistent differences between the two. There is no single character
that consistently separates them. O'Neill (1942, p. 47) stated: "It is ill-advised
to maintain Mariscus as a genus when it is very ill-defined even as a subgenus."
More recently, Koyama (1962b) and Vorster have recognized Mariscus at the
generic level.
Rikli surveyed the anatomy of the leaves and culms of many genera of the
Cyperaceae. He divided Cvperus into two genera, Eucyperus (= Cyperus) and
Chlorocvperus. The latter was characterized by having radiate chlorenchyma
(i.e., kranz anatomy), while the former had nonradiate. Lerman & Raynal
examined the distribution of the C, photosynthetic pathway in the family and
found that Cyperus contained both C, and C4 species. These physiological
differences were correllated with the division that Rikli had based on anatom-
ical information. Subgenus PYCNOSTACHYS corresponds to "Pars Pycnostachys"
(not a valid taxonomic rank) in Kuikenthal's monograph of the genus. Lye
1987]


JOURNAL OF THE ARNOLD ARBORETUM
concurred with O'Neill that .lariscus could not be maintained even at the
subgeneric rank but ought to be included in subg. CYPERUS. The recognition
ofsubgenera PYCNOSTACHYS and CYPERIUS (rather than subgenera Mariscus and
CYPERUS sensu Kiikenthal) is a natural classification that reflects current knowl-
edge of the phylogeny of the genus, as outlined by Raynal (1973).
Van der Veken surveyed variation in embryo shape within the subfamily,
including 162 species of Cvperus. Throughout this genus the embryos were
broadly ellipsoid. There were interspecific differences in size, but these did not
follow taxonomic lines. Van der Veken's data supported a broad concept of
the genus.
Harborne and colleagues surveyed the distribution of flavonoids in South
American, African, and Australian species of C(yperus. They examined about
150 species and reported that each subgenus had a distinct profile of com-
pounds. Subgenus PYCNOSTACHYS is characterized by flavonols, which are ab-
sent in the other subgenera (these have flavones instead). Aurones, which give
a yellowish hue to the inflorescences, are present in subgenera CYPERUS (in-
cluding subg. tlariscus) and ToRUI Nl MsiM but lacking in subgenera PYCREUS
and PCNOSTACHYvs. These investigators believed the differences they reported
confirmed the recognition of PYCNOSTA(HYS as a subgenus distinct from subg.
CYPRUsL. They also concluded that the flavonoid data indicated that no sub-
genus was sufficiently unlike the others to merit generic status. Thus, these
authors also favored a broad concept of the genus.
Chromosome numbers have been reported for about 40 species of (Cperus.
However, even this limited number of counts gives some information about
evolution in the genus. One significant trend is that subg. PYCNOSTA( HYS has
haploid numbers from 8 to 28 (mostly 15-20), while subg. CYPERUS has n =
8-86 (mostly 45-60). The generally lower chromosome numbers of subg.
PYC NOSTACHYS suggest that it is the most primitive subgenus; this is also
indicated by its being the only subgenus with the C, pathway. Different chro-
mosome numbers have been reported for several species. In some species (e.g.,
C. rotundus, n = 16. 48, 54, 76) polyploid races are indicated; in others (e.g.,
C. HIoughtonii Torrey, n = 84, 85, 86), mixoploid.
C('perus in the southeastern United States comprises 63 species in four sub-
genera: five species are adventives from the Old World, seven are endemic, 17
are shared with the northeastern states, 15 are shared with the neotropics, and
the remaining ones have either pantropic or cosmopolitan distributions.
Subgenus PvYNOSTACHYS (C, photosynthesis, spikelets in glomerules or dig-
itate clusters, achenes trigonous). with 150 species worldwide (Lye), includes
14 in our area. Eight of these belong to the New World sect. LUZEOLOIDEI
(Kunth) Clarke (spikelets in glomerulate clusters, scales with proximal abaxial
groove, stamen one per flower). The group has been revised by Denton (1978,
1983). who has also investigated the morphology of the achenes and leaf blades.
She showed that epidermal features of the achenes could be used to distinguish
species. Only one chromosome count is available for this section: (vperus
Era rostis Lam., 2n = 42. This species has been collected as a waif in South
Carolina: it is native to the Pacific coast of the United States and temperate
South America and is naturalized in southern Europe and southeastern Texas.
398
[vo.. 68


TUCKER, CYPERACEAE
The remaining six southeastern species of the subgenus are scattered among
four sections. Section HASPANI (Kunth) Clarke"' (wetland plants; spikelets dig-
itate; achenes ovoid, papillose), is represented in our area by three species.
C('perus IIaspan L. occurs in Coastal Plain wetlands from Virginia southward.
It is one of the few truly pantropic species and is believed to be native to
southeastern Asia, tropical Africa. and the New World tropics. Cyperus dentalus
Torrey, 2n = 34, is a northeastern species of pond shores that extends southward
to South Carolina and Tennessee. This is the only species of the subgenus with
tuberiferous stolons. It is closely related to the southeastern endemic C. Lecontei
Torrey ex Steudel," a Coastal Plain species ranging from North Carolina to
Louisiana.
Section Fusci (Kunth) Clarke' (plants annual; scales ovate: styles and stigmas
very short: achenes ovoid, glossy) is represented in the Southeast by one in-
troduced species. Cyperus difformis L., 2n = 34, a weedy Asian species, was
first collected in the eastern United States in Norfolk Co., Virginia, in 1935 by
Fernald (Tyndale). Lipscomb (1980b) has provided an interesting account of
the spread of this species in North America. The species was first collected in
the New World in New Mexico in 1850. It is a significant weed of rice fields
in California but has not yet become a problem in the southern rice-producing
states (Bryson). In contrast to the other weedy species of the genus (e.g., C.
esculentus), C. diffbrmis is an annual that is capable of completing its life cycle
in only one month; a single plant can produce thousands of achenes. The species
is adapted to ground that is frequently flooded, such as rice fields. The seeds
germinate best under shallow water (Mclntire). The type species of the section,
C. fuscus L., 2n = 72, is Eurasian; it is sparingly adventive from Massachusetts
to Nebraska and Virginia but has not yet been reported from the Southeast.
Subgenus PYCREUS is characterized by having lenticular, laterally compressed
achenes and C, photosynthesis. There are about 120 species worldwide, of
which eight occur in our area. All our species are fibrous-rooted annuals, mostly
less than 30 cm tall, of disturbed wet soils. One. C(yperus louisianensis Thieret,
is endemic to southeastern Louisiana. Five pantropic species occur in our area:
C. flavescens L., 2n = 50, C. pumilus L., 2n = 94, C. flavicomus Michx. (C.
albomarginatus "Nees," see Tucker, 1985a), C. polystachyos Rottb.. and C.
lanceolatus Poiret. Cyperus bipartitus Torrey (C. rivularis Kunth, see Tucker,
1983a), n = 27, is a widespread North American species that also occurs in
the mountains of Mexico, Central America, and southern South America
(Tucker, 1983a). Cyperus filicinus Vahl is endemic to eastern North America
(tidal marshes from Maine to Louisiana).
Subgenus JUNCELLUS has only about six species worldwide. The pantropic
Cyperus laevigatus L., 2n = 80-84, was collected as a ballast plant in Wil-
mington, North Carolina (G. McCarthy s.n. in 1888, Gn!). It apparently never
�C(perus sect. HASPANI (Kunlh) Clarke, Jour. Linn. Soc. Bol. 21: 119. 188.4. TYPE SPECIES: C.
Ilaspan L.
"The name has been attributed to Torrey. but he published it provisionally under (. d:enatus var.
multiradiatus Torrey (Ann. Lye. Nat. Hist. New York 3: 273. 1836). The name (C leconte,' was first
validly published by Steudel (Syn. Pl. Glum. 2: 17. 1854).
"Cvperu.s sect. Fusci (Kunth) Clarke, Jour. Linn. Soc. Bol. 21: 131. 1884. TYPE SPECIES: C('.. / us L
1987]


JOURNAL OF THE ARNOLD ARBORETUM
became established in the eastern United States. This species, which grows in
alkaline or brackish soils, is native to the area from western Texas to southern
California and southern Mexico, to the Lesser Antilles, and to South America.
Subgenus CYPERLIS contains about 400 species worldwide and about 35 in
the Southeast. Among these are pantropic, neotropical, and cosmopolitan rep-
resentatives. About half of the 35 are endemic to the United States, and many
of these are endemic to the Southeast: four are introduced from the Old World.
Plants of sect. UMBELLATI C. B. Clarke are characterized by their caespitose
habit, deciduous rachillas, and appressed, mostly persistent scales. This pan-
tropic group has twelve species in the southeastern United States: C(pcrus
croccus Vahl (C. iglobulosus auct., non Aublet), C. echinatus (L.) Wood (('.
ovularis (Michx.) Torrey), C. P/lukenetii Fern., C. ovatus Baldwin (C. Pollardii
Britton), C. hystricinus Fern., C. refractus Torrey, C. retrofractus (L.) Torrey
(C. dipsaci/ormis Fern., see Carter & Jarvis), C. lancastriensis Porter, C. re-
trorsus Chapman (C. Nashii Britton), n = ca. 90 (Marcks, 1972a), C. thyrsi-
/lorus Jungh., C. retrofle.xus Buckley (C. uni/lorus Torrey & Hooker, non Thunb.).
and C. lentiginosus Millsp. & Chase. Carter (1984) revised the North American
representatives, some ofwhich were also studied by Marcks (1972b) and Tucker
(1983a, 1985b).
Plants of sect. LAXIGIUMI" are characterized by their rhizomatous, single-
stemmed habit, deciduous rachillas, and spreading, more or less deciduous
scales. Species of this section infrequently hybridize with those of the preceding
one (Marcks. 1972a. 1972b). Eight species occur in the eastern United States,
of which four are in our area; there are ten in the mountains of the southwestern
United States, Mexico, and Central and South America. The plants typically
grow in open, dry. sandy or gravelly habitats. The American species were
studied biosystematically by Marcks (1972a, 1972b), and the Mexican and
Central American ones by Tucker (1983a. 1984. 1985a). The species are cy-
tologically similar: all are n = 82 except ('(perus Schtweminiuzi Torrey, n = 84,
85 (Marcks, 1972b). C('perus filicutnmis Vahl (C. .1lartindalei Britton), C. lu-
pulinus (Sprengel) Marcks (C'. filiculnis auct., non Vahl), C'. Grayi Torrey,. and
C. Gravoides Mohlenbrock occur in our area.
The remaining southeastern species are scattered among six mainly pantropic
sections. Section CYPRRUS (sects. E.sculenti Kuikenthal and Rotundi C. B. Clarke)
is most diverse in Australasia (Blake, J. H. Kern). In members of this section
both the scales and the spikelets are persistent (a combination of characters
unknown elsewhere in the genus), and the stolons are tuberiferous. C(perus
routtndus L., purple nut-sedge, is generally acknowledged to be the world's worst
weed. It occurs throughout the Southeast, except in the mountains, but extends
only as far north as southern Missouri and southeastern Virginia. It does not
grow north of the mean 1�C January isotherm (Stoller). C(perus esculentus L..
yellow nut-sedge, is able to tolerate winter air temperatures as low as - 18�C
and is a serious weed in much of the world, especially in cooler regions where
"( p \ rus sect. I AXI UMI ((. i. (Clarke) Kuikenthal. Pllantenr. IV. 20( Heft 101): 220. 1936: based
on lariious subsect. laxi/ltnmi C(. F. Clarkc. Kel Bull. Add. Set 8: 103. 1908. " L .ai.lumaru'.
LE('TOTYPI : SPECIFS (here designated): .a31ari.cuI ta ma, (HBK.) C. B. Clarke ( ('. lammanw e HBK.).
400
[VOL. 68


TUCKER, CYPERACEAE
the more tropical C. rotundus does not grow. These two species also differ in
their thermal optima for growth. In Mexico C. esculentus is found from sea
level to about 2600 m, while C. rotundus occurs from sea level to about 1500
m (Tucker, 1985b). It is unclear whether these species are native to the New
World. Cvperus esculentus now occurs in all 50 states and in southern Canada.
The stoloniferous nature of these two species underlies their success as weeds.
A single tuber can produce a population covering 2-4 m' in two months
(Horowitz). The sharp-pointed stolons can cause puncture wounds in the hands
of farm workers and curious agronomists and penetrate root crops such as
potatoes and yams. In 1821 Elliott noted that Cyperus rotundus was a great
problem for farmers in Georgia and South Carolina. He outlined a method for
removing an infestation by cultivating a fallow field weekly for a year (including
winter), thus allowing the tubers to be killed by exposure to drying and cold
air. Mulligan & Junkins provided a thorough summary of its biology, empha-
sizing weed control and management. Horak & Holt analyzed isozymes in ten
widely separated populations of C. esculentus in California. Genetic variation
served to determine the relative importance of sexual and asexual reproduction.
Results indicated that reproduction by seeds is unimportant in maintenance
of populations in croplands. Stolons and tubers are the primary means of
reproduction. Germinability of seeds from northeastern populations ranged
from 7 to 95 percent; such variation was believed to be genetic (Mulligan &
Junkins). Seeds from a 50-year-old herbarium specimen had 5 percent ger-
mination (Mulligan & Junkins). Cyperus esculentus is self-incompatible (Horak
& Holt).
Members of sect. COMPRESSI Nees" are caespitose annuals with cuspidate
scales and emarginate achenes. Most of the species are native to the Old World
tropics. The pantropic Cvperus compressus L.. n = 64, is the only representative
in the United States. It is found throughout the Coastal Plain and Piedmont,
as far north as Pennsylvania and Missouri. The only other New World species.
C. Wilburii G. Tucker, is endemic to the lowlands of southern Mexico. Its
larger size suggests that it may be a tetraploid derived from C. compressus.
Section IRIOIDEI Nees" comprises several tropical and temperate eastern
Asian species. The plants are annual and have ascending-appressed spikelets
and three-nerved, orbiculate scales. Cyvperus Iria L. is an adventive in all
tropical and temperate regions of the New World and is a common weed
throughout the southeastern Coastal Plain and Piedmont. Apparently, the plants
are cleistogamous. The staminal filaments elongate only enough to bring the
minute anthers into contact with the very short stigmas, which remain inside
the scales at anthesis. Often the anthers are later found agglutinated to the
stigmas.
Section Viscosi C. B. Clarke'" is endemic to the New World and is represented
4( vper'i sect. COMPRESSI Necs. Linnaea 9: 234. 1834. TYE SPE CIES: C. comnpressus L.
(Cy'perus sect. IRIOIDEI Nees, Linnaca 9: 235. 1834. TYPE SPECIES: C. Iria L. Synonym: sect. Iriae
(Kunth) C. B. Clarke. Kew Bull. Add. Ser. 8: 99. 1908.
"( rperu.s sect. Viscosi C. B. Clarke. ,lour. Linn. Soc. Bot. 21: 114. 1884. TYPE SPECIES: C. v'sco' us
Aiton (- C. eleans L.). Synonym: sect. (Gtlunosi (Bock.) Kilkenthal, Pflanzenr. IV. 20(Heft 101):
163. 1936.
1987]


