Download The Genera of Chloridoideae (Gramineae)

The Genera of Chloridoideae (Gramineae)
in the Southeastern United States1,2
Gordon C. Tucker3
1
Prepared for the Generic 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 BSRxxxxx (Norton G. Miller, principal investigator), under which preparation of this account was
commenced, and BSR- xxxxxx (Walter Judd, principal investigator). This treatment, the
umpteenth in the series, follows the format established in the first paper (Jour. Arnold Arb. 39:
296-346. 1958) and continued to the present. The area covered by the Generic Flora includes
North and South Carolina, Georgia, Florida, Tennessee, Alabama, Mississippi, Arkansas, and
Louisiana. The descriptions are based primarily on the plants of this area, with information about
extraregional members of a family, subfamily, or genus in brackets [ ].
Norton Miller, Carroll Wood, Ken Robertson, Tom Rosatti, Gerrit Davidse, John Ebinger,
Dave Seigler, Ihsan Al-Shehbaz, Wilma Campbell, and Barbara Nimblett have provided
suggestions, encouragement, and technical assistance and I wish to thank them for making my work
on the Generic Flora enjoyable and rewarding. William J. Crins has shared his extensive familiarity
with grasses, particularly of Muhlenbergia and Eragrostis. Thanks are extended to the staffs of the
New York State Library, the Botany Libraries of Harvard University, Missouri Botanical Garden
Library, Booth Library of Eastern Illinois University, and the Illinois Natural History Survey
Library for providing access to many references. I thank the curators of the following herbaria who
have sent specimens or provided hospitality and access to the collections under their care during my
visits: A, CLEMS, CONN, ECON, EIU, GH, ILL, ILLS, IND, ISM, MO, MU, MUHW, NEBC,
NY, NYS, SIU, and WIS.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 1
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The illustrations used here were published earlier as Figures x-x in Christopher Campbell's
account of the subfamilies and tribes of grasses prepared for the Generic Flora (see Footnote 2
below). The figures were drawn by Karen Stoutsenberger during a previous grant period (NSF
BMS-21469, Carroll Wood, principal investigator) under the supervision of Carroll Wood, Kenneth
R. Robertson, and Christopher Campbell, who also prepared the materials.
-----------------------------------------------------------------2
For an account of the family, see C. S. Campbell, The subfamilies and tribes of the Gramineae
(Poaceae) in the southeastern United States. Jour. Arnold Arb. 66: 123-199. 1985.
---------------------------------------------------------------3
Stover-Ebinger Herbarium, Department of Biological Sciences, Eastern Illinois University,
Charleston, Illinois 61920; e-mail: [email protected]
Chloridoideae Rouy, Fl. France 14: 2. 1913.*
Chloridoideae Kunth ex Beilschm. {{fide FNA 23}}
Perennials or annuals of diverse habitats. Roots fibrous; rhizomes often present.
Microhairs short and stout; anatomy kranz (PS type), the photosynthetic pathway exclusively C4
(except one Eragrostis sp.). Stems erect or decumbent, simple or sometimes branched near base,
sometimes rooting at the nodes, the internodes hollow. Leaves cauline or basal, few or many, not
distinctly distichous (distinctly distichous in Distichlis and Monanthochloë); sheath open, longer or
shorter than the corresponding internode; ligule membranaceous; blade linear or lanceolate, not
auriculate, flat, convolute, or involute [terete]. Inflorescence a panicle, spike, raceme, or group of
racemes, often secund, often partly included in the subtending sheath. Spikelets laterally
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 2
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compressed, 1-15 (30) flowered; glumes 2, typically equal, about as long as the first lemma; rachilla
articulated above (infrequently below) glumes; flowers bisexual (unisexual, the plants dioecious in
Distichlis, Monanthochloe, and some Eragrostis); lemmas oblong to lanceolate, 1-3 nerved; paleas
elliptic, mostly about as long as the lemmas, usually 2-nerved. Lodicules 2, cuneate or truncate
(less often rounded or acuminate), fleshy. Stamens 3 (infrequently 1 or 2); anthers ellipsoid to
linear. Ovaries ellipsoid to oblong, glabrous or hairy apically; styles 2 [3 in Munroa], separate,
long, short, or obsolete, stigmas terminal, plumose. Fruit a caryopsis (achene with free pericarp in a
few genera), subterete to cylindric, sometimes flattened laterally, linear, oblong, or ellipsoid; hilum
punctiform to linear, from 1/5 to nearly as long as grain; embryo large, typically 1/3 –3/4 the length
of the grain; endosperm solid. Base chromosome number 10 (7, 8, 9, 12 in one or more genera).
(Eragrostoideae Pilger, Nat. Pfl. ed. 2, 14d: 167. 1956*) Type genus: Chloris Sw.
A large subfamily, worldwide in temperate and tropical regions. The hallmark of the
Chloridoideae is the typically three-nerved lemma, consistently kranz anatomy, and distinctive
microhairs. There is some similarity to the Arundinoideae. The Chlorodoideae is one of the three
largest subfamilies of the grasses, along with the Pooideae and Panicoideae. The subfamily in its
present sense was first proposed by Pilger (1954).
Five tribes were recognized by Clayton and Renvoize. The Eragrostideae Stapf (76 genera)
and the Cynodonteae Dumort. (58 genera) are both large and widely distributed. In contrast, the
Pappophoreae Kunth (five genera), Leptureae Holmberg (monotypic) and Orcuttiae Reeder (three
genera) are quite small.
All genera of the Chloridoideae have the C4 photosynthetic pathway, the only exception
being Eragrostis walteri, which most likely is a case of reversion (Chapman). Three of the eight
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 3
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recognized variants of C4 occur in the subfamily. The “Classical NAD-ME” type has centripetally
arranged chloroplasts in the bundle sheath cells, the decarboxylating enzyme is NAD malate
dehydrogenase; this type occurs in Buchloë, Cynodon, and some species of Leptochloa. The
“Classical PCK” type has centrifugal or uneven arrangement of the chloroplasts, and the
decarboxylating enzyme is phosphoenolpyruvate carboxykinase; this occurs in Chloris,
Dactyloctenium, Spartina, Zoysia, and some Leptochloa. The third type, the “Triodia type”, has
centrifugally arranged chloroplasts, and NAD malate dehydrogenase as the decarboxylating
enzyme. This type occurs in several Australian genera, including Triodia and Monodia, but not in
any genera native or naturalized in our area.
Molecular systematics and integrity of subfamily have received some attention. A
monophyletic origin for the subtribe Muhlenbergiinae is supported by restriction site analysis of
chloroplast DNA. Duvall et al. investigated of 25 species of Eragrostideae, one species of
Cynodonteae (Eustachys distichophylla), and one species of Pooideae. They detected 124
restriction sites, of which 67 were variably present and shared by two or more species. Cladistic
analyses established that members of the Muhlenbergiinae, including the genera Muhlenbergia,
Blepharoneuron, Bealia, Chaboissaea, Lycurus and Pereilema, share seven restriction site
mutations and are strongly supported by the data as a monophyletic subtribe. The monotypic
Great Plains genus Redfieldia also clustered with the subtribe Muhlenbergiinae in their analysis,
perhaps indicative of a past intergeneric hybridization event or convergent evolution. Their
restriction site data also weakly support a relationship (six shared mutations) between
Erioneuron, Munroa, and Dasyochloa.
The reed-like genus Neyraudia, previously included in the Arundinoideae and treated in
the GFSEUS account of that subfamily, is now generally placed in the Chloridoideae. It appears
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 4
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to be most closely related to …. (Barkworth Grass Manual).
--------------Biogeography; 17 genera in the Intermountain West
In several genera, the mature pericarp is more or less free from the seed, being fused to the
seed only in the vicinity of the hilum. For such fruits, the term achene has been used. Also the terms
modified caryopsis and cistoid pericarp have been proposed, though scarcely taken up. In Crypsis,
Eleusine, and Sporobolus, for example, the mesocarp and endocarp disintegrate during
development and maturation leaving only the exocarp. When moistened, the seed may actually be
extruded from the mature pericarp (Sendulsky et al.).