JOURNAL OF THE ARNOLD ARBORETUM
by two species in the Southeast. Plants of this section have spicate inflores-
cences; the spikes are short and dense and appear glomerulate, which apparently
caused Kukenthal to believe them to be closely related to plants of sect.
LUZEOLOEIDi (subg. PYCNOSTACHYS). The plants have kranz anatomy, further
supporting their placement in subg. (C ERIt (Tucker, 1985b). They secrete a
viscid fluid and are sticky when living, hence the appropriate sectional name.
Two species occur in the Southeast. Cy'perus c/legans L. grows from southern
Florida and Texas south to Ecuador. C'yperus o.vylepis Nees ex Steudel is a
South American species that has recently become an adventive in the United
States, where it was first noted in Texas (O'Neill). More recently it has been
reported in Louisiana (Thieret. 1964) and in Charleston County, South Carolina
(,lacl)ouyal 1501, 5 Aug. 1981, DUK!E, N(ti, NYS).
Subgenus TORtI iNiUM has a single representative in our area, the pantropic
and warm-temperate Cyperus odoratus L. It is a common species of disturbed,
wet soils, especially pond shores and stream banks. Five segregate species (e.g.,
C. Engelmannii Steudel. C. Jerruginesccns Bock.) have been recognized at
various subspecific ranks. Evidence for treating these segregates as conspecific
with C. odoratus has been published (Tucker, 1984). Three other species of
this subgenus occur in the New World tropics: C. Correllii (Koyama) G. Tucker
in the Bahamas, C. rhizophorae (C. B. Clarke) Standley along the Pacific Coast
of Central America, and C. filifrmins Sw. in the Greater and Lesser Antilles.
Section REMIREA (Aublet) Kern contains a single pantropic species, Cyperus
pedunculatus (R. Br.) Kern (Remirca niaritima Aublet), beach-stars. In our
area it occurs only in Peninsular Florida. The rhizomatous plants form mats
that bind sand dunes. This species has been treated as constituting a monotypic
genus, Remirea, which Kuikenthal placed in the Rhynchosporoideae. Metcalfe
and Oteng-Yeboah showed convincingly that the anatomy of C. pedunculatus
is similar to that of the kranz species of Cyperus. Within Cyperus, the thickened
upper internode ("corky organ") of the one-flowered spikelets suggests a re-
lationship with subg. TORUIi.iNIUM (C. odoratus typically has spongy, thickened
rachilla internodes). Such internodes may serve to make the achenes buoyant,
thus contributing to dispersal by water, but experimental evidence for this
supposition is lacking.
Ri I RT N( "S:
Under family references see Ai IAN ct al. : BARNARI): BARROS (1938): BI AL: BENTHAM:
BIi ASFR (1940, 1941 a, 1941 c): HBRxsiI i ON; BREwBAKI R: BROwN: BURKHAI.TER; CAROLIN
et al.: ('I ARKi (1908, 1909) ( CLIF ORD & HIARBORNE : EITN (1976a): EvYLs & ROBER ISON:
1ASS ITT; FERNA I): GiODtRI RA & WOOTIEN; (jON(t AROV ci a!.; joo)I et al.: HARIORNI;
HARiORNE et al.: Ho i M et al.; , oi-ti st: HI AN(: J. HIi('HINSON: F. D. KE:RN: J. H.
KERN; KESSiHR & S ARBUmK: KOVYAMA (1962b); Li MAoti & DE(CASNF: LlRMAN &
RAYNAI.: LoIROiNON: M -ATi;!: M li.st; MiL(.AIt.; NAPi'IR (1965); Nhis VON Es-
E NliB K; NoI.ii & MiRPiHY ()O'Nl:II.L; PATn(i; RA NALi(1972, 1973); RiKH; S('HiI.Z/. -Mort:I
(1959. 1964): SMITHi c al.: SIANOI)I ; Ti RI et al.: TII z:; TORREY; VAN I)I:R VI EKiN: and
WINFREY & SAMSEi.
AvYIRS, B. The genus C('pcrus in Mexico. ('athol. Univ. Am. Biol. Stud. 1: 1-103. 1946.
[Eighty species.]
402
[vol. 68


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Southeast, sheds dormant seeds. Dormancy broken by stratification, scarification,
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& . The possible ecological significance of the light requirement for ger-
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& . Seasonal changes in the germination response of Cvperus inflexus
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Cyperus inflexus. Ecology 63: 248-252. 1982. [Wetting and drying cycles decrease
time needed for germination once other conditions are met.]
BENDIXEN, L. E. Anatomy and sprouting of yellow nutsedge tubers. Weed Sci. 21: 501-
503. 1973.
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CHERMEZON, H. Sur la position systematique du genre Remirea. Bull. Soc. Bot. France
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CHETRAM, R. S., & L. E. BENDIXEN. Gibberellic acid plus cytokinins induces basal bulbs
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COSTA, J., & A. P. APPLEBY. Response of two yellow nutsedge varieties to three her-
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COUR, P. Cvperus esculentus L.. C. rotundus L.. et ('. rotundus var. brevibracteatus Legr.:
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DENTON, M. F. A taxonomic treatment of the Luzulae group of Cvperus. Contr. Univ.
Mich. Herb. 11: 197-271. 1978.


JOURNAL OF THE ARNOLD ARBORETUM
Anatomical studies of the Luzulae group of C'perus (Cyperaceae). Syst. Bot. 8:
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Jour. Hot. 57: 691-703. 1970a. [= C. involucratus Rottb., a species cultivated in
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Xylem derived from intercalary meristem of (tperu.s alternifbIius. Bull. Torrey
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SControl of the internodal intercalary meristem of (C'vprus alterndiolius. Am.
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. (perus difformis L. (Cyperaceae) in North America. Ibid. 320-327. 1980b.
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distribution maps.]
S & H. H. ILTIS. Post-glacial hybridization of Cyperus Schweinitzii and C. maci-
lentus. (Abstract.) Am. Jour. Bot. 54: 659, 660. 1967.
McGivNEY, M. V. A revision of the subgenus Eucyperus found in the United States.
Cathol. Univ. Am. Biol. Ser. 26: 1-74. 1938. [Keys. descriptions, specimen citations;
illustrations of scales and achenes.]
MCINTIRE, S. Seed reserves in temperate Australian rice fields following pasture rotation
and continuous cropping. Jour. Appl. Ecol. 22: 875-884. 1985. [C. diflormis is one
of the two most abundant weeds; its seeds germinate best under flooded conditions.]
405
1987]
TUCKER, CYPERACEAE


JOURNAL OF THE ARNOLD ARBORETUM
M(LAU(HItN, A. D. The genus ('vperus in the West Indies. Cathol. Univ. Am. Biol.
Stud. 5: 1-108. 1944.
MOHAN RAM, H. Y.. & M. BAFRA. Stimulation of flower formation by cytokinins in
the excised immature inflorescences of (Cperus rotundus. Phytomorphologv 20: 22-
29. 1970.
Mom [NBROCk. R. H. A new species of( iperus from the Illinois sand prairies. Brittonia
11: 255. 256. 1959. [C. Grayoides, illustrated, also in Louisiana and eastern Texas:
see MAR KS (1972a).]
The Cyperaceae of Illinois. I. Cvpcrus. Am. Midl. Nat. 63: 270-306. 1960.
[Keys, descriptions, illustrations.]
Mui.iiG.AN, G. A., & B. F. JI NKINS. The biology of Canadian weeds. 17. (C'perus
ccudentus L. Canad. Jour. Pl. Sci. 56: 339-350. 1976.
O'NEii, 11. T. The status and distribution of some Cyperaceae in North and South
America. Rhodora 44: 43-64. 1942.
OTir-N,-YEBOAH, A. A. Morphology, anatomy, and taxonomy of the genus Renurea
Aublet (Cyperaceae). Boissiera 24: 197-205. 1975.
OVERTON, J. B. Studies on the relation of the living cells to transpiration and sap-flow
in ('perus. Blot. Ga/. 51: 28-63. 1911.
PaX)DHY, M. D. Studies in Cyperaceae. I. Embryology of ( perus Iria L. Nat. Inst. Sci.
India Proc. 37: 1-10. 1971.
PHATIAK, S. C., D. R. St MNiR. I. D. WVii s, I). K. BEIti. & N. C. GLAzF. Biological
control of yellow nutsedge with the indigenous rust fungus Puccinia canaliculata.
Science 219: 1446, 1447. 1983.
Pu otE, F., & R. GERDOL. Rice-field weed communities in Ferrara Province, northern
Italy. Aquatic Bot. 10: 317-328. 1981. [(C )perus diut/rmis L. and (. serotinus Rottb.
are important weeds.]
PORCHeR, F. P. Resources of the southern fields and forests. ed. 2. [i + ] xv x 733 pp.
Charleston, South Carolina. 1869. [( perus escule'ttus, 684, as C. repens: C. rotun-
dus, 685, as C. hydra.]
PRKASII., N. A survey of the leaf structure and its relationship to photosynthetic path-
ways in certain Malaysian plants. Malaysian Jour. Sci. 4(A): 67-73. 1976. [C. dif/usus
Vahl is C,.]
RAYMOND, M. A note on > Crpcrus I'Ieatherhianus. Rhodora 64: 349, 350. 1962.
RAYNAI, J. Notes cyperologiques: V. Sur un groupe de C (yprus montagnards afro-
americains. Adansonia 6: 385-392. 1967. [Illustrations; distribution map of C. pro-
IA.vus HBK. omits occurrences in Louisiana and Central America, see TU(iCKER (1983a).]
Notes cyperologiques: 31. Melanges nomenclaturaux (Cyperoideae). Adansonia
17: 43-47. 1977. [Notes on typilication of '. , ihantueus Vahl and C. odoratus, both
occurring in the Southeast.]
RIEIo, M. S. The genus C perus in North Carolina. Jour. Elisha Mitchell Sci. Soc. 52:
295-306. 1936.
SCANION, G. M. A study of the genus C(iperus in the Hawaiian Islands. Cathol. Univ.
Am. Biol. Ser. 41: 1-62. 1942. [Keys, descriptions, specimen citations.]
SHARMA, O. P., & R. SHIAM. Occurrence of cuticular papillae in C(perus. Curr. Sci.
Bangalore 50: 236. 1981. [C. pilousx Vahl has one to four papillae surrounding each
abaxial stoma: such papillae are rarely present in C. digitatus Roxb.. C. e.xaltatus
Retz., and C. rotundus.]
SoNi, S. 1., P. B. KAIUEMAN, & W. C. lBiti ow. Electron microprobe analysis of silica
cells in leaf epidermal cells of ( perus alterni/olius. Plant Soil 36: 121-128. 1972a.
-- - , & - . Electron microprobe analysis of silicon and other elements
in developing silica cells of leaf and internode of Crperus alternifolius. Ann. Bot.
(London) 36: 611-619. 1972b.
STIot.iuR, E. W. Effect of soil minimum temperature on differential distribution of
406
[VOL. 68


TUCKER, CYPERACEAE
Cvperus rotundus and (C'perus esculentus in the United States. Weed Res. 13: 209-
217. 1973.
, D. P. NEMA. & V. M. BAHN. Yellow nutsedge tuber germination and seedling
development. Weed Sci. 20: 93-97. 1972.
TAYLOR, J. R., & D. K. EVANS. A taxonomic study of the genus C(perus (Cyperaceae)
in West Virginia. ASB Bull. 25: 64. 1978. [Fourteen species: (C. croceus (C. globu-
losus), new state record.]
THIERET, J. W. More additions to the Louisiana flora. Sida 1: 294. 295. 1964. [Cyperus
d(iformis, (. oxylepis, and C retro(/h'xus (as C. uniflorus).]
SCyperus louisianensis (Cyperaceae), a new species from southern Louisiana.
Proc. Louisiana Acad. Sci. 40: 23-26. 1977. [Subgenus Pvcrcus: related to C. hi-
partitus Torrey: illustrations.]
THOMPSON, K., P. R. SHEWRY. & H. W. WVooinHot se. Papyrus swamp development in
the Upemba Basin, Zaire: studies of population structure in C(perus papyrus stands.
Bot. Jour. Linn. Soc. 78: 299-316. 1979.
TICKER. G. C. Taxonomy of the genus Cyperu. (Cyperaceae) in Costa Rica and Panama.
Syst. Bot. Monogr. 2: 1-85. 1983a. [Fifty species, keys, descriptions, distribution
maps: subgeneric classification.]
. Two new species of Cvperus (subgenus I'rotocvperus) from Mexico and Central
America. Bull. Torrey Bot. Club 110: 343-347. 1983b. [C. microbrunneus, C. na-
yaritensis of subg. Pycnostachys; illustrations.]
.Taxonomic notes on two common neotropical species of C(perus. Sida 10: 298-
307. 1984. [C. odoratus.]
- . (perus /lavicomus, the correct name for (Cyperus abonmarginatus. Rhodora 87:
539-541. 1985a. [A pantropic species occurring in the Southeast.]
A revision of the Mexican species of Cvperus L. (Cyperaceae). 285 pp. tLnpubl.
Ph.D. Dissertation. Duke Univ. 1985b. [Eighty-five species.]
The correct name for Cyperus cavennens.is (C. /larus). Cyperaceae. Southw. Nat.
30: 607, 608. 1985c. [C. aggregatuus (Willd.) Endl. the correct name for a neotropical
species that occurs northward to Texas and Louisiana.]
The species of Cvperus described by Liebmann in "Mexicos halvgraes." Syst.
Bot. 11: 14-19. 1986a.
The distribution of C, and C, species of C(perus (Cyperaceae) in North and
Central America. (Abstract.) Am. Jour. Bot. 73: 792. 1986b. [Abundance calculated
from number of herbarium collections: relative abundance of C, species increases
with latitude.]
New records of Cyperus (Cyperaceae) from West Virginia. Castanea 52: 145.
146. 1987. [C(. Hlougohtonii. C. iria. C. plystachyos.]
TUMBLESON, M. E., & T. KoMMEDAHtl. Reproductive potential of C(perus esculentus L.
by tubers. Weeds 9: 646-653. 1961.
TYNDALE. R. W. Distribution of C'vperu dilArnims L. (Cyperaceae) in the southeastern
United States. Castanea 48: 277-280. 1983.
VERMA, S. C.. A. PAL, & B. L. SRIvA sIvs. Anatomical studies on some species of
Cvperus L. Pl. Sci. 5: 52-59. 1973.
VORSTER, P. J. Revision of the taxonomy of llariscus Benth. and related genera in
southern Africa. 348 pp. Unpubl. D.Sc. Dissertation, Univ. Pretoria. 1978.
WILLIAMS, R. D. Intraspecific competition of yellow nutsedge. Proc. So. Weed Sci. Soc.
34: 231-238. 1981.
WILLS, G. D., & G. A. BRISCOE. Anatomy of purple nutsedge. Weed Sci. 18: 631-635.
1970.
WEEDON, R. R., & H. A. STEVENS. C(perus iuscus in Nebraska and South Dakota.
Rhodora 71: 433. 1969.
YANSEN, L. L. Morphology and photoperiodic responses of yellow nutsedge. Weed Sci.
19: 210-219. 1971.
407
1987]