Reproductive biology of the subfamily is distinct. In 13 genera, cleistogamous spikelets
have been reported noted in one or more species, including only Leptochloa and Muhlenbergia in
the southeast (Connor). Cleistogamous subterranean spikelets are known from only four genera of
the family; one of the representatives is a species of Chloris. Gynomonoecy is known from only one
genus, Munroa (Connor, 1981), which is apparently also self-incompatible. Blepharidachne has
three monoecious species, and the fourth, B. kingii, has bisexual flowers (Connor). Stamens
number 1, 2, or 3, depending on the species (Anton & Hunziker). The Chlorideae have a high
number of dioecious genera, seven (Bouteloa, Buchloë, Buchlomimus, Cyclostachya, Opizia,
Pringleochloa, Soderstromia), more than any other tribe in the family, while in the Eragrostideae
only Neeragrostis and Scleropogon are dioecious (Connor). Apomixis is documented from three
genera of the Chloridoideae (Bouteloua, Chloris, Eragrostis).
References:
Amarasinghe, V., & L. Watson. Variation in salt secretory activity of microhairs in grasses.
Austral. Jour.Pl Physiol. 16: 219-229. 1989.*
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 5
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Anton de Triquell, A. Grass gametophytes: their origin, structure, relation with the sporophyte. Pp.
11-20 in T. R. Soderstrom, K. W. Hilu, C. S. Campbell, & M. E. Barkworth, eds., Grass
Systematics and Evolution. Washington, D.C. 1987.
Anton, A.M. Notas criticas sobre Gramineas de Argentina. Kurtziana 10:51-67. 1977.
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(Reprint, J. Cramer, Weinheim. 1961.)
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fusca (L.) Kunth. Environm. Exper. Bot. 31: 437.*
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Mexico. Magnoliophyta: Commelinidae (in part): Poaceae, part 2. xxv + 783 pp. New York and
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pollen. quantitative analysis. Geographic rhode island. Abstract While flower predators
can limit the sexual expression and seed production of salt marsh grasses, the relationship
between these two effects of consumers has not been explored. At our study site,
predation on Spartina patens, Spartina alterniflora, and Distichlis spicata was twice as
high in 1985 (approximately 70% ovule destruction) as in 1986 (approximately 35%
ovule destruction). In both years consumers destroyed flowers before maturity, reducing
sexual expression, and particularly suppressed male sexual expression. Sexual
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 7
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suppression of males was much more pronounced in 1985 when flower predation was
severe and the seed production of undamaged ovules was dramatically reduced. A
number of lines of evidence suggest that predator limitation of male sexual expression
and pollen supply contributed to low seed output in 1985. 1) Undamaged ovules of all
three grasses protected from consumers but exposed to ambient windborne pollen set
many more seeds in 1986 than in 1985, suggesting that pollen was more abundant in
1986; 2) Artificial pollinations revealed that marsh grasses are generally pollen-limited
and that pollen limitation at our study site was more severe in 1985 than 1986; and 3)
Caging stands of marsh grasses generally led to less predator damage, increased male
densities and seed sets similar to those for hand-pollinated flowers. Our results support
the hypothesis that flower predators can indirectly limit seed production by decreasing
pollen availability
----- & -----. Competition and facilitation in marsh plants. Am. Nat. 142: 718-724. 1993.
[Spartina patens, Distichlis spicata in California.]
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------. Morphology and Anatomy of the Western Australian Species of Triodia R. Br., II:
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(Gramineae: Chloridoideae). Aliso 17(2): 99-130. 1998.*
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48-74. 1981. [Good review of breeding systems for all grasses; monoecy, andromonoecy,
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-----. Reproductive biology in grasses. Pp. 117-132 In T. R. Soderstrom, K. W. Hilu, C. S.
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----- & -----. Embryology of Chloris roxburghiana Schult. (Poacaeae). Taiwania 36: 303-310.
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Eragrostis.]
Watson, L., & E. M. Bell. A surface-structural survey of some taxonomically diverse grass
pollens. Austral. Jour. Bot. 23: 981-990. 1975. [Photomicrographs of pollen of Cynodon
dactylon, Chloris truncata, Sporobolus airoides; sadly, chloridoid pollen grains scarcely
distinguishable from those of Arundinoideae and Panicoideae.]
Watson, L., H. T. Clifford, & M. J. Dallwitz. The classification of Poaceae: subfamilies and
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 27
06/27/17
supertribes. Austral. Jour. Bot. 33: 433-484. 1985. [720 currently recognized genera
distributed among 5 subfamilies and 6 supertribes; detailed descriptions.]
Watson, L. & M.J. Dallwitz. The grass genera of the world. Rev. Ed. 1081 pp. Cambridge,
England. 1992. [Detailed parallel descriptions of all chloridoid genera.]
Whigham, D.F. & Nusser, S.M. The response of Distichlis spicata (L.) Greene and Spartina
patens (Ait.) Muhl. to nitrogen fertilization in hydrologically altered wetlands. pp. :3138. In D.F. Whigham, R.E. Good and J. Kvet, eds., Tasks for vegetation science. 1990..
[Distichlis spicata, Spartina patens in Maryland.]*
Wolfe, J., &, J.M. Zajicek. Are ornamental grasses acceptable alternatives for low maintenance
landscapes? Jour. Environm. Hort. 16(1): 8-11. 1998. [Includes references; Eragrostis
superba, Muhlenbergia lindheimeri.]
Wunderlin, R. P. Guide to the vascular plants of Central Florida. viii + 472 pp. Tampa and other
cities. 1982. [Poaceae, 51-96.]
Yates, H.O. Uniola. Southwest. Natur. 11: 372-394. 1966.
Yatskievych, G. Steyermark’s Flora of Missouri. Volume 1. (Revision of J.A. Steyermark, Flora
of Missouri. 1963.) xii + 991 pp. Jefferson City, Missouri. 1999. [Chloridoideae, 673-742;
includes many genera and species in Southeast; keys, illustrations, detailed descriptions;
generic boundaries mostly similar to those herein; Neeragrostis included in Eragrostis.]
Yeoh, H.-H., & L. Watson. Systematic variation in amino acid compositions of grass caryopses.
Phytochemistry 20: 1041-1051. 1981. [Profiles of Chloridoids distinguishable from other
subfamilies.]
Yuvaniyama, A. & S. Arunin. Growth of three halophytic grasses on salt-affected soil in
northeast Thailand. ACIAR proceedings 42: 32-35. 1991. [Includes Sporobolus spp.,
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 28
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Spartina patens, Distichlis spicata..
Judd BI, 1975., New World tropical forage grasses and their management: Bermudagrass, giant
stargrass, St Augustinegrass, Jaraguagrass. World Crops 27. (2): 69-72
-------------------------------------------------------------------------doubtful genera – some of these may be in the Southeast
Tribe Pappophoreae
Pappophorum, southern US fide c+R
Cottea, same note
Vaseyochloa Hitchc., is monotypic, V. multinervosa (Vasey) Hitchc. is endemic to the coast of
southern Texas (does not get close to Louisiana).
Blepharidachne Hack. includes three species, and is found in western Texas and adjacent northern
Mexico.
Blepharoneuron Nash western Texas to Arizona and south into Mexico.
Pappophorum Schreb., 2 species in southwestern U.S., Texas and Arizona southward (Gould);
“Southern USA” to Argentina (Clayton & Renvoize).
Scleropogon western Texas to Nevada
Erioneuron Texas to Colorado
Blepharoneuron southwestern states
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 29
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Munroa Great Plains, in Kansas and Oklahoma
Blepharidachne (not in Louisiana)
.Trichoneura occurs in TX, but not in Louisiana
Key to the genera in the Southeastern United States
Some notes:
Bouteloua, single spike of secund spikelets
Calamovilfa
Chloris + Eustachys have spikelets in two rows
Crypsis
Cynodon
Diplachne spikelets in two rows
Eleusine same
Eragrostis
Muhlenbergia
Spartina spikelets in two rows
Sporobolus grain falls from chaff
Tridens axis of panicle sticky; lemma villous basally
Triplasis midvein of palea bearded distally
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 30
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Tribe Eragrostideae Stapf, Fl. Capensis 7: 316. 1898.
1. Uniola Linnaeus, Sp. Pl. 71. 1753; Gen. Pl. 32. 1754.
Perennials of sandy coastal dunes. Rhizomes present, horizontal. Stems erect , cespitose,
unbranched; leaves basal and cauline, few, not distinctly distichous [except the basal ones in U.
pittieri]; sheath glabrous but villous in the throuat; ligule a fringe of short white hairs [absent];
blade linear, coriaceous, flat or plicate, tapering to a point. Inflorescence a panicle, with the
branches ascendant, or spreading and drooping, secondary branches spreading. Spikelets strongly
laterally compressed, (3) 6-18 (30) flowered; glumes 2, ± equal, lanceolate to ovate-lanceolate, 3-5
nerved, the second about as long as the first lemma, 2-5 nerved; rachilla disarticulating below
glumes; flowers bisexual (the lower [1] 2-6 sterile, and the upper ones generally sterile as well);
lemmas narrowly elliptic to oblong-lanceolate, 3-9 nerved, acute, smooth except for the serrulate
midvein, glabrous; paleas elliptic [oblong-ovate], shorter than the lemmas, 2-keeled, the keels
ciliate or smooth. Lodicules 2, cuneate, truncate, fleshy. Stamens 3; anthers ellipsoid. Ovaries
ovoid to ellipsoid, glabrous; styles 2, separate, short, terminal stigmas plumose. Fruit an achene,
terete, oblong [oblong-ovate]; hilum linear, nearly as long as grain; embryo large, about ½ the
length of the grain. Base chromosome number 10. Type species: U. paniculata L. (see Nash in
Britton & Brown, Ill. Fl. No. U.S. ed. 2. 1: 248. 1913; Linnaeus included both U. paniculata and U.
spicata). – Sea-Oats.
Discussion:
U. paniculata L.
VA to FL and TX and Tamps. to Tabasco (MO)
U. pittieri
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 31
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Bahamas and West Indies
Oaxaca south to Colombia on Atlantic and Pacific Coasts (MO)
L. virgata (Leptochloopsis)
U. condensata
References
Under subfamily references see: Laegard & Sanchez Vega.
Hovanesian, J. Micropropagation of Uniola paniculata L. Plant cell reports. 1986. v. 5 (5)
:385-386. ill. Note Includes references. uniola paniculata. duneland plants. tissue
culture. methodology. Other Torres, K.C.
Wilsey, M.H. Propagation of sea oats (Uniola latifolia). The Plant propagator. June 1985. v. 31
(2) :15. uniola. duneland plants. propagation. sand stabilization. coastal areas.
OtherSubject erosion control plants Geographic alabama.
Tyndall, R.W., A.H. Teramura, C.L. Mulchi & L.W. Douglass. Effects of salt spray upon
seedling survival, biomass, and distribution on Currituck Bank, North Carolina.
Castanea 52: 77-86. 1987.. [Uniola paniculata, Spartina patens, Triplasis purpurea.
Rowan, W.T. Insects found on sea oats, Uniola paniculata, in south Florida. Florida
entomologist 63: 341. 1980.
Hester, M.W., & I.A. Mendelssohn. Water relations and growth responses of Uniola
paniculata (sea oats) to soil moisture and water-table depth. Oecologia. 1989. v. 78 (3) :289296. ill. Note Includes references. uniola paniculata. soil moisture. growth. water table. water
relations. drought. dunes. OtherSubject inundation Geographic louisiana. Other
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 32
06/27/17
Wade, M.R., D.F. Gilman, & J.C. Colosi. Germination response of Uniola latifolium.
HortScience. Aug 1990. v. 25 (8) :915-916.. uniola. chilling requirement. potassium
nitrate. seed germination. temperature relations. diurnal variation. germinability. seed
treatment. seed stratification.
___, ____, & _____. The seed germination patterns of two wide ranging coastal dune dominants.
Bull. Ecol. Soc. Amer. 58: 25. 1977.*
Hester, M.W. &, I.A. Mendelssohn. Effects of macronutrient and micronutrient additions on
photosynthesis, growth parameters, and leaf nutrient concentrations of Uniola paniculata
and Panicum amarum. Bot. Gaz.. 151: 21-29. 1990. Note Includes references. uniola
paniculata. panicum. duneland plants. npk fertilizers. trace element fertilizers. nutrient
removal by plants. nutrient contents of plants. leaves. stems. growth rate. dry matter
accumulation. photosynthesis. Geographic louisiana.
_____ & _____. Expansion patterns and soil physicochemical characterization of three
Louisiana populations of Uniola paniculata (Sea oats). Jour. Coastal Res. 7: 387-401.
1991. [Low seed production and vegetative growth of less than 1 m per year limits range
expansion on eroding shoreline.]
Gilreath, J.P. Response of container grown sea oats to selected preemergence herbicides.
Proceedings of the annual meeting of the Florida State Horticultural Society. Florida State
Horticultural Society. Meeting.. 105 : 202-204. year [Uniola paniculata.]
Seneca, E.D. Germination and seedling response of Atlantic and Gulf coast populations of
Uniola paniculata. Am. Jour. Bot. 59: 290-296. 1972.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 33
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Seneca, E.D. Germination response to temperature and salinity of four dune grasses from the
Outer Banks of north Carolina. Ecology 50: 45-53. 1968.*
Van der Valk, A.G. The floristic composition and structure of foredune plant communities of
Cape Hatteras National Seashore. Chesapeake Sci. 16: 115-126. 1975.*
Sylvia, D.M., A.G Jarstfer, & M. Vosatka. Comparisons of vesicular-arbuscular mycorrhizal
species and inocula formulations in a commericial nursery and on diverse Florida
beaches. Biology and fertility of soils.. 16 (2): 139-144. 1993. Note Includes references.
uniola paniculata.
Oertel, G.F., & M. Larson. Developmental sequences in Georgia coastal dunes and distribution of
dune plants. Bull. Georgia Acad. Sci. 34: 35-48. 1976. [U. paniculata in zone nearest high
tide.]
Woodhouse, W.W., Jr., E.D. Seneca, & A.W. Cooper. Use of sea oats for dune stabilization in the
southeast. Shore and Beach 36: 15-21. 1968.*
2. Distichlis Rafinesque, Jour. Phys. Chim. 89: 104. 1819.*
Dioecious (rarely monoecious) mat or sward forming perennials of tidal marshes [alkaline
or saline inland wetlands]. Rhizomes present, scaly; stolons present. Stems erect , cespitose,
unbranched; leaves cauline, distichous, few; sheath glabrous or ciliate; ligule an erose, ciliate
membrane; blade linear, flat or involute (or convolute). Inflorescence a panicle (raceme or single
spikelet) with the branches ascendant; secondary branches appressed. Spikelets laterally
compressed, 3-17 (-20) flowered; glumes 2, subequal, ovate-lanceolate, the first (1) 3-7 nerved, the
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 34
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second 5-9 nerved (lateral nerves difficult to discern); rachilla disarticulating above glumes;
flowers unisexual (the upper ones generally sterile); lemmas elliptic to oblong-lanceolate, 5-11
nerved, acute, smooth, glabrous; paleas elliptic, a little shorter than the lemmas, 2-keeled,
(membranous in the staminate flowers, chartaceous in the carpellate). Lodicules 2, cuneate,
truncate, fleshy. Stamens 3; anthers ellipsoid. Ovaries ovoid to ellipsoid, glabrous; styles 2,
separate, short, terminal stigmas plumose. Fruit a caryopsis, terete, oblong; hilum linear, nearly as
long as grain; embryo large, typically1/3 –3/4 the length of the grain. Base chromosome
number 10. (Name from Greek distichos, ‘two-rowed’, referring to the conspicuously two ranked
arrangement of the leaf blades.) Type species: D. maritima Raf. (=D. spicata (L.) Greene). –
Saltgrass.
A genus of 5 species, only one of which occurs in North America. Distichlis spicata ranges
Nova Scotia south to Florida and west along the Gulf coast to Texas, occurs in all the southeastern
states bordering the Atlantic Ocean and Gulf Coast; also
Plants from the interior of North America have been distinguished as D. stricta, based on having
more compact or congested inflorescences than those in coastal areas.