JOURNAL OF THE ARNOLD ARBORETUM
9. Kyllinga Rottboell. Descr. Icon. Rar. Nov. PI. 12. 1773, non. cons.
Small, rhizomatous or tufted perennials (1 species annual). Culms trigonous
or roundly trigonous, smooth. Leaves 1-5, basal: sheaths short, closely fitting
the culms, ligule lacking; blades flat or V-shaped in cross section [lacking], the
margins and keels scabrellate. especially distally: chlorenchyma radiate; bundle
sheaths 2-layered ("(Cperus type"). Involucral bracts 2-4, leaflike. horizontal
to slightly reflexed or erect. Spikes 1-4, sessile, densely ovoid to cylindrical.
Spikelets 15-150 per spike. not readily distinguishable without magnification,
ovate to lanceolate. decidedly flattened. Scales 4, the 2 basal minute, the 2
distal much longer, making up the bulk of the spikelet, the lower of these (the
third scale of the spikelet) subtending a perfect flower, the upper (fourth scale)
slightly smaller, sterile or infrequently bearing I or 2 (often abortive) stamens.
The fertile scale of the spikelet ovate, conduplicate. with a conspicuous smooth
or spinulose-scabrellate [fimbriate or erose] keel terminating in a mucronate
or mucronulate [aristate] apex, laterally 2- to 4-nerved. Flowers perfect. Peri-
anth lacking. Stamens 1-3: filaments ribbonlike. about as long as the subtending
scales; anthers oblong-elliptic to linear, the apices of the connectives not pro-
longed: pollen grains 4-aperturate [uniaperturate], obovoid, psilate, trinucleate.
Styles capillary, smooth; stigmas 2, about as long as the styles. Achenes len-
ticular, laterally compressed, narrowly ovoid to oblong or ellipsoid, about ,
the length of the subtending scale, the apex obtuse, apiculate, the base cuneate
to rounded. barely to decidedly stipitate, the surface puncticulate. Embryos
narrowly ellipsoid. Base chromosome number 60. (Cy'pcrus subg. Kyl/inga
(Rottb.) Valck.-Suringar.) TYPE SPE(IES: K. monocephala Rottb., nom. illeg.
(= K. nemoralis (J. R. & G. Forster) Dandy ex Hutchinson & Dalz., typ. cons.).
(Named for Peter Kylling, Danish botanist, d. 1696.)
A genus of about 40-45 species, nearly all of which are tropical. The greatest
diversity is in tropical East Africa and Madagascar. where there are 30-35
species. Eight occur in southern Asia, three or four in eastern Asia, and two
in Australasia. Two (neither endemic) grow in the Hawaiian Islands, but none
occurs in Europe. There are eight species in the New World: three of these.
Kyllinga pumila Michx., K. odorata Vahl, and A. hreviol/ia. 2n = 120, which
occur in the Southeast, are pantropic. Ky/linga vaginata Lam. and K. tibialis
Ledeb. are species of littoral habitats in the Caribbean. South America, and
tropical West Africa. Kyllinga nudiceps C. B. Clarke is endemic to Isla del
Coco, in the Pacific some 300 km southwest of Costa Rica. Kyllinga squumulata
Thonn. ex Vahl (Cyperus .lh'eli Mattf. & Kfikenthal), from tropical Asia. is
introduced in Florida and the West Indies; K. breviholioides (Delahoussaye &
Thieret) G. Tucker,' from temperate eastern Asia. has become sparingly es-
tablished in the eastern United States in the area from Connecticut to western
North Carolina and Tennessee. The four southeastern species are mostly weedy
plants of disturbed, usually moist. sunny places. Ky/lingu pumila is a common
A /htlim,'a hn'�rtiti udt (iih)elahmossaN c & I hicret (. T ucker, comb. no\.. based on11 ('p rui h/ c
f/lo //ii I )clahoussac &. I hicret. Sida 3: 1 I. 1)67.
408
[voi. 68


weed of lawns and croplands in the eastern United States from Pennsylvania
and Missouri south to the Gulf Coast.
Kvllinga differs from Cyperus, with which it has been combined by some
workers, in its very short rachilla and in the two lowest sterile scales of its
spikelets being greatly reduced. Taxonomically useful characters have been
reviewed by Tucker. The most important of these are habit (rhizomatous
perennials or caespitose annuals), length and orientation of the involucral bracts,
and length of the anthers. Such characters as number of stamens and presence
of spinulose prickles on the keels of the scales have previously been used
(Delahoussaye & Thieret) but frequently vary within individuals of the same
species and sometimes within spikes of a single plant.
The plants are probably at least partly wind pollinated. However, because
of the close spacing of the spikelets within an inflorescence, some anthers
probably shed their pollen directly onto stigmas of adjacent spikelets. Insect
pollination may be important in some species with conspicuous, whitish or
cream-colored spikes (e.g., Kyllinga odorata), as it is in many species of Rhyn-
chospora sect. DICHROMENA. Syrphid flies have been observed visiting indi-
viduals of K. tibialis in Costa Rica (AlacDougal 1190, DUKE) and K. odorata
in Mexico (Tucker 2222, DUKE).
REFERENCES:
Under family references see BARROS (1935): BEAT: BENTHAM: BLASER (1940, 1941a):
CAROLIN et al.; CLARKE (1908, 1909); Ei EN (1976a); FASSETT: GODFREY & WOOTEN;
GONCHAROV et al.; HARBORNE: HARBORNE ci al.; HOLM et al.; HOLTTUM; HUANG;
J. HUTCHINSON; KUNTH; LE MAOUT & DECAISNE: LERMAN & RAYNAL; NAPPER (1966);
NEES VON ESENBECK, O'NEILL, RAYNAL (1972, 1973); RIKLI; SCHULZE-MOTEL (1959,
1964); TORREY: and VAN DER VEKEN.
DELAHOUSSAYE, A. J., & J. W. THIERET. C(perus subgenus Kyllinga (Cyperaceae) in the
continental United States. Sida 3: 128-136. 1967. [Synopsis; illustrations of spikelets
and achenes, distribution maps.]
GOVINDARAJALU, E. The systematic anatomy of south Indian Cyperaceae: Cyperus sub-
genus Kvllinga (Rottb.) Suringar. Jour. Linn. Soc. Bot. 62: 41-58. 1969.
LYE, K. A. New taxa and combinations in K/llinga. Nordic Jour. Bot. 1: 741-747.
1981.
MCNAUGHTON, S. J. Ecology of a grazing ecosystem: the Serengeti. Ecol. Monogr. 55:
259-294. 1985. [During the wet season, leaves of Kvllinga nervosa provide forage
for the Thomson's gazelle.]
PADHYE, M. D. Studies in the Cyperaceae. IlI. Life history of Kvllinga brevifolia Rottb.
with a brief discussion on the taxonomic position of Kvllinga. Bot. Gaz. 132: 172-
179. 1971.
TUCKER, G. C. A revision of the genus Kylling/a Rottb. (Cyperaceae) in Mexico and
Central America. Rhodora 86: 507-538. 1984. [Six species; keys, descriptions, dis-
tribution maps. extensive specimen citations.]
10. Lipocarpha R. Brown in Tuckey, Narr. Exped. Congo 5: 459. 1818, nom.
cons.
Small, caespitose annuals of wet sandy or peaty soils. Roots fibrous, rhizomes
absent. Culms 1-20(-100), usually densely clustered, erect, spreading, or curved,
TUCKER, CYPERACEAE
409
1987]


JOURNAL OF THE ARNOLD ARBORETUM
filiform, terete, glabrous. Leaves I or 2, basal, filiform, about as wide as the
culms, the lower reduced to a bladeless sheath or a sheath bearing merely an
involute appendage, the upper with blade up to '/ as long as the culm, or
reduced like the lower one: stomata paracytic: chlorenchvma radiate: the bundle
sheaths 2-layered ("C('pIrus type"). Inflorescences unbranched, a sessile cluster
of 1-4 dense spikes; bracts 1-4, filiform, 1-4 times as long as the spikes, leaflike,
the longest erect, appearing as a continuation of the culm, the other(s) shorter
than or equaling the spikes, borne approximately perpendicular to the culm:
rays none. Spikes ("spikelets") 1-4. sessile, ovoid [globose]: denuded rachis
persistent, with rhombic scars where the spikelets were attached. Spikelets
("flowers") [20-]50-150, densely spirally arranged, borne approximately per-
pendicular to the rachis, deciduous. Scales (1, 2, or) 3: outer scale lanceolate
to ovate-lanceolate, planar or nearly so. with 2 conspicuous medial veins and
a less conspicuous central one. laterally weakly 1- or 2-nerved or essentially
nerveless, mucronulate [aristate]; inner scale hyaline, equaling or shorter than
the outer, or reduced to a scalelike appendage much shorter than the outer,
with 3-5 inconspicuous veins or veinless. or absent: third scale present between
the outer scale and the achene in some species, similar to or smaller than the
second. Flowers perfect. Perianth lacking. Stamens I or 2; filaments capillary,
about ` as long as the outer scale: anthers ovoid, the apices of the connectives
not prolonged: pollen grains 4-aperturate. obovoid to subspheroidal. psilate or
scabrate. Styles filiform: stigmas 2, about /. as long as the styles, minutely
swollen apically, glabrous, deciduous before the achenes mature. Achenes tri-
gonous to terete, obovoid to cylindrical, slightly shorter than the outer scale,
the base sessile to stipitate,. the apex obtuse to subtruncate, apiculate. the surface
papillose. Embryos ellipsoid. Base chromosome number 6. (Incl. Ascolepis Nees
ex Steudel, Hemicarpha Nees ex Arnott.1') TYPE SPECIES: L. sene~ialensis(Lam.)
T. & H. Durand (L. ar'genteum (Vahl) R. Br., nom. illeg.; see Haines & Lye).
(Name from Greek, lipo, to fall, and carpha, chaff, referring to the deciduous
hyaline inner scale of the spikelet.)
A genus of about eight species occurring in tropical and warm-temperate
regions. Five grow in North America: Lipocarpha maculala (Michx.) Torrey,
on the Coastal Plain from Virginia to Texas, southward into the tropics: L.
occidenlalis, restricted to the Pacific coast: L. Drummondii, from Oklahoma
and Texas west to New Mexico: .. aristulata, across the United States from
South Carolina and Florida west to Washington and California; and L. mi-
'"he inclusion of lriniitur/tha in i;prarpha necessitates the Iollo ting new combinations for
species occturing in the Ne' W\ orld:
I ip ti , ati/it i r ah i alad cH (( ox .)( i. ucker. based on lrtino' rpic a noron thfl/ta \ar. ai nitulatia ( o . Bull.
I orre'. Club 21: 3". 111 4.
L.. )rniiinii nitdu (Necs) (i. Tucker I ased on h ionTrpha l)ruit nu it/ Nos i Nees Martius, I. Brasil.
2(1): (, 1S42.
/. nicrantha (Vahl) (. Tucker. hased on .Sict>71 niurantthtis Vahl, Enum. 2: 254. 1806.
I. tad'/eualtis ( rao ) ( i. fucker, based on ll/ I ii'iarpha i. ci/ n'uta/it (ira ., Proc. Am. Acad. 7: 391.
l S/i iihinhuri' u (Ericdl.) (i. I ucker, k ased on llH im piha rp / Sctiio/iir ki Friedl. Am. .lou. lBot. 28:
b0(). 1941.
410
[voL. 68


TUCKER, CYPERACEAE
crantha, throughout the United States and southeastern Canada, southward to
tropical South America. Lipocarpha Schomburgkii is known only from the
Guyana region of northern South America.
All species are small, inconspicuous plants of disturbed wet soils, especially
shores of ponds and pools. Because of their small size (less than 30 cm tall,
and often less than 1 cm!), they are easily overlooked and are probably more
frequent and widely distributed than available collections indicate.
Raynal's view that Lipocarpha is a highly reduced derivative of Cyperus
seems well founded and is accepted here. The fact that both genera have
"Cvperus-type" kranz anatomy (Metcalfe) further strengthens this conclusion.
The achene and subtending scales of Hemicarpha are probably homologous to
a single spikelet of Kyllinga or ('yperus. Friedland suggested that the inner
hyaline scale represented five perianth members that correspond to the bristles
subtending the achenes in some species of Scirpus. Raynal's interpretation of
the inner scales of Lipocarpha (and Ilemicarpha) as reduced scales of a spikelet
appears more plausible than Friedland's view.
Haines & Lye studied the African species previously assigned to Hemicarpha
and Lipocarpha and concluded that the two genera should perhaps be merged.
Goetghebeur (pers. comm.) has recently studied all the Old World species of
these genera, as well as those of the closely related genus Ascolepis. He con-
cluded, as I had from my independent investigations, that the three genera
should be combined.
Chromosome numbers have been reported for Lipocarpha argentea R. Br.
(2n = 26) and L. microcephala Kunth (2n = 46). This suggests a base chro-
mosome number of 6.
No species is gathered as food or for medicinal purposes. Lipocarpha argentea
and L. microcephala (R. Br.) Kunth are recorded as weeds in eastern Asia
(Holm et al.).
REFERENCES:
Under family references see BARROS (1938); BEAL; BENTHAM; BLASER (1940. 1941a);
BROWN; CAROLIN e/ a/.; CLARKE (1908); ElIEN (1976a); FASSETT; GODFREY & WOOTEN;
HOLM et al.; HUANG; J. HUTC HINSON; J. H. KERN; KOYAMA (1962b); KUNTH; LE MAOUT
& DECAISNE; LERMAN & RAYNAL; METCALFE; NAPPER (1965); NEES VON ESENBECK;
O'NEILL; SCHULZE-MOTEL (1959, 1964); STANDLEY; TEERI et al.; TORREY; and VAN DER
VEKEN.
FRIEDLAND, S. The American species of Ilemicarpha. Am. Jour. Bot. 28: 855-861. 1941.
[Revision of the North and South American species: keys, distribution map, de-
scriptions; discussion of morphology of the spikelets; no specimen citations.]
HAINES, R. W., & K. A. LYE. Studies in African Cyperaceae IV, Lipocarpha R. Br.,
Hemicarpha Nees, and Isolepis R. Br. Bot. Not. 124: 473-482. 1971.
KOYAMA, T. The genus Lipocarpha R. Br., its morphology and systematic position in
the family Cyperaceae. (In Japanese: English abstract.) Acta Phytotax. Geobot. 33:
218-226. 1982.
PALLA, E. Ober den morphologischen Wert der Bliite der Gattungen Lipocarpha und
Platylepis. Ber. Deutsch. Bot. Ges. 23: 316-323. pl. X7I'. 1905. [Floral diagrams.]
RAYNAL, J. Notes cyperologiques: VII. Sur quelques Lipocarpha africains. Adansonia.
II. 7: 81-87. 1967. [Two new species; illustrations.]
1987]


JOURNAL OF THE ARNOLD ARBORETUM
Il I
,1 \ /
Y )
/,
/1 ;
//
I
1/
]
' /
MK-S [ /,
FIGUt:R 3. Rh ynchospora sect. DIIIROMENA. a-c, R. colorata: a. habit (note rhizomes
to right). x /:2 b. mature achene, tubercle scarcely decurrent on body of achene, x 20:
c, same, in longitudinal section, the 2 layers of the achene wall separated by dotted line,
seed coat unshaded, endosperm stippled, embryo unshaded, x 20. d-k, R. Iloridensi.:
d. head of spikelets subtended by involucral bracts, x 3: e, I large and I small spikelet
enclosed by 2 scales, x 6: f, abaxial surface of spikelet. 2 scales removed, flowers pro-
tandrous, x 10: g, same spikelet, adaxial surface, I stamen and 3 scales removed. x 6:
412
[VOL. 68
d <


Tribe SCHOENEAE Dumortier, Fl. Belg. 144. 1827. (Tribe Rhynchosporeae Fenzl
in Endlicher, Gen. Pl. 2: 115. 1836.)
11. Rhynchospora Vahl, Enum. Pl. 2: 229. 1806, nom. cons.
Small to large, caespitose or single-stemmed, perennial [annual] herbs of
moist open woods, bogs, pocosins, ditches, and pond shores. Roots fibrous;
rhizomes or stolons present in a few species. Culms trigonous, subtrigonous,
or terete, smooth throughout or ribbed just below the inflorescence, glabrous.
leafy [leafless]. Leaves numerous, basal, cauline, or both; basal leaves with
blades flat to conduplicate or involute-filiform, the margins and midveins
generally scabrellate with unicellular [multicellular] prickles, the surfaces gla-
brous or with prickles like those on the margins, or pubescent with long, flexible,
unicellular hairs, or papillose (in R. alba); cauline leaves shorter than but
otherwise similar to the basal ones; stomata paracytic, generally confined to
the abaxial surface; chlorenchyma not radiate [radiate in some tropical species].
Inflorescences terminal (sometimes also lateral, the lateral ones smaller and
less branched than the terminal), fasciculate or cymose: bracts 1-6, leaflike
(sometimes basally whitened); rays slender, terete, smooth or scabrellate; heads
loosely to densely ovoid or capitate. Spikelets solitary, globose, ellipsoid, or
slenderly lanceolate, the 1-5 basal scales sterile. Scales spirally arranged, closely
imbricate, ovate to lanceolate, entire or mucronulate at apex, nerveless to rather
prominently nerved, the midvein most conspicuous. Flowers perfect (the ter-
minal 1 or 2 scales sterile or subtending rudimentary ovaries and functional
stamens). Perianth bristles lacking or 1-6(-20), smooth, barbed, or plumose,
persistent. Stamens (1-)3(-12); filaments capillary or ribbonlike; anthers elliptic
to oblong, the apices of the connectives not prolonged; pollen grains uniaper-
turate, obovoid, psilate or scabrate, binucleate. Styles glabrous; the stigmas
longer than, equaling, or much shorter than the style. Achenes lenticular (dor-
siventrally flattened), ovoid to slenderly ellipsoid, crowned with a pyramidal
to subulate tubercle shorter than to 3 times longer than the body of the achene,
the base sessile to conspicuously stipitate, the lateral edges often raised to form
a conspicuous ridged margin, the surface alveolate to cancellate (rarely smooth
or nearly so), transversely rugulose or not. Base chromosome number 5. (Incl.
Psilocarya Torrey, Dichromena Pers., Calyptrostylis Nees.) TYPE SPECIES: R.
alba (L.) Vahl (Schoenus albus L.), typ. cons. (Name from Greek, rhynchos,
snout, and spora, seed, in reference to the prominently beaked achenes.)
A genus of about 225 species, worldwide in distribution, with greatest di-
versity in the New World tropics; about 60 occur in the southeastern United
States. Temperate North America. especially the southeastern Coastal Plain,
is rich in species, and there are many others in the Old World tropics. Only a
h. flower with subtending scale, anthers fallen. 5 scales and rachilla of spikelet removed,
x 10; i, flower removed from spikelet, anthers dehiscing, styles not yet elongated, stigmas
not receptive, x 12; j, nearly mature achene with persistent style and stigmas, x 20: k,
mature achene, tubercle decurrent on body of achene, x 20.
413
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JOURNAL OF THE ARNOLD ARBORETUM
few species are indigenous to temperate Eurasia-three species in Europe and
four in the Soviet Union east of the Urals.
Kukenthal's worldwide monograph (1949, 1950, 1951) provided a basis for
identification and further study of the genus Rhvnchospora. Gale, in her careful,
well-illustrated monograph, did much to clarify the taxonomy of the North
American species. Thomas (1984) has recently investigated the tropical section
D[) HROMENA (Pers.) Pfeiffer and confirmed its inclusion in Rhynchospora.
The genus is little known cytologically: chromosome numbers have been
published for only ten species (summarized by Thomas, 1984). These suggest
a base chromosome number of 5, in keeping with the base number for other
genera of the family.
There are three subgenera in Rhynchospora (Kukenthal, 1949, 1950, 1951).
The largest of these, including about 54 of the 60 species in our area, is subg.
RHYNCHOSPORA (Eurhytnchosporac Gray), species ofwhich have papery spikelet
scales and stigmas equaling or longer than the styles. Complete descriptions of
the southeastern species were provided by Gale.
Species of sect. DIC'ROMENA have sessile capitate inflorescences and whitish
spikelets often subtended by whitish bracts and lack perianth bristles. The
section is primarily neotropical in distribution and contains 23 species, ofwhich
four are present in the United States. Three occur in the Southeast. Insect
pollination has evolved in plants of this section, as was first noted in 1893 by
DI)e Lagerheim and later studied by Uphof and Leppik.
Thomas (1984) reviewed previous investigations of entomophily in species
of sect. DI)CHROMFNA and made thorough field and laboratory studies. Fifteen
species of bees (Hymenoptera) visit flowers of plants included in this section.
The bees exhibit constancy, visiting four to ten inflorescences in a population
before leaving. The flowers have no fragrance and no nectar: the white color
of the bracts and spikelets attracts the bees, and pollen is the only reward. The
pollen grains have a sticky "pollenkil": thus, they aggregate and stick to the
bee's body and legs. There is probably some transfer of pollen by the wind.
All species of sect. DICHROMENA are self-compatible. Thomas (1984) postulated
that the evolution of entomophily may have permitted the species to radiate
into shaded tropical forests, where a lack of air movement necessary for wind
pollination is compensated for by insect and self-pollination.
No species of Rhynchospora is gathered for food or medicinal uses. Several
species are detrimental weeds in rice fields, both in the Old World and in the
southeastern United States.
REiFRENCEs:
Under family references see BADEN cl al.: BEAI: B:NTHAMI: BER(.(;REN: BLASER (1940.
1941 b) BREWBAKER: (C ARKE (1908. 1909): Eni iN (1976a): EY IEs & ROBERTSON: FASSETT :
G(io)FREY & WooriTN: GON( HiARV ct al.: (GOO)D t al.: HARBORNE: HARHORNF Ct a!.:
HESLA C( a!.: iiEUSSER: HOLM ( ail.: HoLTTIUM: IOTCHKISS: G. E. HUTCHINSON:
J. Hti TilNSON: J. H. KERN: KOYAMA (1961): KUKKONEN (1969. 1986): KUNTH: LE
MAOUT & DECAISNE: IERMAN & RAYNAL: MEEUSE: METCALFE: NAPPER (1964b): NEES
VON ESENBECK: O(,DEN; S(-Ht LzE-Mo IL (1959, 1964): SMITH et al.: STANDI EY: TEERI tI
al.: ITORREY: VANHECKFE and VAN DER VEKEN.
414
[VOL. 68


TUCKER, CYPERACEAE
Under Eleocharis see KUKENTHAL.
GAL E, S. Rhynchospora section lurhvnchospora in Canada, the United States. and the
West Indies. Rhodora 46: 90-134, 159-197, 207-249, 255-278. 1944. [The basic
monograph; distribution maps, descriptions, keys, and illustrations.]
GORDON-GRAY, K. D., & L. L. BANo t. Silica deposits in Rhynchospora species. Proc.
Electron Microscop. Soc. S. Afr. 8: 83, 84. 1978.
GOVINDARAJALU, E. Observations on new kinds of silica deposits in Rhynchospora
species. Proc. Indian Acad. Sci. B. 70: 28-36. 1969.
GUAGLIANONE. E. R. Contribuci6n at estudio del genero Rhynchospora Vahl (Cypera-
ceae) II. Darwiniana 22: 499-509. 1980. (English abstract.) [R. Brittonii and R.
microcarpa: SEM photographs of achenes.]
Alphabetisches Verzeichnis fur Rhynchospora Vahl. 19 pp. San Isidro, Argen-
tina. 1981. [Index to generic and specific names, including synonyms, for KUKENTHAL
(1949. 1950, 1951).]
Contribuci6n al estudio del genero Rhynchospora Vahl (Cyperaceae) IV: R.
iberae, nueva especie de America Austral. Darwiniana 24: 469-473. 1982. [New
species related to R. californica Gale; cross sections of leaves and SEMs of achenes.]
HILL, E. J. The perianth of Rtnchospora capillacea var. leviseta. Rhodora 8: 186, 187.
1906.
HOLM. T. Studies in the Cyperaceae. VI. Dichromena leucocephala Vahl. and D. latifolia
Baldw. Am. Jour. Sci. 154: 298-305. 1897. [Taxonomic history: morphology and
anatomy.]
KRAL, R. A new species of Rhynchospora (Cyperaceae) from southwestern Georgia.
Sida 7: 42-50. 1977. [R. Thornei, from Baker Co.; illustrations, key to new species
and relatives: R. diverg'ns, R. pusilla. R. rarillora, and R. stenophylla.]
LAGERHEIM, M. G. DL. Note sur un Cypcracee entomophile. Jour. Bot. (Morot) 7: 181-
183. 1893.
LEPPIK, E. E. Dichromena ciliata, a noteworthy entomophilous plant among the Cyp-
eraceae. Am. Jour. Bot. 42: 455-458. 1955.
MACBRIDE, J. F. Some Peruvian sedges. The status of Rhynchospora. Fieldiana Bot. 4:
165-168. 1929.
RAGONESE. A. M., E. R. GUA(IIANONE. & C. DIZEO DE STRITTMATTER. Desarollo del
pericarpio con cuerpos de silice de dos especies de Rhynchospora Vahl (Cyperaceae).
(English abstract.) Darwiniana 25: 27-41. 1984. [Developmental study of the peri-
carp in R. corvmhosa (L.) Britton and R. scutellata Griseb., emphasizing the origin
and differentiation of the silica bodies in the outer cell walls: line drawings and
SEMs.]
TAKEDA, T., O. OENL, & W. AGAxTA. The occurrence of Cf species in the genus Rhyn-
chospora and its significance in kranz anatomy of the Cyperaceae. Bot. Mag. Tokyo
93: 55-65. 1980.
THOMAS. W. W. A new species of Rhynchospora (Cyperaceae) from Venezuela. Contr.
Univ. Mich. Herb. 15: 311-313. 1982.
. The systematics of Rhynchospora section Dichromena. Mem. N. Y. Bot. Gard.
37: 1-116. 1984. [Mostly neotropical section with three species occurring in the
Southeast.]
UPHOF, J. C. T. Die Entomophilic der Cyperaceengattung Dichromena Michx. Ber.
Deutsch. Bot. Ges. 50: 208-214. 1932.
12. Dulichium Persoon, Syn. Pl. 1: 65. 1805.
Perennial herbs of swamps, fens, and shores. Roots fibrous; rhizomes hor-
izontal. Culms 1-3, terete, hollow, glabrous. Basal leaves bladeless: sheaths
appressed; cauline leaves several, the blades lanceolate, about 1-2 times longer
415
1987]


JOURNAL OF THE ARNOLD ARBORETUM
than the sheaths, auriculate, planar, with margins and midveins densely sca-
brellate abaxially; stomata confined to the adaxial surfaces (sometimes a few
present near the margins on the abaxial surface): chlorenchyma not radiate;
air cavities present. Inflorescences solitary in the axils of the upper leaves: rays
and rachises slender, compressed, scabrellate on the edges; spikes loosely ovoid,
appearing flattened from the distichous arrangement of the spikelets. Spikelets
3-20, linear-lanceolate, flattened: rachilla persistent, the internodes with hya-
line margins, the lowermost scale sterile (except in the terminal spikelet). Scales
3-9, deciduous as the achenes mature, lanceolate, conduplicate, acute, 5- to
9-nerved, the midveins scabrellate. Flowers perfect. Perianth bristles 6-9, 1-
2 times as long as the mature achene, retrorsely barbed. Stamens 3; filaments
ribbonlike, nearly as long as the scales: anthers linear, the apices of the con-
nectives minute. Style capillary, glabrous; stigmas 2, about as long as the style.
glandular-pubescent. Achenes planoconvex, narrowly ellipsoid, the apex acute.
the base stipitate, the surface puncticulate. Embryos turbinate. Base chromo-
some number 16. TYPE SPECIES: D. arundinaceum (L.) Britton. (Name from
the Greek duo, two, and leichon, scale, referring to the two-ranked scales of
the spikelets.)
A monotypic genus of wetland plants endemic to temperate North America.
Dul/ichium is easily distinguished from other Cyperaceae by its characteristic
distichous spikelet scales and its three-ranked cauline leaves. An interesting,
apparently uninvestigated feature of the plants is that in adjacent culms arising
from the same rhizome, the leaves are spiraled clockwise in one and counter-
clockwise in the next. The single species, 1). arundinaceum, is distributed from
Newfoundland to southeastern Manitoba, south to southern Florida and eastern
Texas, and disjunctively in the area from northwestern Montana and south-
western British Columbia south, mostly west of the Cascades and the Sierra
Nevada. to central California (Wood. 1972, map). The genus had a wider
distribution during the Pleistocene when it occurred in Europe (Wood, 1971,
map). Fossils of this species are known from the Pliocene in the Soviet Union
(Daghlian). Infraspecific variation in fossil achenes from Europe has been stud-
ied by Truchanowiczowna.
Dulichiun, has usually been placed in the tribe Cypereae, near C(vperus.
Linnaeus (Sp. Pl. 1: 45. 1753) included the species in (Cperus, presumably
because of its distichous spikelet scales. The two genera differ, however, in
several important features: Dulichium has widely spaced axillary inflorescences
subtended by leaflike bracts with conspicuous sheaths, while Cy'perus has api-
cally clustered inflorescence branches subtended by sheathless bracts: Duli-
chium has one sterile scale at the base of each spikelet, and Cyperus has two;
Dulichium has perianth bristles, but C(perus does not.
The embryos of Dulichium resemble those found in Rhynchospora, rather
than those of any genus of the Cypereae (Van der Veken). A new monotypic
tribe, the Dulichieae, has recently been proposed for this genus by Schulze-
Motel (1959).
Plants of this genus have been neither reported to have economic use nor
noted as weeds.
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REFERENCES:
Under family references see BEAL; BENIHAM: BLASER (1940, 1941a, 1941b); CLARKE
(1908); COOK; DAGHLIAN: EYLES & ROBERTSON; FASSETT; GODFREY & WOOTEN; GooD
et al.; HOTCHKISS; KUNTH; LE MAOUT & DECAISNE; LERMAN & RAYNAL; MATTFELD;
METCALFE; NEES VON ESENBECK; OGDEN: RADFORD et al.; SCHULZE-MOTEL (1959, 1964);
TORREY; and VAN DER VEKEN.
Under Eleocharis see WOOD.
BELL, F. G. Fossil of an American sedge, Dulichium arundinaceum (L.) Britt., in Britain.
Nature 227: 629, 630. 1970. [Illustrations.]
SCHULZE-MOTEL, W. Dulichieae. eine neue Tribus der Cyperaceae-Scirpoideae. Will-
denowia 2: 170-175. 1959.
TRALAU, H. Extinct aquatic plants of Europe. Bot. Not. 112: 385-406. 1959.
TRUCHANOWICZOWNA, J. Variability of the recent and fossil fruits of the genus Duli-
chium. (Polish and English summaries.) Acta Palaeobot. 14: 119-143. 1973.
WOOD, C. E., JR. Some floristic relationships between the southern Appalachians and
western North America. Pp. 331-404 in P. C. HOLT. ed.. The distributional history
of the biota of the southern Appalachians. Part II. Flora. Blacksburg, Virginia. 1971.
[fig. 1, extant and known former distribution of D. arundinaceum.]
13. Schoenus Linnaeus, Sp. P1. 42. 1753: Gen. Pl. ed. 5. 26. 1754.
Caespitose perennials of open sunny wetlands. Rhizomes short, oblique.
Culms terete, hollow, glabrous. Leaves all basal: sheaths tough, glossy, glabrous,
ligule lacking; blades linear, subcylindrical, upper surface flat or broadly convex;
stomata paracytic, on both surfaces [mostly adaxial]; chlorenchyma not radiate.
Inflorescences terminal, sessile, capitate [diffusely branched]; bracts 1 or 2,
oblique to erect, sheathless or essentially so, basally expanded and partly clasp-
ing the spikelets, distally linear; rays lacking. Spikelets (1-)10-25, oblong-
ellipsoid, flattened, the 2 or 3 basal scales sterile; rachilla wingless, more or
less deciduous at maturity. Scales distichous. 3-8, oblong, acute but not mu-
cronate, distally scabrellate, laterally nerveless, medially 1-nerved. Flowers
perfect. Perianth bristles lacking to 6, smooth or scabrellate. Stamens 3; fila-
ments ribbonlike; anthers linear, the apices of the connectives subulate, con-
spicuous; pollen grains 4-porate, obovoid, finely scabrate (pore areas frustillate).
Styles trigonous to subtrigonous, glandular; stigmas 3, capillary, shorter than
the styles, glandular. Achenes roundly trigonous to subterete, ovoid to ellipsoid,
the apex broadly rounded, the base gradually tapered to a stipe, the surface
smooth or barely reticulate, glossy. Base chromosome number 20(?). TYPE
SPECIES: S. nigricans L.; see Britton & Millspaugh, Bahama Fl. 56. 1920. (Name
from Greek schoinos, for a rushlike plant.)- BLACK-HEADED SEDGE.
A genus of about 80 species, mostly restricted to Australasia but with a few
occurring in Africa, Eurasia, and the New World. Schoenus nigricans L., 2n =
54, 55, is present in North America. It is common in southern Florida but rare
in the Florida Panhandle, where it grows in wet grasslands over limestone
outcrops; it also occurs in the southwestern United States in the mountains
and valleys of western Texas, southern California, and southwestern Nevada,
where it grows in marshes and thermal springs. It is also reported from the
West Indies, Europe, and Asia.
417
1987]