References
Under subfam. see Bertness & Shumway; Johnson-Green et al., Lefor et al.; Smart & Barko;
Whigham & Nusser, and Yuvaniyama, & Arunin.
Barkworth, M.E. Distichlis. Page 25 in Barkworth, M.E., K.M. Capels, S. Long, & M.B. Piep, eds.
Flora of North America, north of Mexico. Magnoliophyta: Commelinidae (in part): Poaceae, part
2. New York and Oxford. 2003.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 35
06/27/17
Beetle, A.A. The North American Variations of Distichlis spicata. Bull. Torrey Bot. Club
70:638-650. 1943.
-----. The Grass Genus “Distichlis.” Revista Argentina de Agronomía, 22:86-94. 1955.
Daines, R.J., & A.R. Gould. The cellular basis of salt tolerance studied with tissue cultures of the
halophytic grass Distichlis spicata. Jour. Pl. Physiol. 119: 279-280. 1985.*
Enberg, A., & L. Wu. Selenium assimilation and differential response to elevated sulfate and
chloride salt concentrations in two saltgrass ecotypes. Ecotoxicology and environmental
safety 32: 171-178. 1995. [D. spicata in California.]
Ho, I. Vesicular-arbuscular mycorrhizae of halophytic grasses in the Alvord Desert of Oregon.
Northwest Sci. 61:148-151. 1987. [D. stricta.]
Ketchum, R.E. B., R.S. Wareen, L.J. Klima, F. Lopez-Gutiérrez, & M.W. Nabors. The mechanism
and regulation of proline accumulation in suspension cell cultures of the halophytic grass
Distichlis spicata L. Jour. Pl. Phys. 137: 368-374. 1990.*
Miyamoto, S., E.P. Glenn, & M.W. Olsen. Growth, water use and salt uptake of four halophytes
irrigated with highly saline water. Jour. arid environm. 32(2) :141-159. 1996. [Distichlis
palmeri in Mexico.]
Prodgers, R.A., &, W.P. Inskeep. Heavy metal tolerance on inland saltgrass (Distichlis spicata).
Great Basin Natur. 51: 271-278. 1991. [Tolerance to copper, manganese, zinc in Montana
Schwarz, A.G. & R.W. Wein. Threatened dry grasslands in the continental boreal forests of
Wood Buffalo National Park. Canad. Jour. Bot. 75:1363-1370. [Distichlis stricta in
Northwest Territories.]
Seelig, B.D., J.L Richardson, & W.T. Barker. Characteristics and taxonomy of sodic soils as a
function of landform position. Soil Sci. Soc. Am. Jour. 54:1690-1697. 1990. [North Dakota
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 36
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study; D. stricta associated with highest sodium levels along gradient.]
Warren, R.S. Photosynthesis, respiration, and salt gland activity of Distichlis spicata in relation
to soil salinity. Bot. gaz. 150:346-350. 1989.
Wu, L., A.W. Enberg, & X. Guo. Effects of elevated selenium and salinity concentrations in root
zone on selenium and salt secretion in saltgrass (Distichlis spicata L.). Ecotoxicology
and environmental safety. 37:251-258. 1997 . [Nutrient uptake.]
Wu, L., & Z.Z. Huang. Selenium accumulation and selenium tolerance of salt grass from soils
with elevated concentrations of Se and salinity. Ecotoxicology and environmental safety.
22: 267-282. 1991. [Metal tolerance of Distichlis spicata in California.]
Yensen, N.P. & L.L. Bedell. Considerations for the selection, adaptation, and application of
halophyte crops to highly saline desert environments as exemplified by the long-term
development of cereal and forage cultivars of Distichlis spp. (Poaceae). Tasks for
vegetation science 28: 305-313. 1993.* [High salinity tolerance noted.]
Yensen, S.B., &, C.W. Weber. Composition of Distichlis palmeri grain, a saltgrass. Jour. food
science 51(4):1089-1090. 1986. [Proximate analysis of seed composition.]*
Zhao, Z., J.W. Heyser, & J.W. Bohnert. Gene expression in suspension culture cells of the
halophyte Distichlis spicata during adaptation to high salt. Pl. Cell Physiol. 30: 861-867.
1989.*
3. Monanthochloë Engelmann, Trans. Acad. Sci. St. Louis 1: 436. 1859, pl. 13.1
Dioecious mat or sward forming perennials of seashores and tidal marshes [alkaline or
1
Engelmann included the following syntypes: Texas, Drummond; Matamoros, Berlandier 3227;
Florida, Key West, Blodgett; Texas, Galveston, May, Lindheimer.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 37
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saline inland wetlands or flats]. Rhizomes present, scaly; stolons present, long, wiry. Stems
decumbent to erect , solitary, i.e., one per node, unbranched; leaves distichous, cauline, clustered on
short axillary shoots; sheath rounded, longer than internode, glabrous or puberulent; ligule a short
ciliate membrane; blade linear, conduplicate or involute. Inflorescence a single spikelet, partially
inclosed by the uppermost leaf. Spikelets laterally compressed, 2-5 (7) flowered; glumes absent (or
one rudimentary glume present; rachilla disarticulating at base of lowest lemma; flowers unisexual
(the upper ones generally sterile); lemmas elliptic to oblong-lanceolate, vaguely 7-9 nerved, acute,
smooth, glabrous, those subtending carpellate flowers leaf-like in texture; paleas narrowly elliptic, a
little shorter than the lemmas, 2-keeled, coriaceous or indurate). Lodicules absent. Stamens 3;
anthers ellipsoid. Ovaries ovoid to ellipsoid, glabrous; styles 2, separate, short, terminal stigmas
plumose. Fruit a caryopsis, terete, oblong, enclosed by coriaceous palea; hilum linear, nearly as
long as grain; embryo large, typically1/3 –3/4 the length of the grain. Base chromosome
number 10. (Name from Greek, mono, one, anthos, flower, and chloë, grass, thus single-flowered
grass, a reference to the solitary spikelet.) (Halochloa Griseb. ???) Type species: M. littoralis
Engelm., the only species included in the protologue. – Shoregrass.
A genus of two species with an amphitropical distribution. Monanthochloë littoralis
Engelm., shoregrass, 2n = 40, occurs in Florida (Merritt Island south to Keys, and on the Gulf Coast
from Tampa Bay southward), and Louisiana, and Cuba, and from Texas to California and Mexico.
The other species, M. acerosa (Griseb.) Speg. is endemic to south central Argentina (Thieret,
Villamil). The genus is probably related to Distichlis, but differs from that genus, and all other
grasses in North America, in its highly reduced inflorescences.
References
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 38
06/27/17
Under subfamily references see Clayton & Renvoize; Gould; and Hitchcock & Chase.
Owens, A.G., & S. Riche. Monanthochloë littoralis (Gramineae) in Louisiana. Sida 1: 182, 183.
1963.
Thieret, J. W. Distichlis. P. 28-30 in Barkworth, M.E., K.M. Capels, S. Long, & M.B. Piep, eds.
Flora of North America, north of Mexico. Magnoliophyta: Commelinidae (in part): Poaceae, part
2. New York and Oxford. 2003.
Villamil, C.B. El género Monanthochloë (Gramineae). Estudios morfologicos y taxonomicos con
especial referencia a la especie argentina. Kurtziana 5: 369-391. 1969. [Detailed study; M.
australis transferred to Distichlis.]
4. Tridens Roemer & Schultes, Syst. Veg. 2: 34; [599?] 601. 1817.2
Perennials of grasslands, savannas, and disturbed open places, typically in well drained
soils. Rhizomes often present. Stems erect , cespitose, unbranched (upper stem and panicle
branches viscid when fresh); leaves cauline, few; sheath glabrous or pubescent; ligule a fringe of
short white hairs or ciliate membrane; blade linear, flat or involute. Inflorescence a panicle
[raceme], with the branches spreading and slightly drooping, or ascendant and appressed; secondary
branches spreading. Spikelets laterally compressed, (3) 4-11 (16) flowered; glumes 2, unequal or
subequal, lanceolate to ovate-lanceolate, the first 1-nerved, the second 1-3 (9) nerved; rachilla
disarticulating above glumes; flowers bisexual (the upper ones generally sterile); lemmas elliptic to
oblong-lanceolate, 3-nerved, emarginate, mucronate, smooth, the nerves pubescent proximally
[glabrous]; paleas elliptic, a little shorter than the lemmas, 2-keeled, the keels sometimes ciliate.