JOURNAL OF THE ARNOLD ARBORETUM
Kiikenthal published a worldwide revision of Schoenus, and the genus has
received little subsequent systematic attention. The European species have been
investigated ecologically. Schoenus nigricans requires aluminum ions for growth.
and its range in the blanket bogs of the British Isles is thus limited to the coastal
region of western Ireland.
Plants of the genus have little economic significance. Wet meadows domi-
nated by Schoenus e/rru,'incus L. are mowed for fodder in northern and central
Europe. The species is adapted to low nutrient levels and is quickly displaced
by grasses when fertilizers are regularly applied.
RI FI RI NC's:
Under family references see BENTHAM; BERGGREN; C( ARKE (1908, 1909); GODFREY
& WooIN: N (CoNtAROV ('l al.: HARBORNI: HARIO)RNE (et a!. ;.. Hltr IC'INSo)N KtlKKONEN
(1986): KiNi H: Lt MAoi i & D C'AISNE: Li RMAN & RAx NAi.; Mi IeA FF;: NEs VON
Ei NB'CK; Sc'H mI z-Moei (1959, 1964): TORREY: and VANHECKE.
(GAN/Rr, (C., & J. PI ADE)NHAUER. Seasonal dynamics of shoot nutrients in Schoenus
i'rruriino'us (Cyperaceae). Holarctic Ecol. 9: 137-142. 1986. [Seasonal patterns of
biomass in an annually mowed calcareous fen in southern Germany: peak standing
crop in mid-July.]
Ki'K ENTHAI., G. Vorarbeiten iu einer Monographic der Rhynchosporoideae. Schocnus.
Repert. Sp. Nov. 44: 1-32, 162-195. 1938. [Worldwide monograph: keys, descrip-
tions; 83 species.]
SPARI INO, J. H. The occurrence of Schoe'nus nigr ians L. in blanket bogs. I. Environ-
mental conditions affecting the growth of S. ninricas in blanket bogs. Jour. Ecol.
55: 1-13. 1967a. II. Experiments on the growth of S. nirican.s under controlled
conditions. Ihid 14-31. 1967b.
WHEin :ER, B. D. An ecological study of .Schocnuo firrlugineus L. in Scotland. Watsonia
14: 249-256. 1983. [Autecology of a rare species.]
14. Cladium P. Browne, Civ. Nat. Hist. Jamaica. 114. 1756.
Stoloniferous, single-stemmed or loosely clustered, medium to large peren-
nials of sunny wetlands. Culms terete, roundly trigonous, or thickly crescen-
tiform, hollow, glabrous. Leaves all cauline: sheaths glabrous, much shorter
than the blades: blades flat or slightly conduplicate to subinvolute, the margins
and midveins sparsely scabrellate to harshly scabrous; chlorenchyma not ra-
diate: alternate bundles inverted. Inflorescences pedunculate, terminal or both
lateral and terminal, diffusely branched: bracts leaflike but with shorter blades:
primary rays terete, wirelike and slightly drooping, glabrous: secondary rays
similar to primary but shorter and more slender; tertiary and quaternary rays
regularly produced in some species, these subtended by lanceolate scalelike
bracts and sheathing prophylls (involucels). Spikelets in glomerules of 1-5.
narrowly ellipsoid to lanceolate; rachilla wingless. Scales 3-5, the basal 1-3
sterile, ovate to oblong-lanceolate. Flowers perfect or imperfect (the distal
flower of a spikelet perfect. the subdistal staminate). Perianth lacking. Stamens
2 or 3: filaments about as long as the subtending scale, flattened; anthers linear,
the apices of the connectives subulate: pollen grains 4-porate, narrowly obovoid
(sometimes with a peculiar apical appendage containing the degenerate nuclei).
scabrate. Styles subtrigonous, glabrous; stigmas 3, longer than the styles, glan-
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[voL. 68


TUCKER, CYPERACEAE
dular. Achenes terete, ovoid, the apex broadly round (the withered style base
sometimes persistent), the base truncate and impressed, sometimes stipitate,
the surface smooth or nearly so. Embryos small, broadly obovoid, scarcely
differentiated (the first leaf not developed). Base chromosome number 20. TYPE
SPECIES: C. Mariscus (L.) Pohl (Schoenus Mariscus L.; see Britton & Brown,
Illus. Fl. No. U. S. Canada, ed. 2. 1: 347. 1913). (Name from Greek clados,
branch, referring to the highly branched inflorescences.)-TWIG-RUSH,
SAW-GRASS.
Cladium is here accepted in the strict sense-i.e.. consisting of three species:
C. Mariscus, C. mariscoides (Muhl.) Torrey, and C. jamaicense Crantz. Ku-
kenthal treated the genus more broadly, including Machaerina Vahl. Recent
studies by Vanhecke and Metcalfe argue against such a broad circumscription.
Species of Cladium consistently differ from those of Machaerina in their small-
er, less differentiated embryos and their isobilateral leaves with inverted bun-
dles (illustrated by Metcalfe).
Two species occur in our area. Cladium jamaicense, the saw-grass of the
Florida Everglades, grows in tidal marshes and coastal wetlands from eastern
Virginia to Mexico and the West Indies. Some authors (KUkenthal, Raynal)
included C. jamaicense in the European C. Mariscus; Kern also included the
Australasian C. procerus S. T. Blake. The second species in our area, C. ma-
riscoides, occurs in brackish wetlands and inland fens and marshes from New-
foundland to Saskatchewan to Florida and Missouri; it is rare in the Southeast.
Raynal, without discussion, treated C. mariscoides and C. jamaicense as syn-
onyms of C. Mariscus, an extreme view not followed by anyone else.
Cladium jamaicense is important as the dominant species of much of the
Florida Everglades. The culms and leaves of C. Mariscus are gathered and
used in the manufacture of paper products in the Danube Delta, Romania.
REFERENCES:
Under family references see BEAL: BENTHAM: CLARKE (1908); CLIFFORD & HARBORNE;
ERDTMAN; EYIES & RouERTSON; FASSETT: GODFREY & WOOTEN; GOOD et al.; HARBORNE:
HARBORNE et al.: HETTruM; HOTC HKISs: G. E. HuITHINsoN: J. HUTCHINSON: J. H. KERN:
KUKKONEN (1969); LERMAN & RAYNAL: MEEUSE: METCALFE: SCHULZE-MOTEL (1959,
1964); TORREY; and VANHECKE.
CONWAY, V. M. Biological flora of the British Isles: Cladium Mariscus (L.) R. Br. Jour.
Ecol. 30: 211-216. 1942.
DEVILLEZ, F., & J. R. DESLOOVER. Influence de pretraitements chauds et froids sur
germination des graines de Cladium Mariscus. (English summary.) Bull. Soc. Bol.
Belg. 113: 45-58. 1980. [Warm followed by cold pretreatment gives best results.]
GUICHARD, A. Sur l'existence de faisceaux libero-ligneux a l'orientation inverse dans la
feuille vegetative de Cladium Mariscus P. Br. Compt. Rend. Acad. Sci. Paris 187:
509-511. 1928a. [Illustrations; description of inverted vascular bundles in leaf blades.]
Origine, parcours et torsion des faisceaux libcro-ligneux inverse du Cladium
Mariscus P. Br. Ibid. 567-569. 1928b. [Illustrations; basipetal differentiation of
vasculature in leaf blades.]
KOKENTHAL, G. Vorarbeiten zu einer Monographic der Rhynchosporoideae. XI. 10.
Cladium Crantz [sic]. Repert. Spec. Nov. 50: 1-17, 139-193. 1942. [Worldwide
revision of the genus in the broad sense: 47 species; C. jamaicense treated as sub-
species of Eurasian C. Mariscus.]
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JOURNAL OF THE ARNOLD ARBORETUM
Pi ADFNHAUER, J., & 1). ESKA. Untersuchungen /um Nahrstotlhaushalt eines Scheidried-
Bestandes (Cladietum marisci). Veroff. (ieobot. Inst. Riibel 87: 309-327. 1986.
[Ecological study; maximum aboveground biomass in September.]
RAYNAL, J. Notes cyperologiques 17. Revision des (adnium P. Browne s. lat. (C'yper-
aceae) de Madagascar et des Mascareignes. Adansonia. 11. 12: 103-112. 1972.
RuiDESc , L. The use of sawgrass for paper product manufacture: an examination of
properties. Pp. 191-195 in J. TOUHRifI :R & R. W. PIERsoN, JR., eds., Biological control
of water pollution. Philadelphia. 1976.
STEWAR), K. K. Physiological, edaphic, and environmental characteristics of typical
stands of sawgrass. Aquat. Ecol. Newsl. 9: 22, 23. 1976. [Tested for use in filtering
waste water: only 12 percent of phosphorus incorporated by plants; system saturated
after just eight weeks.]
--- & W. H. ORNES. The autecology of sawgrass in the Florida Everglades. Ecology
56: 162-171. 1975.
Subfam. CARICOIDEAE Pax, Bot. Jahrb. 7: 307. 1886.
Tribe SCLERIAE Kunth ex Fenzl in Endlicher, Gen. PI. 2: 114. 1836.
15. Scleria Bergius, Sv. Vet.-akad. Handl. 26: 142. 1765.
Small to medium, erect [scandent], perennial or annual herbs of grasslands,
open woods, fens, and shores. Roots fibrous; rhizomes regularly present in
many species, indurate, sometimes tuberlike, simple or branched. Culms trigo-
nous, glabrous, pubescent, or scabrellate [retrorsely scabrous], sometimes bul-
bous basally. Basal leaves bladeless or nearly so. Cauline leaves several; sheaths
3-angled, glabrous or more often scabrellate or pubescent; blades lanceolate to
linear or filiform, flat to slightly conduplicate [involute or thickened], glabrous,
scabrellate, or pubescent; chlorenchyma not radiate. Inflorescences paniculate,
I to several, terminal or lateral and terminal; bracts leaflike but shorter than
or equaling the cauline leaves; rays trigonous, scabrellate on the angles or
smooth, secondary rays regularly produced in some species. Spikelets 1-6,
lanceolate to linear or oblong. Scales 1-6. ovate-deltoid, acute, mucronulate
to cuspidate, conspicuously medially 1-nerved, laterally nerveless, glabrous or
pubescent. Flowers imperfect; carpellate flower(s) 1 (or 2), borne at the base
of the spikelets or in separate spikelets. Perianth bristles lacking. Stamens 1-
3; filaments capillary; anthers narrowly ellipsoid to linear, the apices of the
connectives frequently prolonged as slender, subulate, reddish appendages;
pollen grains uniaperturate, obovoid to subspheroid, psilate. Hypogynium, if
present, pebbled or warty, entire or with 3 acute to obtuse [truncate or acu-
minate], ciliate or glabrous lobes clasping the base of the achene. Styles slender,
glandular; stigmas 3, capillary, shorter than the styles. Achenes roundly tri-
gonous to terete, globose to ellipsoid, the apex broadly rounded (sometimes
apiculate), the base sessile to broadly stipitate, the surface smooth, reticulate,
trabeculate, rugose, glabrous, or pubescent. Base chromosome number 7(?).
TYPE SPECIES: S. jlagellum-nigrorum Berg.: see Britton & Brown, Illus. Fl. No.
U. S. Canada, ed. 2. 1: 348. 19 13. (Name from Greek skleros, harsh, the culms
of the type species being bound together into whips for beating slaves in Sur-
inam; often incorrectly said to be derived from Greek skleria, tough, in reference
to the achene walls; see Holm, 1898).-NtrT-RUSu.
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A predominantly tropical genus of some 200 to 225 species. Centers of
diversity are tropical South America, tropical Africa, and southeastern Asia.
Twelve species occur in the United States, all east of the Great Plains. All are
present in the Southeast. Several range northward into northeastern North
America, reaching Massachusetts, southern Ontario, and southern Minnesota.
Two of our representatives occur southward into the West Indies. Many of our
species are endemic, as are most other taxa of Scleria. Many African species,
for example, occur only in Africa, and several are restricted to a single country
or are known from only one collection. Such endemism contrasts with the
distribution of the other large, mostly tropical genera of the family, such as
Cvperus, in which about one-fifth of the species are pantropic. Only two species
of Scleria, C. lithosperma (L.) Sw. and S. hirtella Sw., are reported from both
the Old World and the New.
The morphology of the achenes and the hypogynia has traditionally provided
the chief criteria for the circumscription of species. Core noted that some
species-for example, the South American Scleria leptostachya Kunth-pro-
duced both smooth and verrucose achenes, sometimes within a single collection
and sometimes within the same inflorescence. Nelmes (1955, 1956) reported
similar problems with certain African species, and he relied on features of the
rhizomes, ligules, and inflorescence (in addition to achene morphology) in his
classification of the African species. The hypogynium is apparently derived
from receptacular tissue, as is shown by its vascularization (Blaser, 1940, 1941b).
Robinson (1966) indicated that many of the southern African species of
Scleria are strong calcicoles. This autecology contrasts with that of the Amer-
ican species, most of which grow in acidic coastal plain habitats. Apparently
only one American species, S. nitida Willd. (which Fairey treated as a synonym
of S. verticillata) is a calciphile (Fernald).
Core recognized five sections in the genus, of which two, sects. SCLERIA (sect.
Euscleria Endl.)and HYPOPORvM (Nees) Endl., are represented in the Southeast.
In sect. HYPOPORUM the species have androgynecandrous spikelets (carpellate
flowers below the staminate) and lack hypogynia. There are five species in our
area: S. verticillata Willd., S. hirtella Sw.. S. Baldwinii (Torrey) Steudel, S.
georgiana Core, and S. lithosperma (L.) Sw. Species of sect. SCLERIA have
unisexual spikelets and three-lobed, entire hypogynia. In our area this section
includes seven species: S. triglomerata Michx., S. minor Stone. S. oligantha
Michx., S. ciliata Michx., S. pauciflora Willd., S. Curtissii Britton, and S.
reticularis Michx.
The genus is scarcely known cytologically. Reports are available only for
Scleria tesselata, 2n = 28, of southeastern Asia. This suggests the base number
x = 7 for the genus.
Species of Scleria have unusual embryological features (Nijalingappa). In S.
foliosa A. Rich. the embryos have both chalazal and micropylar haustoria.
Wall formation in the endosperm is complete in the Cyperaceae. except in
Scleria, where it is incomplete. The surface of the cotyledon is papillose in
Scleria but smooth in other genera of the family.
Robinson (1966) stated that several southern African species had "citrus-
scented" foliage: in fact, he used this as a lead characteristic in his key. Thus,
further investigation of the chemistry of these plants might be fruitful.
421
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JOURNAL OF THE ARNOLD ARBORETUM
The fruits of Scleria triglomerata are dispersed by ants. The hypogynium
functions as an elaiosome (Gaddy). However, Robinson (1962) suggested that
the hypogynium provided buoyancy for the achenes of several southern African
species and was thus an adaptation for dispersal along water channels that
might later provide appropriate conditions for germination and growth of
seedlings.
No species of Scc'ria is gathered for food. Rhizomes of S. hirtella have been
employed medicinally in Colombia (Core). The tough, scabrous foliage of Sc/e-
ria is unsuitable for cattle forage. About ten species are noted as significant
weeds in Central and South America, tropical Africa, and southeastern Asia.
Scleria sumatrensis Retz. is a detrimental weed in Borneo (Holm et al.).
REt iERI N( TS:
Under family references see BEINTHAM: BLASI.R (1940, 1941b): CLARKE (1908. 1909):
Cl IFI()RI)o EITEN (1976a); FERNAIii): GA)I)Y; HARBORNE: HARBORNF et at.: HotiM et al.:
Hoi iiUM: Hi AN: J. Hi I('HINSON: J. H. KERN: KOVAMA: KUKKONEtN (1969): L: MAOI:I
& DE('AISNi:: LRMAN & RAYNAL; Ml:t sE: MiliAI.E:; NAPPER (1964b): O'NEitL:
ScHIilze-MOTEL (1959. 1964): STANDIt)Y: TiiRi e al.: and TORRiY.
CORE, E. L. The American species of.Sclcria. Brittonia 2: 1-105. 1936. [Basic monograph
for the New World species: keys, descriptions, representative specimens; illustrations
of achenes.]
FMIREY, J. E., III. The genus S'c'ria in the southeastern United States. Catanea 32: 37-
71. 1967. [Keys, descriptions, distribution maps: taxonomy closely follows CORE.]
HO LM, T. Studies in the Cyperaceae. VII. On the inflorescence of the genus Scleria. Am.
Jour. Sci. 155: 47-52. 1898; VIII. On the anatomy of some North American species
of Scleria. Ibid. 157: 5-12. 1899. [Discussion of several southeastern species: illus-
trations.]
N 1 M :S, E. Notes on Cyperaceae: XXX V II. Scleria Berg. sect. IIHpoporum (Nees) Endl.
in Africa. Kew Bull. 10: 415-453. 1955; XXXIX. African species ofScleria excluding
sect. Ilipoporum. Ibid. 11: 73-111. 1956. [Keys. descriptions, discussions, specimen
citations; illustrations of many species.]
NUAiiN(GAPPA, B. H. M. Embryology of Sclera /oliosa ((yperaceae). PI. Syst. Evol. 152:
219-230. 1986. [Illustrations.]
ROBINSON, E. A. Notes on Scleria: I. The African species of sect. Tesselatae. Kirkia 2:
172-192. 1961: III. Scleria lirtl/la and some allied species. Ibid. 4: 175-184. 1964.
- -. Scleria in Central Africa. Descriptions and notes: II. Ibid. 3: 8-14. 1962.
A provisional account of the genus Seleria Berg. (Cyperaceae) in the "Flora
Zambesiaca" area. Kew Bull. 18: 487-551. 1966. [Keys, descriptions, illustrations.]
Tribe CARICEAE Kunth ex Dumortier, Fl. Belg. 144. 1827.
16. Cymophyllus Mackenzie in Britton & Brown, Illus. Fl. No. U. S. Can. ed.
2. 1: 441. 1913.
Loosely caespitose perennials of mesic montane forests. Rhizomes oblique.
Culms subterete, smooth, aphyllopodic. Leaves several; lowest with papery
sheath only, bladeless; uppermost sheathless, the blade broadly lanceolate,
broadly rounded at apex, undulate at margins (especially so when dried), con-
spicuously multinerved but lacking a differentiated midvein and ligule. Inflo-
rescences single densely ellipsoid spikes. I per culm, terminal, with the pistillate
flowers below the staminate: bracts single broadly deltoid entire scales, 1 per
422
[VOL. 68