2
Roemer and Schultes included as a synonym under Tridens the following: Tricuspis P. Beauv., t.
xv, f. 10. Tricuspis was elsewhere in this work, and the only entry for it in the index is to p. 34.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 39
06/27/17
Lodicules 2, cuneate, rounded, fleshy. Stamens 3; anthers ellipsoid. Ovaries ovoid to ellipsoid,
glabrous; styles 2, separate, short, terminal; stigmas plumose. Fruit a caryopsis, dorsiventrally
compressed, oblong; hilum linear, nearly as long as grain; embryo about 2/3 the length of the
grain. Base chromosome number 10. (Name from Latin tres, three and dens, tooth, referring to the
three-toothed lemmas.) Type species: T. flavus (L.) Hitchc. – Tall Purpletop, tridens.
A genus of 14 species, found in North and South America (and Angola?). The North
American species of Tridens were treated under Triodia in the first edition of Hitchcock’s Manual,
and under Tridens in the 1952 revision by Chase. Triodia is now interepreted as a genus restricted
to Australia. The genera Erioneuron Nash3 and Dasyochloa Willd. Ex Rydb. of the southwestern
U.S. and Mexico, were included in Triodia as well by Hitchcock; Tateoka showed that because of
morphological differences and differing chromosome base number (8 rather than 10), these two
genera should be excluded from Tridens.
In North America there are 10 species, of which 6 occur in the Southeast. Tridens muticus
(Torr.) Nash [T. elongatus (Buckl.) Nash], slim tridens, 2n = 40, occurs from Carroll Co., Arkansas
(Smith) to Utah and California south to Texas (but not Louisiana). Plants in the Southeast are var.
elongatus (Buckley) ShinNers, which has larger glumes with3-7 veins, in contrast to the western
and Mexican typical variety, which has smaller, 1-nerved glumes.
T. flavus, tall purpletop, 2n = 40, is the most widely ranging species, occurs from New Hampshire,
Ontario, and Nebraska, south to Florida and northeastern Mexico; it is widespread in all states in
our region. The living plants are easy to recognize because of the viscid or tacky feel of the upper
3
Erioneuron pilosum (Buckley) Nash has been collected in eastern Kansas, not far from the Arkansas
state line, and might conceivably be found in western Arkansas or northwestern Louisiana.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 40
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stem and inflorescence axis and branches. Southeastern plants with panicles erect and pulvini with
hairs on both sides, are distinguished as var. chapmanii (Small) Shinners; this variety occurs from
Missouri to Virginia southward to Florid and texas.
T. strictus (Nutt.) Nash, longspike tridens, 2n=40, occurs from North Carolina, Illinois, and Kansas
south to Georgia and Texas. It is known from all the southeastern states except Florida. This species
and the preceding one hybridize, yielding T.  oklahomensis, which has been found in Kansas,
Louisiana, and Oklahoma.
Tridens oklahomensis has been recently reported from Kansas (Freeman et al.)
Tridens ambiguus (Ell.) Schult., pine-barrens tridens, 2n=40, occurs from North Carolina
(Virginia?) to central Florida and west to east Texas.
Tridens carolinianus (Steud.) Henrard, North Carolina to northern Florida, west to Louisiana.
Tridens eragrostoides (Vasey & Scribn.) Nash, 2n=40, occurs from Texas to Arizona, south to
Mexico and Cuba; it is known in our region from southern Florida and southern Alabama.
References
Under subfamily references see: Burbridge (1946a, 1946b, 1953), Freeman et al.
Caro, J.A. Rehabilitación de Trichloris Parodi (Gramineae). Dominguezia 2: 21-23. 1981. check
at MO
Crooks, P., & C.L. Kucera. Tridens  oklahomensis (T. flavus  T. strictus), an interspecific sterile
hybrid in the Eragrosteae (Gramineae). Am. Jour. Bot. 60: 262-267. 1973.
McKenzie, P.M., et al. Tridens  oklahomensis (T. flavus  T. strictus) (Poaceae) new to
Louisiana. Sida 12: 424. 1987.
Renvoize. --------------------------- Kew Bull. 33: 525. 1979.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 41
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Tateoka, T. 1961. A Biosystematic Study of Tridens (Gramineae). Am. Jour. Bot. 48: 565-573.
1961.
Valdes-Reyna, J. Tridens. pp. 33-40 in Barkworth, M.E., K.M. Capels, S. Long, & M.B. Piep, eds.
Flora of North America, north of Mexico. Magnoliophyta: Commelinidae (in part): Poaceae, part
2. New York and Oxford. 2003.
5. Triplasis Palisot de Beauvois, Ess. Agrost. 81., pl. XVI, fig. x. 1812
Perennials or annuals of beaches and other natural or disturbed sandy, open places.
Rhizomes often present. Stems erect , cespitose, unbranched, glabrous or pubescent at the nodes;
leaves cauline, few; sheath scabrous but otherwise glabrous; ligule a fringe of short white hairs;
blade linear, flat or involute. Inflorescence a panicle (exserted or partly included in the upper
sheath), with the branches spreading; secondary branches appressed-ascendant. Spikelets laterally
compressed, 2-4 flowered; glumes 2, unequal, lanceolate to ovate-lanceolate, the first a little
shorter (?) than second, 1-nerved, the second about as long as the first lemma, 1-nerved; rachilla
disarticulating above glumes; flowers bisexual (the upper ones generally sterile); lemmas elliptic to
oblong-lanceolate, 3-nerved, deeply emarginate, aristate, smooth, the midvein pubescent, the lateral
nerves silky-pubescent; paleas elliptic, a little shorter than the lemmas, 2-keeled, silky with long
hairs distally. Lodicules 2, cuneate, truncate, fleshy. Stamens 3 (lacking in cleistogamous flowers);
anthers ellipsoid. Ovaries ovoid to ellipsoid, glabrous; styles 2, separate, short, terminal stigmas
plumose. Fruit a caryopsis, dorsiventrally compressed, oblong; hilum linear, nearly as long as
grain; embryo large, typically 1/3 –3/4 the length of the grain. Base chromosome number 10.
Type species: T. americana Beauv., the only species included in the protologue. (Name from Greek
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 42
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triplasios, three-parted.) – Sandgrass.
A genus of two species, found in North and Central America. T. americana, perennial sandgrass
T. purpurea (Walt.) Chapm., purple sandgrass, 2n=40.
The disarticulating culms distinguish this genus from Tridens, to which it is probably closely
related.
References
Hatch, S.L. Triplasis. pp. 41, 42 in Barkworth, M.E., K.M. Capels, S. Long, & M.B. Piep, eds.
Flora of North America, north of Mexico. Magnoliophyta: Commelinidae (in part): Poaceae, part
2. New York and Oxford. 2003.
6. Leptochloa P. Beauvois, Ess. Agrost. 71. 1812, pl. XV, fig. 1.
Perennials or annuals of disturbed, sunny, wet open places. Rhizomes absent. Stems erect ,
cespitose, often branched, glabrous; leaves cauline, few; sheath flatened or terete, glabrous, mostly
longer than the corresponding internode; ligule membranous, ciliate or not; blade linear [auriculate
in L. viscida (Scribn.) Beal], flat (often becoming involute in drying), glabrous or with a few long
hairs at the summit. Inflorescence a panicle, linear to narrowly ovate, with the branches spreading
or ascendant appressed; secondary branches appressed-ascendant; small inflorescences of
cleistogamous spikelets formed in the axils of lower leaves in several species. Spikelets laterally
compressed or sub-terete, (1) 2-10 flowered; glumes 2, unequal to nearly equal, lanceolate,
generally scabrid on the midnerve, the first shorter than the second, 1-nerved, the second nearly as
long as the first lemma, 3-nerved; rachilla disarticulating above glumes; flowers bisexual (the
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 43
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upper ones generally sterile); lemmas narrowly elliptic, 3-nerved, emarginate, aristate, mucronate,
or awnless, glabrous or hairy; paleas elliptic, a little shorter than the lemmas, 2-keeled, glabrous.