TUCKER, CYPERACEAE
i7
\~t t
\ n'rs n just
FIouRE 4. Cvmophyvllus. a-n, C. Fra.seri: a. habit (portion of plant, leaf of preceding
season, plus new shoot terminated by inflorescence just past anthesis), x '/; b, detail of
undulate leaf margin, x 6; c, longitudinal section of inflorescence, staminate flowers
above, carpellate below, x 2; d, staminate flower with subtending scale, x 3; e, anther
(basilixed). x 12; f, 3 carpellate flowers enclosed in perigynia, each in axil of a scale,
x 3; g, longitudinal section of perigynium to show carpellate flower (note bristlclike
rachilla), x 5: h. stigma (note lack of papillae-species is insect pollinated), x 12; i,
longitudinal section of gynoecium to show single basal anatropous ovule, x 12; j, early
stage ofdeveloping fruit, growth of gynoecium producing kink in style, x 5: k. perigynium
enclosing mature achene, x 6; I, immature achene (note rachilla at base), x 6; m, achene,
x 6; n, embryo, dissected from base of achcne, 2 views, x 25.
spike, broader than but otherwise like the pistillate scales immediately above
it. Flowers imperfect. Perianth lacking. Scales oblong-ovate, entire, without
conspicuous midvein or nerves. Stamens 3: filaments slender, 1-3 times as
long as the subtending scales; anthers slenderly ellipsoid, the connectives not
423
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JOURNAL OF THE ARNOLD ARBORETUM
prolonged. Perigynia 10-30, broadly ellipsoid, roundly trigonous, abruptly con-
tracted to a short, entire beak, weakly 20- to 30-nerved. glabrous: rachilla
filiform, A-/: as long as the perigynium. Styles slender: stigmas 3, slightly
longer than the style, exserted from the beak of the perigynium. Achenes trigo-
nous, broadly ellipsoid, the apex broadly rounded, the base abruptly stipitate,
the surface smooth, glossy. Chromosome number unknown. TYPE SPECIES: C.
Fraseri (Andrews) Mackenzie ((Carex Frascri Andrews: see Britton & Brown,
Illus. Fl. No. U. S. Canada, ed. 2. 1: 441. 1913.) (Name from Greek kuma,
wave, and phl//on, leaf, in reference to the undulate margins of the leaves.)
- FRASER'S SEDGE.
A monotypic genus endemic to the southern Appalachians. The sole species,
Cy'mophy'llus Fraseri. is well known for its attractive white spikes that are
conspicuous when the plants flower in the spring. The plants grow in mesic to
somewhat damp soils in mixed hardwood forests, particularly on northern and
western slopes at middle elevations. The species is known from eastern Ten-
nessee and northwestern South Carolina, north through the Ridge and Valley
and Blue Ridge provinces to extreme south-central Pennsylvania (Somerset
County). Clarkson listed known collections arranged by state and county.
The systematic position of the genus has been disputed. Kukenthal treated
the species as C'arcx lFraseri (sect. Leucocephali Holm of subg. Primaocarex
Kiikenthal). Mackenzie, Fernald, Metcalfe, and Reznicek (pers. comm.) rec-
ognized Cymophyll/us as a distinct genus. The conspicuous white inflorescences
of('. F'raseri, while unique among North American species of the tribe Cariceae,
are also known in at least one Old World species of Care (C(. ha/densis L.).
White inflorescences are associated with insect pollination (discussed below)
and have evolved in Crperus and Rhynchospora. In ('. Fraseri there is a rachilla
within the perigynium. While a rachilla is not present in any temperate North
American species of Carex, it does occur in several other species (e.g., C.
micro'lochin Wahlenb. (boreal North America, cold-temperate Eurasia, south-
ern South America, tide Fernald)). Anatomical evidence (summarized by Met-
calfe) gives the strongest support for the generic status of Cymophll/us. In
Cymophy/llus Fraseri the culms are terete (trigonous (rarely hexagonal) in Car-
ex): the leaves lack ligules (which are always present in Carex): the uppermost
leaflacks a sheath and consists of blade only (sheaths are always present in the
cauline leaves of (Carcx): the large leaf blade is broadly rounded apically (acute
in ('arc') and lacks the differentiated midrib and the adaxial laver of bulliform
cells typical of Carex (Holm: Metcalfe). In (l'rmophyllus Frascri the median
vascular bundle has an incomplete adaxial sclerenchyma cap. and there is an
abaxial sclerenchyma girder (Metcalfe). The presence of perigynia in Carex
and ( Cmophyllus clearly indicates that they are closely related, although it is
unclear how. The presence ofa rachilla in Cy'mophyll/us suggests that this genus
might be closer to the Southern Hemisphere Lncinia Pers. than to Carex.
('vmnophyllus Fraseri has long been suspected of being entomophilous (Clark-
son), although there has been only a single field study documenting entomophily
(Thomas). Four bee and one fly species were observed to visit spikes of this
species, which flowers from late April to mid-June. The insects collect pollen
424
[vol.. 68


for food and transfer it from plant to plant. They land on the lower, relatively
broad carpellate portion of the spikes, where they deposit pollen on the stigmas.
They then crawl up to the anthers, collect pollen, and fly to another inflores-
cence. The pattern of stigmas first, then anthers, probably enhances outcrossing
(Thomas).
REFERENCES:
Under family references see BENTHAM, CLARKE (1908), FERNALD, METCALFE,
SCHULZE-MOTEL (1964), and TORREY.
CLARKSON, R. B. Fraser's sedge, Cymophyllus Fraseri (Andrews) Mackenzie. Castanea
26: 129-136. 1961. [Ecology: summary of literature and known distribution.]
HOLM, T. Studies in the Cyperaceae, III. Carex Fraseri Andrews, a morphological and
anatomical study. Am. Jour. Sci. IV. 4: 121-128. pl. IV. 1897. [Detailed description
with taxonomic comments.]
HORN, G. S. VAN, & L. G. WILLIAMS. New county records for endangered and threatened
species in Tennessee. Castanea 46: 343-345. 1981. [C. Fraseri in Polk Co.]
JOHNSON, R. H., & J. W. WALLACE, JR. The flavonoid profile of Cymophyllus Fraseri
(Cyperaceae). (Abstract.) Am. Jour. Bot. 73: 727, 728. 1986. [Contains methylated
apigenins.]
KOKENTHAL, G. Cyperaceae-Caricoideae. In: A. ENGLER, ed., Pflanzenr. IV. 20(Heft
38): 1-824. 1909.
MACKENZIE, K. K. Cyperaceae: Caricoideae. N. Am. Fl. 18(2, pts. 1-7): 1-478. 1931-
1935. [The standard monograph: descriptions and keys for 533 species.]
RAYNER, D., et al. Native vascular plants: endangered, threatened, or otherwise in
jeopardy in South Carolina. So. Carolina Mus. Bull. 4. 22 pp. 1979. [C. Fraseri
extirpated in South Carolina.]
SIMS, J. Carex Fraseriana. Fraser's carex. Bot. Mag. 33: no. 1391. 1811. [C. Fraseriana
Sims, a synonym of C. Fraseri Andrews.]
THOMAS, W. W. Insect pollination of Cy:mophyllus Fraseri (Andrews) Mackenzie. Cas-
tanea 49: 94, 95. 1984.
17. Carex Linnaeus, Sp. PI. 2: 972. 1753; Gen. Pl. 280. 1754.
Caespitose or single-stemmed, small to medium-sized perennials of wet to
dry woods, grasslands, rock outcrops, pocosins, fens, bogs, marshes, and swamps.
Roots fibrous, smooth or pubescent; rhizomes (infrequently lacking) short and
oblique or long and horizontal, with closely appressed, lanceolate scales. Culms
loosely to densely clustered or solitary, fertile or both vegetative and fertile,
trigonous [hexagonal], the angles smooth or scabrellate. Basal leaves several to
many; sheaths smooth; ligule hyaline. glabrous; blades flat, conduplicate, pli-
cate, or involute, scabrellate (or smooth) on margins and midveins, sometimes
microscopically papillate on 1 or both surfaces, infrequently glaucous; stomata
paracytic, present on one or both surfaces; chlorenchyma not radiate; air cham-
bers frequently present; cauline leaves similar to basal ones but shorter and
fewer, sometimes lacking. Inflorescences simple or compound, monoecious
(rarely dioecious); bracts lacking or 1-6; spikes 1 to several, loosely to densely
ovoid to slenderly cylindrical, sessile or borne on simple [branched] erect to
pendent peduncles; each spike subtended by a leaflike or filiform basal bract;
spikes wholly carpellate or wholly staminate or gynecandrous or androgynous.
TUCKER, CYPERACEAE
425
1987]


JOURNAL OF THE ARNOLD ARBORETUM
{x
a K
" t{I�
'S
I
\t K
V
j
xYK.
K
U
FI-R, lu5. ( Caret subg. VI(,NI A: 8 species shown, each representing a dillfferent section
(A. ( Carc retro/llea (sect. Pi. sitol o IIN): B, ('. vu( lpinoidea (sect. Mt ninn OR A,): (',
(' (Ico 1ipousiti (sect. HI El E I)( ( IN); I), ('1 lue''ivta inataw (sect. V I 'IPINAE): E, (C. bran-
m"e ve' subsp. sphaero.stachla (sect (i.AREosAi ); F, I(. bromoidet (sect. DEWEYANAE):
426
[vol.. 68
\
I i �l
I"� li
(�t
B� ff.
ipt
i 1 t 1
!
,:
'..
j
: � I


TUCKER, CYPERACEAE
Staminate scales lanceolate (the margins rarely fused basally), hyaline to char-
taceous, 1- (to 3-)nerved; carpellate scales lanceolate to broadly ovate, char-
taceous, 1 - (to 3-)nerved. Flowers imperfect, protogynous or protandrous. Peri-
anth lacking. Stamens 3; filaments capillary or ribbonlike, longer than the
subtending scales; anthers broadly to slenderly ellipsoid; pollen grains 1- or
4-aperturate, obovoid or subspheroidal, psilate, trinucleate. Perigynia solitary
in the axils of carpellate scales, lenticular, subterete, trigonous, or slightly to
strongly compressed (beak, when present, less than to equaling or sometimes
longer than the body), coriaceous to chartaceous, the faces nerveless or with
1-15 nerves, minutely papillose or not, scabrellate or essentially smooth, dull
or glossy. Styles capillary, straight or curved; stigmas 2 or 3 [or 4], equaling or
exceeding the styles in length, smooth, papillose, or glandular, at anthesis
exserted through the orifice of the perigynia. Achenes lenticular or trigonous
[4-sided], ovoid to ellipsoid, 1/4 as long as to nearly as long as the body of the
perigynium, sessile or stipitate, apiculate or entire, the faces flat, convex, or
concave, the edges obtuse or acute (invaginate in a few species), the epidermal
cells translucent, opaque, or glossy. Embryos obconical, the radicle basal. Base
chromosome number 5. TYPE SPECIES: C. hirta L., not C. pulicaris L.; see
Hitchcock & Green, Prop. Brit. Bot. 187. 1929, and comments by Voss, Mich.
Bot. 11: 31, 32. 1972. (The classical Latin name, perhaps derived from the
Greek keirein, to cut, due to the sharp margins and keels of the leaf blades.)
- SEDGE.
A very large, cosmopolitan genus, reported to contain from 1000 to 2000
or even 2500 species (Standley, 1985a), including 165 that occur in the South-
east. Four subgenera have been recognized, of which two are represented in
the United States. Subgenus INDOCAREX Baillon (inflorescences richly branched,
branches subtended by tubular prophylls) comprises about 50 species of the
Old World tropics. Subgenus VIGNEA (Lestib.) Kukenthal (spikes all either
gynecandrous or androgynous, sessile, stigmas two, perigynia and achenes len-
ticular) includes about 500 species; it is worldwide in distribution but is most
diverse in the northern temperate and boreal regions. Subgenus CAREX (subg.
Eucarex; spikes sessile or pedunculate, some exclusively staminate or pistillate,
stigmas 3 (rarely 2), perigynia and achenes trigonous) is the largest subgenus.
with about 800 species. Subgenus PRIMOCAREX Kiukenthal (spikes solitary,
terminal, stigmas 2 or 3, achenes lenticular or trigonous) is not represented in
our area.
The evolution of the tribe Cariceae is largely unclear. Due to shared features
of the inflorescences, Smith & Faulkner suggested that it arose from ancestors
akin to the Scleriae or the Hypolytreae. Kukkonen (1963). because of similar
G, C. Hfowei (sect. STELLULATAF): H, C. scoparia (sect. OVALES)). Four or 5 items illus-
trated for each, these drawn at same magnification throughout: 1, inflorescence, x 1: 2,
scale subtending perigynium, abaxial surface, x 10; 3. mature perigynium, abaxial sur-
face, x 10; 4, mature achene, abaxial surface. x 10; 5, longitudinal section of mature
perigynium and achene (C and D only), x 10.
427
1987]