Lodicules 2, cuneate, truncate, fleshy. Stamens 2 or 3; anthers ellipsoid. Ovaries ovoid to ellipsoid,
glabrous
or puberulent on the nerves; styles 2, separate, short, terminal stigmas plumose. Fruit a
caryopsis, dorsiventrally compressed or laterally compressed, oblong; hilum linear, nearly as long
as grain; embryo large, typically1/3 –3/4 the length of the grain. Base chromosome number 10.
(Diplachne Palisot de Beauvois, Ess. Agrost. 80. 1812, pl. XVI, fig. IX.) Type species: L. virgata
(L.) Beauv. (see Nash in Britton & Brown, Ill. Fl. No. U.S., ed. 2, 1: 229. 1913; species included by
Beauvois in protologue: Cynosurus capillaceus, Eleusine filiformis, E. virgata.) (Name from Greek
leptos, slender and chloa, grass, referring to the slender spikes.) –Sprangletop.
Leptochloa, treated here in the broad sense to include Diplachne, includes about 45 species,
with about 15 in the New World and about – in the U.S. about 8 species occur in our region.
Recognition of the segregate genus Diplachne has varied, with supporters and detractors.
Clayton & Renvoize treat Diplachne as sect. of Leptochloa, a view supported or at least implied by
numerous other students of grasses.
L. dubia, occurs from Oklahoma to Arizona south to Argentina; it is also in southern Florida (e.g.,
Monroe Co., Key Largo, Kral 51801, MO)
Leptochloa fusca (L.) Kunth ssp. fascicularis (Lam.) N. Snow [L. fascicularis (Lam.) A.Gray],
2n=20, (Massachusetts?) Maryland to Missouri, VA, NC, SC, S to Florida and Louisiana
(statewide, MacRoberts, 1989); rare in Arkansas (Smith)
L. filiformis (Lam.) Beauv., red sprangletop, 2n=20, Arkansas (common; smith) and statewide in
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 44
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Louisiana (MacRoberts, 1989). A hybrid between this species and L. virgata , intermediate in
morphology and posessing “defective pollen,” was reported from Costa Rica by Pohl.
L. panicoides , rare in Arkansas, in the eastern half of the state only (Smith), statewide in Louisiana
(MacRoberts)
L. uninervia in Arkansas known only from Jefferson county (Smith) but statewide in Louisiana
(MacRoberts) and recently reported from Kansas (Freeman et al.)
L. nealleyi southern coastal Louisiana (MacRoberts, 1989)
L. scabra, south-central Louisiana (MacRoberts, 1989)
L. virgata Reported from La. by MacRoberts (1989, by voucher, but no county indicated).
Spikelets laterally compressed, small, 1.5-5 (7) mm long, usually imbricate, on clearly secund
racemes .... sect. Leptochloa
Spikelets subterete, large, 5-15 mm long, usually distant, on indistinctly secund racemes..... sect.
Diplachne.
In Venezuela, a decoction of L. virgata Beauv. is used to relieve stomach ache, and L.
dominguensis Trin. is infused to provide a remedy for seasickness (Morton).
References
Under subfamily references see: Ashraf & Yasmin; Bor; Clayton & Renvoize; Freeman et al.;
Hitchcock & Chase; Morton.
Aslam, Z., M. Saleem, G.R. Sandhu, & R.H. Qureshi. Sodicity effects on growth and chemical
composition of Diplachne fusca. Pakistan Jour. Bot. 11: 123-128. 1979.*
Baskin, C.C., J.M. Baskin, & E.W. Chester. Germination ecology of Leptochloa panicoides, a
summer annual grass of seasonally dewatered mudflats. Acta Oecol. 14: 693-704. 1993.*
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 45
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[Dormancy broken by dry storage for 2 months; light required for germination.]
Bhatti, A.S., A. Ara, G. Sarwar, & J. Wieneke. Some leaf structures related to salt regulation in
kallar grass [Leptochloa fusca (L.) Kunth]. Jour. Pl. Nutr. 15: 313-326. 1992.
Blake, S.T. Plinthanthesis and Danthonia and a review of the Australian species of Leptochloa
(Gramineae). Contr. Queensland Herb. 14: 1-19. 1972.*
Catasus Guerra, L.J. Revisión del género Leptochloa (Poaceae) en Cuba. Acta Bot. Cub. 27: 1-5.
1985.*
Kumar, A. Use of Leptochloa fusca for the improvement of salt-affected soils. Exper. Agri. 32:
143. 1996.*
Latif, F., M. I. Rajoka, & K. A. Malik. Saccharification of Leptochloa fusca (Kallar grass) straw
using thermostable cellulases. Bioresource technology. 50: 107-111. 1994.
Lazarides, M. A revision of the Australian Chlorideae. (Gramineae). Austral. Jour. Bot. Suppl. Ser.
5: 1-51. 1972.*
-----. The genus Leptochloa Beauv. (Poaceae, Eragrostideae) in Australia and Papua New Guinea.
Brunonia 3: 247-269. 1980. [Ten spp.; includes American D. filiformis as an introduction.]
Malik, K.A., Y. Zafar, M. Kloss, & R. Bilal. Dinitrogen fixation associated with roots of kallar
grass (Diplachne fusca) growing in saline soils. Proc. 2nd International Symp. N2 fixation
with non-legumes, Banff, Canada, 1982. 1986.*
Nicora, E.G. Los géneros Diplachne y Leptochloa (Gramineae, Eragrosteae) de la Argentina y
países limitrofes. Darwiniana 33(1-4): 233-256. 1995.*
Ortiz, J.J. Estudio sistemático del género Gouinia (Gramineae, Chloridoideae, Eragrostoideae).
Acta Bot. Mex. 23: 1-33. 1993.*
Pane, H., & M. Mansor. Competition between red sprangletop (Leptochloa chinensis) and rice
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 46
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(Oryza sativa) under different nitrogen levels. Pl. Protection Quarterly 11: 109. 1996.*
Singh, G., H.S. Gill, I.P. Abrol, & S.S. Cheema. Forage yield, mineral composition, nutrient
cycling and ameliorating effects of karnal grass (Leptochloa fusca) grown with mesquite
(Prosopos juliflora) in a highly alkaline soil. Field Crops Research 26: 45-55. 1991.
[Excellent forage on soil with pH of 10.4 in India.]
Snow, N. Noteworthy collections: Nevada. Madroño 39: 158, 159. 1992. [Leptochloa filiformis a
new record for Nevada.]
-----. The phylogenetic utility of lematal micromorphology in Leptochloa and related genera in
subtribe Eleusininae (Poaceae, Chloridoidea, Eragrostideae). Ann. Mo. Bot. Gard. 83: 504529. 1996.
-----. Phylogeny and systematics of Leptochloa P. Beauv. Ph.D. thesis. Washington Univ. 1997.*
-----. Nomenclatural changes in Leptochloa P. Beauvois sensu lato (Poaceae: Chloridoideae) Novon
8: 77-80. 1998.
-----. Caryopsis morphology of Leptochloa sensu lato (Poaceae, Chloridoideae). Sida 18: 271-282.
1998.*
-----. Leptochloa. Pp. 51-60 in Barkworth, M.E., K.M. Capels, S. Long, & M.B. Piep, eds. Flora of
North America, north of Mexico. Magnoliophyta: Commelinidae (in part): Poaceae, part 2. New
York and Oxford. 2003. [Includes Diplachne.]
-----, & G. Davidse. Leptochloa mucronata (Michx.) Kunth is the correct name for Leptochloa
filiformis (Poaceae). Taxon 42: 413-418. 1993.