JOURNAL OF THE ARNOLD ARBORETUM
' ,, ) 9/I
) ;n .'
FIuR 6. (arcAs subg. CARIx (subg. 1 ucarex). a-i. ( gtganIca: a, inflorescence,
uppermost 3 spikes staminate, x :: b, staminate flower and subtending scale, adaxial
view, 5: c, perigynium in axil of subtending scale, stigmas of carpellate flower pro-
truding, 5: d. carpellate flower (gynoecium), perigynium removed, x 5: e, ovule, lateral
view, micropylc not visible, x 25: t, mature perigynium enclosing achene, x 5; g. achene
with persistent style, , 5: h, longitudinal section ofachene, seed coat not shown, embryo
basal, endosperm above, x 5: i, seed removed from achene. x 5. j-r, (' . g/aucescens. j,
minlorescence, staminate spike uppermost, ': k, staminate flower and subtending scale.
most of2 stamens removed. x 5:1, perigynium and subtending scale, stigmas of carpellate
flower protruding, x 5: m. carpellate flower, 5 n, ovule, micropyle visible, raphe
behind, x 25: o, mature perigynium enclosing achene, x 5: p, perigynium. detail of
surface, showing globular to ellipsoid cells that produce glaucous effect, x 25: q, achene,
x 5: r, embryo, x 25.
428
[VOL. 68


infestations of smut fungi, indicated a probable close relationship with subfam.
Rhynchosporoideae; Koyama concurred with this opinion. A clearer under-
standing of generic relationships of the genus must await a better picture of
evolution within the genus. Its very large size and worldwide distribution
continue to hamper such studies.
Kfukenthal believed that subg. PRIMOCAREX Kiikenthal was the most prim-
itive within the genus. A succession of more recent cyperologists (Kreczetowicz;
Nelmes; Koyama, 1962a; Le Cohu, 1968; Haines & Lye; Smith & Faulkner;
Reznicek, 1986b) have taken the opposite view. In their opinion the unispicate
condition of subg. PRIMOCAREX was derived (perhaps polyphyletically) from
ancestors with richly branched inflorescences like those of subg. INDOCAREX.
However, the presence of a rachilla within the perigynium of some species of
subg. PRIMOCAREX suggests that it is the most primitive subgenus. Smith &
Faulkner believed that subgenera CAREX and VIGNEA might have evolved from
subg. INDOCAREX by reduction in inflorescence structure (a pattern also sug-
gested for several other genera of the family, e.g., Cyperus and Scirpus). This
would have involved loss of cladoprophylls (tubular prophylls subtending
branches) and reduction of branching. There are contrasting interpretations of
the inter- and infrageneric relationships in Carex.
The morphology of the inflorescences, particularly of the spikes and peri-
gynia, has traditionally been most heavily relied upon in distinguishing species
and circumscribing sections. Anatomical and cytological features are also tax-
onomically useful. Anatomical evidence has long been applied to the system-
atics of Carex. Crawford described the stems and leaf blades of the British
species. Akiyama presented a systematic study of the eastern Asian species,
emphasizing anatomical differences. Several recent revisions have included
anatomical descriptions of culms and leaves. Standley (1985a), in her mono-
graph of the northwestern species of sect. PHACOCYSTIS Dum. (sect. Acutae),
showed that related species differ in the distribution of sclerenchyma and sto-
mata in culms and leaf blades. In certain species stomata are present on one
or both surfaces (Standley, 1986). The importance of anatomical features has
also been discussed by Le Cohu (1972) and by Metcalfe.
Recent studies with the scanning electron microscope have revealed an in-
teresting variety of surface features in leaves, perigynia, and achenes of Carex.
The presence of tubercles (Hoshino, 1986) and papillae (Maloney & Evans)
and the distribution of stomata (Standley, 1986) are useful in distinguishing
species and circumscribing sections.
Cytological studies have been helpful in Carex, but chiefly at the specific
level. Chromosome numbers in the genus range from n = 6 to n = 56. The
base chromosome number is 5, and the commonest haploid numbers in North
American species are 10, 20, 30, and 40 (Wahl). In many instances related
pairs of species differ in chromosome number. Aneuploidy is prevalent within
the genus. Aneuploid series characterize many sections (Wahl; Davies; Dietrich;
Faulkner, 1972). Polyploidy is infrequent.
The pollination biology of Carex has received little attention. Most species
are anemophilous. Honey bees and beetles visit inflorescences to gather pollen
TUCKER, CYPERACEAE
429
1987]


JOURNAL OF THE ARNOLD ARBORETUM
and thus may also be vectors (Leppik). Self-compatible and self-incompatible
species have been noted in the genus (Faulkner, 1973; Handel, 1976, 1978a:
Schmid, 1984b). It is not known whether the incompatibility is sporophytic
or gametophytic. Handel (1976) determined that pollen-flow distances in C.
platyphyvla Carey and C. plantaginea Lam. were rarely more than 10 m.
Little is documented about the dispersal of fruits of Carev. It has been
assumed that species with inflated perigynia are dispersed by floating on water,
but experimental verification is lacking. Several North American species (e.g.,
C(. communis Bailey, C. umbhcllata Willd., and (C. pedunculata Willd.) have
elaiosomes at the base of the perigynia and are dispersed by ants (Handel,
1978; Gaddy. 1986). Carev pauci/lora Michx., widespread in northeastern
North America, has subulate perigynia that at maturity spring away from the
rachis (up to 60 cm) when touched (Hutton).
Flavonoid profiles can be used to distinguish between closely related species.
Toivonen (1974) showed this in the Fennoscandian representatives of sect.
('ANESCENTES (sect. HIeleonastes). Manhart (1985) demonstrated that classifi-
cations based on occurrences of flavonoids were similar to relationships de-
termined by morphology.
The species of Carex fall into three broad ecological groups with regard to
habitat: wetland, forest, and ruderal. In general the species of a section are
ecologically similar. Several sections (e.g., sects. PALUDOSAE G. Don and
LI PULINAE Carey) include mostly wetland species. Section ACROCYSTIS Du-
mort., however, contains species of dry to dry-mesic open or wooded habitats.
Several sections (e.g., sect. AInBAE Ascherson & Graebner) are composed mostly
of calcicoles.
Most species of Carex are rhizomatous perennials. Carex is the only large
genus of the family containing no annuals. Certain species reproduce mostly
vegetatively (e.g.. C. Bigelowii Torrey. plants of which set abundant seed, with
little germination or recruitment of seedlings unless disturbance occurs). In the
boreal C. /lava L. seedlings persist for several years until competition is removed
(by disturbance or herbivorv) and then grow rapidly to fill in the available
space (Schmid, 1986).
The economic importance of the genus lies chiefly in providing fodder for
domestic and wild mammals, especially in colder regions. Many Russian species
are important in this way (Goncharov et al.): Carex stans Drejer and C. discolor
Nylander provide good grazing for cattle and reindeer. In Iceland, meadows
of C. lIynghyei Hornem. are managed and yield up to five tons per hectare.
The nutritional content is very similar to that of common pasture grasses such
as Kentucky bluegrass, Poa pratensis L.
The following is a synopsis of the southeastern species, with chromosomal,
systematic. and ecological references. The order and circumscription of sections
generally follows Mackenzie (1931-1935).
Subgenus VI(NEA (Lestib.) Kiikenthal, represented in the Southeast by species
belonging to ten sections, is characterized by lenticular achenes, dorsiventrally
flattened perigynia, two stigmas, and both carpellate and staminate flowers in
each spike of the inflorescence.
Species of sect. AMMOGLOCHIN Dumort. ( lrenariae Kunth, including sect.
430
[VOL. 68


Divisae) are small rhizomatous plants of grasslands and strands. Two Eurasian
species, Carex arenaria L., n = 29. 58, 60, 64 (Noble) and C. divisa Hudson,
are naturalized in our area. Both grow on coastal sands from eastern Maryland
to eastern North Carolina. Several others occur in Canada and the western
United States, where C. Eleocharis Bailey is an important forage in the Rocky
Mountain region (Hermann. 1970).
Section MACROCEPHALAE Kukenthal comprises two eastern Asian species,
one of which, Carex Kobomugi Ohwi, 2n - 84, 88, is sparingly naturalized
from eastern Virginia (Norfolk Co.) north to Cape Cod; it should be looked
for in eastern North Carolina. Standley (1985b) studied its population biology.
Although previous authors had described the species as dioecious, she showed
that individual rhizomes of a clone were consistently either staminate or car-
pellate (monoecious).
Section PHAESTOGLOCHIN Dumort. (sect. Bracteosae (Kunth) Pax) is one of
the most diverse sections of Carex in North America; it includes 16 species in
our area, all with ranges that extend into the northeastern United States or to
Canada. Plants of these species are mostly caespitose, with one to five sessile
androgynous spikes. Webber & Ball revised the C. rosea complex and corrected
the application of the names C. rosea and C. convoluta. Chromosome numbers
are known for six southeastern representatives of this section: C. sparganioides
Muhl., n = 23; C. cephalophora Muhl. ex Willd.. n = 24; C. retroflexa Willd.,
n = 20; C. rosea Schkuhr (C. convoluta Mack.), n = 26; C. appalachica Webber
& Ball (C. radiata auct.. non (Wahlenb.) Sm.); and C. radiata (Wahlenb.) Small
(C. rosea auct., non Schkuhr), n = 29. David & Kelcey summarized the biology
of the European species. C. muricata L., C. spicata Hudson, and C. divulsa
Stokes, all 2n = 58. Carex spicata and C. divulsa are naturalized in the North-
east south to Virginia. They might be found in North Carolina.
Section MULTIFLORAI (Kunth) Mack. contains three species in our area. The
commonest of these, Carex vulpinoidea Michx.. n = 26, 27, is known from all
of the southeastern states and ranges north into southern Canada. It is also
sparingly naturalized in England (Clapham et al.). The other southeastern species
are C. triangularis Boeck. and C. anneciens Bickn. Both occur in most of the
southeastern states, but neither is as common as C. vulpinoidea.
Section HELEOGLOCHIN Dumort. (sect. Paniculatae G. Don (Hort. Brit. 367.
1830; non Carey) is represented in the Southeast by Carex decomposita Muhl.,
n = 30, 32, 33, which occurs in every state in our area. Plants of this section
are the only North American representatives of C('arex with paniculate inflo-
rescences. Certain extraregional species of the section appear to be cytologically
conservative (cf. Clapham et al.). Carex diandra Schrank, 2n = 60, is circum-
boreal, while C. paniculata L., 2n = 60, 62, 64, and C. appropinquata Schum.,
2n = 64, are European.
In the Southeast, sect. VULPINAE (Carey) Christ is represented by five species
of swamps, marshes, and wet meadows. The plants resemble those of the
preceding two sections but are distinguished by their long, slender perigynia
(1 cm long in Carex crus-corvi Shuttlew.). In several species the bases of the
perigynia are conspicuously enlarged with aerenchyma, which probably makes
the fruits buoyant and allows dispersal by water. Carex crus-corvi, n = 26, C.
TUCKER, CYPERACEAE
431
1987]


JOURNAL OF THE ARNOLD ARBORETUM
laevivaginata (Kukenthal) Mack., n = 23, and C. slipata Willd., n = 26, occur
throughout our area.
Section GLAREOSAE G. Don (sect. IHeleonastes (Kunth) Kukenthal) is a group
of circumboreal species of wet woods and bogs. The plants are small and have
few-flowered inflorescences. Three species, Carex brunnescens subsp. sphae-
rostachya (Tuckerman) Kalela, n = 27, 28, C. canescens L., n = 27, 28, and
C. trisperma Dewey, n = 30, barely reach our area from the north and are
found in the mountains of North Carolina and Tennessee.
Section STELLULATAE (Kunth) Christ consists of perhaps 30 species world-
wide. The plants are caespitose and have gynecandrous spikes of spreading to
rellexed perigynia with serrulate beaks. Reznicek & Ball (1980) revised the
North American species and provided excellent keys and descriptions. There
are seven representatives in our area. Carex Ruthii Mack. is endemic to high
elevations in the southern Appalachian Mountains from West Virginia to Geor-
gia. Carex exilis Dewey is primarily northeastern, occurring from Newfound-
land to Ontario south to Maryland: it is also known from widely disjunct
stations in central North Carolina, southern Mississippi, and southern Ala-
bama. The other southeastern species are C. atlantica Bailey (including C.
Xlohriana Mack.), C. llowei Mack., n = 27, C. incomperta Bickn., n = 22.
and C. angustior Mack., n = 26. The two European species for which counts
are available have similar numbers: C. elongata L., 2n = 56, and C. echinata
Murray, 2n = 56, 58.
Species of sect. DEWEYANAE (Tuckerman) Mack. are probably closely related
to those of sect. STELLULATAF but have fewer, narrower, and appressed rather
than spreading perigynia (Reznicek & Ball, 1980). Carex bromoides Schkuhr,
n = 31 +(4), is the sole southeastern representative; it occurs in every state in
our area. Carex Dewevana Schwein., the only other species of the section, occurs
in northeastern North America.
Section OVALES (Kunth) Christ contains about 50 species in North America.
It is the largest section in our area, and the 16 representatives occurring in the
Southeast have flattened, papery, appressed perigynia in dense, ovoid spikes.
The section is taxonomically difficult and needs revisionary work. Several taxa
recognized by Mackenzie (1931-1935) have been synonymized by later work-
ers. Most of our species are widespread in eastern North America. For example,
Carex tribu/oides Wahlenb., n = 35, and C. reniformis (Bailey) Small occur in
all the southeastern States, while C. argyrantha Tuckerman and C. aenea Fern.
are northeastern and just enter our area in the mountains of North Carolina.
C'arex vexans Herm. is endemic to central and southern Florida. Among our
representatives, chromosome numbers are known only for C. tenera Dewey
(n = 26, 27, 28), C. straminea Willd. (n = 34 t(3)), and C. cristatella Mack.
(n = 35). The type species, the European C. ovalis Good., 2n = 64, 66, 68, is
cytologically similar to eastern North American species of the section.
Subgenus CAREX (subg. Eucarex Cosson & Germ.) includes the remaining
sections of the genus, 26 of which are represented in the Southeast. The plants
are characterized by differentiated spikes in which the terminal spike is wholly
staminate and the others are wholly or partly carpellate. Except in the distig-
[VOL. 68
432