----- & M.G. Lelong. A first report of Leptochloa scabra Nees (Poaceae) from Alabama. Sida 14:
616, 617. 1991.*
----- & B.K. Simon. Taxonomic status and Australian distiribution of the weedy neotropical grass
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 47
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Leptochloa fusca subsp. uninervia, with an updated key to the Australian Leptochloa
(Poaceae, Chloridoideae). Austrobaileya 5: 299-305. 1999.
Zafar, Y., & K.A. Malik. Photosynthetic system of Leptochloa fusca (L.) Kunth. Pakistan Jour.
bot. 16: 109-116. 1984.*
Zafar, Y., M. Ashraf, & K.A. Malik. Nitrogen fixation associated with roots of kallar grass
(Leptochloa fusca (L.) Kunth. Plant Soil 90: 93-105. 1986.
7. Dinebra Jacquin, Fragm. 77. [Tab.121, fig. 1] 1809.
Annuals of disturbed open places, typically in temporarily wet soils. Rhizomes absent.
Stems erect or spreading simple or branched near basal leaves cauline and basal, few; sheath
glabrous, pubescent at the throat; ligule membranous, ciliate; blade linear, flat, convolute, or
involute. Inflorescence a spike, with widely spaced, spreading spikelets; short secondary branches
sometimes present. Spikelets strongly laterally compressed, 3-6 flowered; glumes 2, subequal,
lanceolate, acute to aristate, about as long as the spikelet, 3-nerved; rachilla disarticulating at base
of each spikelet; flowers bisexual; lemmas narrowly elliptic, acute, 1(-3) nerved, keeled, pubescent
on the nerves, or glabrous; paleas elliptic, a little shorter than the lemmas, 2-keeled, the keels often
ciliolate. Lodicules 2, cuneate, truncate, fleshy. Stamens 3; anthers ellipsoid. Ovaries ovoid to
ellipsoid, glabrous; styles 2, separate, short, terminal stigmas plumose. Fruit a caryopsis, trigonous,
ellipsoid; hilum linear, nearly as long as grain; embryo large, 1/2 the length of the grain. Base
chromosome number ? Type species: D. arabica Jacq. [=D. retroflexa (Vahl) Panz.] , the only
species included. (Name from ----.) -- Common name.
A genus of three species, native to Africa and southern Asia. Dinebra retroflexa (Vahl) Panz.,
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 48
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viper-grass, 2n = 20, is native to tropical Africa and southern Asia. It is adventive in North Carolina
[Castanea 52: 104, 1987]. Elsewhere in North America, it has been reported from ore piles at the
Port of Baltimore, Maryland. The genus was not included by Peterson et al.
References
Under subfamily references see
Barkworth, M.E. Dinebra. Pp. 63, 64 in Barkworth, M.E., K.M. Capels, S. Long, & M.B. Piep,
eds. Flora of North America, north of Mexico. Magnoliophyta: Commelinidae (in part): Poaceae,
part 2. New York and Oxford. 2003.
Phillips. ------------------ Kew Bull. 28: 411-418. 1973.
8. Eragrostis N.M. von Wolf, Gen. Pl. Vocab. Char. Def. 23. 1776.*
Perennials or annuals of disturbed open places, typically in well drained soils, but also in
damp places. Rhizomes often present; sometimes stoloniferous. Stems erect , spreading, or
decumbent (repent, rooting at the nodes in E. reptans and E. hypnoides), simple or branched near
basal leaves cauline and basal, few; sheath longer or shorter than internode, glabrous, pubescent at
the throat; ligule a fringe of short white hairs; blade linear, flat, convolute, or involute.
Inflorescence a panicle, open or somewhat contract with the branches ascendant. Spikelets strongly
laterally compressed, 3-30 flowered; glumes 2, subequal or unequal, lanceolate to ovate-lanceolate,
1 (2 or 3) nerved; rachilla persistent, the glumes, lemmas and paleas disarticulating acropetally to
release the caryopses [persistent, the inflorescence ripening intact in E. tef]; flowers bisexual
(unisexual in E. reptans, the plants dioecious); lemmas elliptic to oblong-lanceolate, 3 (-5) nerved;
paleas elliptic, a little shorter than the lemmas, 2-keeled, the keels often ciliolate, persistent on the
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 49
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rachilla or falling with the lemmas. Lodicules 2, cuneate, truncate, fleshy. Stamens 2 or 3; anthers
ellipsoid. Ovaries ovoid to ellipsoid, glabrous; styles 2, separate, short, terminal stigmas plumose.
Fruit a caryopsis, subterete to cylindric, oblong to broadly ellipsoid; hilum linear, nearly as long as
grain; embryo large, typically1/3 –3/4 the length of the grain. Base chromosome number 10.
(Eragrostis Host, nom. superfl.; Erochloë Raf.; Acamptoclados Nash; Erosion Lunell;
Diandrochloa de Winter). Type species: E. eragrostis (L.) von Wolf [= E. minor Host; see Koch,
1978]. (Origin of name uncertain; perhaps from Greek, eri-, very or much, and agrostis, the name
of a grass genus of the Poöideae; see further discussion by Peterson). – Lovegrass.
A genus of about 350species, worldwide in tropical, subtropical and warm temperate regions. There
are about 49 species in the US, and 29 species in the Southeastern States (Peterson). About 40
species occur in India and surrounding countries (Bor). The genus includes a good deal of diversity
in habit and form, with open and narrow inflorescences, and annual and perennial species. The lack
of awns, the strongly three-nerved lemmas, and distinctive pattern of articulation, in which the
lemmas fall before the paleas leaving a partly bare rachilla, is a good hallmark of the genus. Some
have suggested the genus is polyphyletic, but this has not been substantiated.
Four sections were recognized by several workers (Roshevits, Clayton & Renvoize).
Sect. Cataclastos Doell. (Macroblepharus Phill.)
Sect. Pteroëssa Doell. (Megastachya Beauv.)
Sect. Platystachya Benth.
Sect. Lappula Stapf. These sections were not followed or discussed by Peterson.
E. arida Hitchc. western Arkansas (Smith)
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 50
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E. frankii, northwestern Arkansas (Smith)
Following 2 species distinct by having cup shaped glands on pedicels, leaf margins and culm
E. minor Host [E. poaeoides Beauv. ex Roemer & Schultes] This species is worldwide in temperate
and tropical regions, and found in all the southeastern states. Louisiana (Allen); eastern Arkansas
Smith) lemmas 1.5-2 mm long
E. cilianensis (All.) Vignoli-Lutati ex Janch.4, stink grass, 2n=20, is widely distributed in tropical
and warm temperate regions worldwide. Known from all the southeastern states (scarce in Georgia
and Florida), it is distinguished from E. minor by longer lemmas (2-2.8 mm). The fresh plants have
a stinking odor, described as soapy or suggestive of Datura stramonium L. (Solanaceae). Plants of
this species usually have at least some saucer-shaped glands on the foliage, and often on the
glumes, lemmas and inflorescence branches as well.
E. curtipedicellata Buckley, Howard Co., Arkansas (Smith)
E. pilosa (L.) Beauv. Louisiana (Allen); Arkansas , northwest, rare (Smith). Native of Europe but
now widely distributed in tropical and warm-temperate regions worldwide.
E. tenella (L.) Beauv. ex R. & S., Old World annual introduced in the southeast, Texas, and south
to South America.
E. pectinacea (Michx.) Steud., 2n=40, 60, is found throughout the U.S., south through
Mesoamerica and the West Indies to Argentina. A tufted annual, it is a common weedy species of
disturbed damp places.
E. tephrosanthos Schultes Scattered in U.S., LA (Allen), south to tropics (Koch). closely related
to preceding, differing chiefly in its spreading rather than appressed pedicles. Koch stated he had
4
Mosher (Univ. Ill. Agr. Exper. Sta. Bull. 205: 381. 1918) was possibly the earliest author accepting the
new combination suggested by Lutati; see also Shinners. Rhodora 56: 26, 27. 1954, who attributed it to
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 51
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not observed hybrids between them; Pohl noted that immature specimens can be difficult to
distinguish. The species was not recognized by Peterson
E. tracyi A.S. Hitchc., an annual Florida endemic, closely related to E. tephrosanthos
E. cumingii Steud. [E. simplex Scribn.] occurs in Florida, southern Georgia, and Alabama. It is
probably an introduction from southeast Asia, where most of its close relatives occur (Koch 1978).