TUCKER, CYPERACEAE
matic sect. PHACOCYSTIS Dumort. (sect. Acutae), the ovaries and achenes are
trigonous and there are three stigmas.
Section POLYTRICHOIDEAE (Tuckerman) Mack. contains only Carex leptalea
Wahlenb., n = 26, an eastern North American endemic growing in damp,
mossy woods, often in calcareous soils, from Florida and eastern Texas north
into Canada. These are small, thin plants bearing few slender, beakless perigynia
and narrowly oblong, truncate achenes.
Section PHYLLOSTACHYAE (Tuckerman) Bailey has four North American
species, characterized by androgynous spikes and staminate scales with basally
fused margins. Two, C. Jamesii Schwein., n = 35, and C. Willdenovii Schkuhr,
n = 31, occur in the Southeast. In addition to features of the perigynia, these
species are distinguished by the distribution of micropapillae on the leaves and
culms (Maloney & Evans).
Section ACROCYSTIS Dumort. (Montanae (Kunth) Carey) comprises ten species
in the Southeast and nearly 30 worldwide. The plants grow in the most xeric
habitats of any species of Carex in our area, typically dry woodlands and rock
outcrops. They are small and tufted, with the leaves stiff, the carpellate spikes
few flowered, and the perigynia globose to ovoid, closely covering the roundly
trigonous achenes. Chromosome numbers are reported for half of our repre-
sentatives and indicate an aneuploid series: C. communis Bailey, n = 14, C.
nigromarginata Schwein., n = 17, C. artitecta Mack., n = 18, C. pensylvanica
Lam., n = 18, and C. lucorum Willd. ex Link, n = 20. The European species
are more diverse cytologically (n = 9, 15, 19, 33) but are similar ecologically.
The fruits of C. artitecta (Handel, 1978) and C. nigromarginata (Gaddy, 1986)
are dispersed by ants.
Section PICTAE Kiikenthal has two representatives in eastern North America,
Carex picta Steudel and C. Baltzellii Chapman ex Dewey. Both are local, dry-
woodland species of the unglaciated eastern United States. Carex Baltzellii is
endemic to Georgia and northern Florida. Carex picta, occurring from southern
Indiana to Georgia and Louisiana, is a curious species. It is the only native
dioecious representative of Carex in our area. The plants form "fairy rings"
as the rhizomes branch and proliferate dichotomously (see Martens for illus-
tration). Clones from individual rhizomes are consistently staminate or car-
pellate, and carpellate plants do not always flower every year.
Section CLANDESTINAE G. Don (Digitatae (Fries) Carey) consists of four
species of the North Temperate Zone. The plants have purple leaf sheaths and
perigynia with minute beaks and tapered bases. Carex pedunculata Muhl.,
which grows on wooded, mesic, calcareous slopes, is the only representative
of the section in the Southeast. Elaiosomes are borne at the base of the perigynia,
which are dispersed by ants (Handel, 1976; Gaddy, 1986). Mackenzie (1931-
1935) included the only tetrastigmatic species of Carex, C. concinnoides Mack.
of the Pacific Northwest, in this section. St. John & Parker established subg.
Altericarex for this unusual species, but aside from its tetramerous carpellate
flowers, C. concinnoides fits in sect. CLANDESTINAE rather well, both morpho-
logically and ecologically.
Section TRIQUETRAE (Carey) Kuikenthal comprises five species of temperate
433
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JOURNAL OF THE ARNOLD ARBORETUM
North America. The plants are caespitose, and they have greenish sheaths and
short-beaked, pubescent perigynia. There are two representatives in the South-
east, Care dasc'arpa Muhl. and C. tenax. Chapman ex Dewey. Both grow in
pine forests. mostly from South Carolina to southern Mississippi. Another
species of the section, C. hirtifolia Mack., n = 22 (3)+ (3), of the northeastern
United States, reaches its southern limit in the mountains of Virginia.
Section Ain.Ar: Ascherson & Graebner consists of two species, both boreal
calcicoles of dry soils. One of these, C'arex ehurnea Boott, the only North
American representative, is a stoloniferous plant with glabrous perigynia that
is recorded in our area only from Tennessee. The second is C. a/la Scop., 2n =
54, of Eurasia.
Section PANICEAF G. Don (non Christ) is a Eurasian and North American
section of 12 species, five of which occur in the Southeast. The stoloniferous
plants have purple to reddish scales subtending the flowers, and ascending to
spreading, more or less ovoid perigynia. A member of this section, the rare
southern Appalachian endemic Carevx ilimoreana Mack., occurs on wet, shad-
ed cliffs in the Blue Ridge Mountains of North and South Carolina (Gaddy,
1983). Three of our representatives, C. I'oodii Dewey, n = 22. 26. C. tetanica
Schkuhr, n = 26. and C'. .leadii Dewey, are mostly northern in distribution
and just reach our area in the mountains of Tennessee and North Carolina.
The fifth species, C. Chapmanii Steudel, is endemic to the Coastal Plain be-
tween Florida and North Carolina. The European C. panicea L. and C. vaginata
Tausch have lower chromosome numbers: both are 2n 32.
Section LAXiFLORAF (Kunth) Kukenthal, containing about 25 species in east-
ern North America (17 in our area). one in the western United States, and a
few in eastern Asia. is the most diverse section of Care. in our area. The plants
grow in woodlands: they are caespitose and bear conspicuously two-nerved
perigynia. Our species have recently been studied by Bryson, and Manhart
(1986) has investigated their cytology. Handel (1978a). who investigated the
pollination biology of C'are.x planltai'ea Lam. and C. plat'phll//a Carey, re-
ported that both are self-compatible and that apomixis is absent. He studied
the dispersal of pollen by wind and found that pollen was transported twice as
far from C. plantag, inea as from C. platvph'ylla. This difference was attributed
to the greater average height above ground of the staminate flowers in C.
planlaginea. An aneuploid series is evident in those southeastern representa-
lives of the section for which chromosome numbers have been reported: C.
Alanhartii Bryson, n - 14, C. purpurifiera Mack., n = 17, 18, 19, C. leptonervia
(Fern.) Fern., n - 18, 19. C. blunda Dewey, a 18, 19, 20, 21, 22, C. gra-
cilescens Steudel, n = 20, C. lax/ilora Lam., n = 20, C'. laxicunlmis Schwein.,
n . 22. 23, C. digitalis Willd.. n 24, C. plantaginea, n . 25, and C. plaly-
phl//a, n = 33, 34. 35. Carex striatula Michx. and (C. laxillora are mvrme-
cochorous (Gaddy. 1986).
Section GRANIiFARFiS (0. F. Lang) Kukenthal includes five eastern North
American species, of which four are found in the Southeast. They are calcicoles
and have few-flowered pedunculate spikes and perigynia with many line nerves.
C(arex granularis Muhl. ex Willd., n 16+(4), occurs in all the southeastern
states and is the widest-ranging species of the section. The other southeastern
434
[vot.. 68


TUCKER, CYPERACEAE
representatives are C. rectior Mack., C. Crawei Dewey, and C. microdonta
Torrey & Hooker.
To sect. OLIGOCARPAE (Carey) Kuikenthal (including sect. Griseae Bailey)
belong nine species of eastern North America, of which six are present in our
area. Members of this section are ecologically and morphologically similar to
plants of sect. GRANULARES but have lower chomosome numbers. Carex flac-
cosperma Dewey (C. glaucodea Tuckerman), C. oligocarpa Schkuhr, n = 27,
and C. grisea Wahlenb. (C. corrugata Fern.), n = 28, occur nearly throughout
our area and are also found in the northeastern United States.
Species of sect. HYMENOCHLAENAE (Drejer) Bailey (including sects. Sylvaticae
Boott and Gracillimae (Carey) Kukenthal) are widely distributed in the tem-
perate regions of the Northern Hemisphere and in the East African Highlands
(Kilkenthal; Mackenzie, 1931-1935). The plants have slender, drooping spikes
and often strongly beaked perigynia. A European representative of this section,
Carex sylvatica Hudson, 2n = 58, is naturalized in southern New England and
Long Island. There are six species in the Southeast. The eastern North American
representatives are currently being revised (with particular attention to cytol-
ogy) by Waterway (in prep.). Reznicek (1986a) has provided a detailed illus-
trated study of the Mesoamerican species. Chromosome numbers have been
reported for C. gracillima Schwein. (n = 5, 27), C. flexuosa Muhl. ex Willd.
(n = 27, 28), C. aestivalis Curtis (n = 28), and C. prasina Wahlenb. (n = 30),
all of which occur in the Southeast. Carex cherokeensis Schwein., wolf-tail,
reported from every state in the Southeast, and C. Sprengelii Dewey, n = 21,
of the northeastern United States, are sometimes segregated into sect. Longi-
rostres Kiikenthal because of their longer perigynial beaks.
Section VIRESCENTES (Kunth) Carey is represented in temperate North Amer-
ica, Eurasia, and the mountains of northern South America. The plants have
densely cylindrical, stiffly erect spikes. There are six species in eastern North
America, and all occur in the Southeast. Our representatives for which chro-
mosome numbers are known (Carex Bushii Mack., n = 24, C. hirsutella Mack.,
n = 26, C. Swanii (Fern.) Mack., n = 27, and C. virescens Muhl. ex Willd.,
n = 30) provide yet another example of the aneuploidy so frequent in the genus.
Species of sect. CAREX (sect. HIirtae (Tuckerman) Christ) are widespread in
the Northern Hemisphere, and a few are disjuncts in temperate South America.
The plants are stoloniferous and have three to ten spikes of ascending, ovoid
perigynia. The section has only two representatives in the Southeast: the North-
eastern and midwestern Carex lanuginosa Michx., n = 39, is known in our
area only from Arkansas, while C. striata Michx. (non C. striata Gilib., nom.
illeg.; C. Walteriana Bailey), of the Coastal Plain, ranges from Georgia north
to southeastern Massachusetts. The type species of this section and of the genus,
C. hirta L., n = 56, is sparingly adventive in the northeastern United States
(south to the District of Columbia).
Section ANOMALAE Carey includes many species in eastern Asia and Aus-
tralasia, one in the western United States, and another in the eastern United
States, Carex scabrata Schwein., n = 27, recorded in our area from North
Carolina, Tennessee, and northern Alabama. Plants of this species have dense,
1987]
435


JOURNAL OF THE ARNOLD ARBORETUM
cylindrical carpellate spikes and perigynia with bidentate beaks: they are sto-
loniferous and typically grow near woodland springs.
The monotypic sect. SHORTIANAE (Bailey) Kiikenthal contains Carex Shor-
tiana Dewey, an uncommon but attractive species of the Ohio River valley
south to central Tennessee. The plants have culms each bearing four or five
gynecandrous spikes of nerveless, corrugated perigynia with stipitate bases and
entire beaks.
The species of sect. PENDULINAE (Fries) Christ have a circumpolar distri-
bution and are characterized by pedunculate spikes and closely spaced peri-
gynia. The type species is the European Carex pendula Hudson, 2n = 58 or
60. The three representatives in our area, C. Joorii Bailey, C. verrucosa Muhl.,
and C. glaucescens Ell., are all widely distributed.
Species of sect. LIMOSAE (Tuckerman) Christ have drooping, few-flowered
spikes and broadly elliptic, beakless perigynia. Many are circumboreal in dis-
tribution and grow in fens, bogs, or wet woods. A single, primarily northeastern
representative, C'arex Barrattii Schwein. & Torrey, from the mountains of
Tennessee and North Carolina, is known in our area. The type species is the
circumboreal C. iunosa L., 2n = 56.
The diverse and heterogeneous sect. ATRATAE (Kunth) Christ'' contains many
species of the arctic and alpine tundra. The plants are characterized by sessile,
erect or drooping spikes, dark pistillate scales, and beaked or beakless perigynia.
There are many representatives in the southern Rocky Mountains (Hermann,
1970; Murray), but none of these is shared with our area. The single species
of our area, Carex Buxhaunii Wahlenb., n = 37, ca. 50. reaches its southern
limit in North Carolina and Arkansas.
Section PHACOCYSTis Dumort. (sect. ,'cutae Fries) is also a diverse circum-
boreal group. The plants are moderately large and have drooping spikes and
distigmatic, lenticular achenes. Three northeastern species, C(arex strictior
Dewey, n = 34, C. strica Lam., and C. torta Boott ex Carey, n = 33, reach
their southern limits in the northern half of our area. Standley (1985a) revised
the 15 representatives of this section in the Pacific Northwest. While none of
the species she treated occurs in our area, her thorough investigation of inter-
specific differences in leaf and culm anatomy, cytology, morphology, and some
aspects of ecology is informative and provides a model for future studies.
Species of sect. CRYPTOCARPAE (Tuckerman) Kiikenthal are mostly wetland
plants. They have drooping, densely flowered spikes and trigonous achenes.
C'arex ginandra Schwein.. C. tlitchelliana M. A. Curtis, and C. crinita Lam.,
n = 33, occur in the Southeast. These have been treated as a single taxon under
the last name, but there is good evidence for their specific status (Bruederle &
Fairbrothers, 1986). Carex ginandra and C(. crinita hybridize rarely. The hy-
brids produce aborted achenes (Standley, 1983).
Section Coii INSIAE Mack. contains a single species, Carex Collinsii Nutt.,
that grows in swamps on the Atlantic Coastal Plain from Georgia to Rhode
"(N rc~ sect. ATRATIA (Kunth) (hrist, Bull. Soc. Bot. Belg. 24: 15. 1885.
436
[voL. 68


TUCKER, CYPERACEAE
Island (Tucker, 1978). It is characterized by few-flowered inflorescences and
subulate perigynia.
Species of sect. FOLLICULATAE Mack. also have subulate perigynia, but the
spikes are densely many flowered and the plants are taller. There are two
representatives in the Southeast, Carex lonchocarpa Willd. ex Sprengel, found
throughout our area, and C. folliculata L., n = 28, a northeastern species
growing only in the mountains of North Carolina and Tennessee.
Species of sect. PSEUDO-CYPEREAE (Tuckerman) Christ are tall, paludal plants
of circumpolar distribution. They have drooping, slenderly cylindrical spikes
and densely arranged, conspicuously bidentate perigynia. There are two rep-
resentatives in our area, Carex Schweinitzii Dewey, n = 30, and C. comosa
Boott, n = 32. Carex pseudocyperus L., 2n = 66, is widespread in the Northern
Hemisphere and is believed to be native to New Zealand (Clapham et al.).
Section PALUDOSAE G. Don20 has eight species in North America and several
in Eurasia. The plants are stoloniferous and bear firm, many-nerved, slightly
inflated perigynia. There are two representatives in our area, Carex hyalinolepis
Steudel, found in wetlands throughout the Southeast, and C. trichocarpa Muhl.
ex Schkuhr, n = 55, a boreal bog species known in the Southeast only from
the mountains of North Carolina (Core).
Dense spikes of conspicuously inflated perigynia characterize members of
sect. SQUARROSAE Carey, which are endemic to eastern North America. There
are three species in our area, Carex Frankii Steudel, C. typhina Michx., and
C. squarrosa L., n = 28, each occurring in all or most of the southeastern states.
Section VESICARIAE (Tuckerman) Carey is a group of perhaps 20 species,
mostly of eastern North America and Eurasia. The plants generally grow in
shallow water and are characterized by inflated perigynia. Five representatives
occur in our area, but only one, Carex lurida Wahlenb., n = 32, 33, is common
(reported from every state). The others are C. Baileyi Britton, n = 34, C. bullata
Schkuhr, C. Elliottii Schwein. & Torrey, and C. rostrata Stokes, n = 34. The
type species, C. vesicaria L., n = 4 1, and C. riparia Curtis, n = 36, are cyto-
logically similar Eurasian representatives.
REFERENCES:
Under family references see BARNARD; BARROS (1935): BEAL; BENTHAM: BERGGREN:
BLASER (1941 c); BREWBAKER: CLARKE (1908, 1909): CLIFFORD & HARBORNE: COOK: EYLES
& ROBERTSON; FASSETT; GADDY (1986); GIBBS; GODFREY & WOOTEN; GONCHAROV et
al.: GooD et al.: HARBORNE: HARBORNE et al.: HARRIS & MARSHALL: HESSE: HOLTTUM:
HUANG; G. E. HUTCHINSON: .1. HUTCHINSON: KOYAMA (1962a); KRAL; LE MAOUT &
DECAISNE; LERMAN & RAYNAL; LLOYD & WOOLHOUSE; LOVELL: MEEUSE: METCALFE:
NOBLE & MURPHY: OGDEN: PATCH: RAYNAL (1972): RIKLI; SAVILE; SCHULZE-MOTEL
(1959, 1964); STACE; STANDLEY: TEERI ct al.; TIETZ: TORREY: and WINFREY & SAMSEL.
Under Rhynchospora see LEPPIK.
Under Cvmophyllus see K1JKENTHAL: MACKENZIE (1931-1935).
20Carex sect. PALUDOSAE G. Don in Loudon. Hort. Brit. 367. 1830. non (Fries) Christ (1884). TYPE
SPECIES: C. paludosa Good. (= C. acutfonnirrs Ehrh.).
1987]
437


JOURNAL OF THE ARNOLD ARBORETUM
AKIYAMA, S. On the systematic anatomy of the leaves of some Japanese ('arices. (In
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BAHii Y, L. H. A preliminary synopsis of the genus ('arex. Notes on Mexico, Central
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Bi.n, K. 1,., & L. C. l iss. Autecology of Ko/re'sia Bellardii: why winter snow accu-
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--- & E. GORHAM. Life history aspects of primary production in sedge wetlands.
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BrRRY, E. W. Fossil grasses and sedges. Am. Nat. 39: 345-348. 1905. [Including C.
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Bo'HER, T. G. A study of the circumpolar ('arex leleormastes-amblvorh'rncha complex.
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BROWN, L. E., & C. D. Pr iRSON. ('ares rosea (Cyperaceae), Triolium lappaceum
(Fabaceae). and .lira carop/ayllea (Poaccae) new to Texas. Sida 10: 263, 264. 1982.
BRI Diu.RIL, L. P.. & I). FI. FAIRBROI mli RS. Preliminary chemosystematic investigations
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162: 86, 87. 1982. [Summary of techniques; no species mentioned.]
& -- Variability and taxonomic usefulness ofachene and perigynium char-
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suppl. 2): 107. 1983.
- & . Systematic investigations of C arev M1itchelliana (Cyperaceae). (Ab-
stract.) Ibid. 71(5, part 2): 159. 1984.
--- & ---- enetic variation in populations of Care.v crinita Lam. (Abstract.)
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--- & -- . Alloryme variation in populations of the Cuaret crinna complex
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1987]
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