Koch noted that it is cleistogamous, as evidenced by its frequently dehisced anthers and lack of
exserted stamens or stigmas. The closely related Australian E. elongata (Willd.) Jacq. has been
collected in Florida a century ago (Simpson s.n. in 1897, US; exact locality not known)
E.curvula (Schrad.) Nees, weeping lovegrass, is native to southern Africa. Apomixis is well known
in the complex of Eragrostis curvula in South Africa (Covas & Cairnie) , although sexual
reproduction also occurs. This species is taxonomically complex; several segregates have been
named, and chromosome counts range from 2n = 20 to 80, as well as 42 and 63. A tufted perennial,
the species is widely planted for forage and to stabilize soils. It has reached as far north as southern
New England and Illinois, and is found in all states in the southeast, as well as Texas and the
southwestern states. The species is extensively planted for forage and erosion control in the
southern Great Plains and southwest, as well as in argentina and southern Africa. Analysis of
electrophoretic data from seed protein showed three major groupings in the species, corresponding
to the cultivars ‘Weeping,’ ‘Boer,’ and ‘Robusta.’ The distinctness of E. lehmanniana Nees,
Lehmann lovegrass, was also supported by this eletrophoretic study (Poverene & Voigt).
E. lugens Louisiana (Allen)
E. hirsuta, similar to lugens; Arkansas, statewide (Smith)
E. capillaris northern Arkansas, widespread; Louisiana , rare (Allen)
Hitchcock.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 52
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E. elliottii Wats. [E. acuta A.S. Hitchc.], Se. U.S., Caribbean, Mexico. Previously recognized as an
endemic of western Florida, Koch (1987) concluded that such plants were within the range of
variation for E. elliottii, agreeing with Harvey (1948)
E. atrovirens (Desf.) Trin. ex Steud. [E. chariis sensu Hitchc., not Schultes; cf. Koch 1978.], thalia
lovegrass, 2n = 60, is a naturalized species in Florida5, southern Alabama, and southern
Mississippi. A cespitsoe perennial, it is native to northern Africa.
E. pilifera Scheele, Louisiana, fide Hitchcock (Allen could not confirm)
E. refracta, southeastern U.S.
E. intermedia, Arkansas, statewide (Smith)
E. caroliniana
E. secundiflora Presl ssp. oxylepis (Torrey) S.D. Koch [E. oxylepis Torrey], red lovegrass, 2n = 40,
ranges from South Carolina to Missouri and Colorado, south to northern Mexico. Ssp. secundiflora
occurs in southern Mexico, Central and South America (Koch 1978; Peterson). The two subspecies
differ in the shape of the caryopsis and the pubescence of the sheath.
E. bahiensis (Schrad.) Schultes (E. expansa Link), Bahia lovegrass, is known from the Gulf Coast
region of Louisiana, Mississippi, Alabama, and Florida6; it is native to Bolivia, southern Brazil and
Argentina.
E. ciliaris (L.) R. Br., 2n=20, 40, is more reminiscent of a pooid grass such as Phleum than other
Eragrostis with branched inflorescences. The stiff conspicuous hairs on the nerves of the paleas
give the inflorescence a bristly appearance. It is a native of the Old World, introduced from Florida
to Louisiana (St. Tammany and Vernon parishes) south to Peru and Brazil (Pohl).
5
6
Hillsboro Co., 0.7 mi W of Mango, R.E. Perdue Jr. 1743 (ILL), det. J.R. Swallen.
Escambia co., vic. Pensacola, 12 March 1953, Ahles 7185 (ILL)
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 53
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E. glomerata (Walt.) L.H. Dewey, 2n=20, occurs from the southeastern U.S. south to Bolivia and
Argentina. Louisiana (Allen); Arkansas, statewide (Smith)
Eragrostis. hypnoides (Lam.) B.S.P, teel lovegrass, 2n=20, occurs throughout the U.S. (although
sporadic in much of the west) and south to Argentina. It is a fairly common species of muddy
shores in the southeast. The stolonifreous habit, and open, sparsely branched inflorescences suffice
to distinguish it from congeners. A close relative, E. reptans (Michx.) Nees, creeping lovegrass, 2n
= 60, occurs primarily in the Great Plains States (extending east to western Florida and Middle
Tennessee) south to northern Mexico. It has unisexual flowers and is dioecious. Both species occur
on wet shores, and may be especiaaly abundant on drawn down reservoirs.
E. trichodes, central Arkansas, 2 counties (Smith)
E. spectabilis Arkansas, statewide (Smith)
E. peregrina
E. plana Nees, 2n = 20, has been collected as a waif at the Santee Wool Combing Mill in South
Carolina (Gould & Lonard). It is native to southern Africa.
E. scaligera Salzm. ex Steud.,2n=40, a species of Brazil and Frech Guiana, has been collected in
Collier and Lee counties, Florida (Koch 1975; Peterson).
E. gangetica (Roxb.) Steudel, native of tropical Asia and Africa, has been collected in Mississippi
(Jackson Co., Fontainebleau, 24 Sept. 1953, R.L. Diener 1390, ILL) and in Florida (Collier Co., see
Koch 1978). E. virescens Presl, a Chilean species, was collected in Appalachicola, Florida by A.W.
Chapman, probably in the late 19th Century (Koch 1978).
Eragrostis tef (Zucc.) Trotter [E. abyssinica (Jacq.) Link], teff or t’ef, 2n=40, is a grain in
northeastern Africa. It is little cultivated elsewhere, but is a staple grain in Ethiopia, where it
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 54
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accounts for half the annual grain production (Purseglove). The seeds are about 1-1.5 mm long,
much smaller than familiar grains such as wheat or rice, and about the size of millet. The grain is
about 9 % protein, contains little gluten, but is high in iron and calcium. It is used to make a
flatbread or pancake called ingera, a staple of Ethiopian cuisine. Teff flour is mixed with water and
allowed to ferment for 1-2 days, before being baked in covered trays. The resulting flatbread is
about 50 cm in diameter, a few mm thick, and elastic with a slightly sour taste. A self-pollinated
species, teff is also grown as an annual hay crop in India, southern Russia and Ukraine, southern
and eastern Africa, southeastern Australia, and South America. The seeds germinate rapidly,
producing a forage crop in only 9 weeks in Kenya (Bogdan). In recent decades teff has been planted
as forage in several western states. It has great potential as a forage crop, wit crude protein analyses
ranging from 14-19 percent (Boe et al., 1986). In South Dakota, teff has been infested at rates up to
30 percent by a stem borer, Eurytomocharis eragrostidis Howard, which limits its potential value as
a forage until control methods can be devised (Boe et al. 1992). It is similar to E. minor and E.
cilianensis in having glands on the pedicels, leaves and culms and may be closely related. Or
perhaps derived from E. pilosa (Clayton & Renvoize). Various strains of teff show considerable
genetically based variation in features of panicle size, grains per panicle, grain weight, and tiller
number; this variation can be used for improvement of the crop (Tefera et al.).
In addition to Eragrostis tef, a number of species are used as forage, including E. pilosa, E.
megastachya, E. minor, and E. suaveolens Beck (Roshevits).
References
Under subfamily references see Austin, Bor, Clayton & Renvoize; Harlan (1986);
Hitchcock (1920), Hitchcock & Chase; Purseglove.
??????. E. poaeoides in Tenn. Castanea 46: 334. 1981.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 55
06/27/17
Ahles, H.E., C.R. Bell, & A.E. Radford. Species new to the flora of North or South Carolina.
Rhodora 60: 20-32. 1958. [E. lugens.]
-----, and A. E. Radford. Species new to the flora of North Carolina. Jour. Elisha Mitchell Sci. Soc.
75: 140-147. 1959. [E. curvula.]
Ahring, R.M., N.L. Dunn, Jr., & J.R. Harlan. Effect of various treatments in breaking seed
dormancy in sand lovegrass, Eragrostis trichodes (Nutt.) Wood. Crop Sci. 3: 131-133.
1963.*
Amarasinghe, V., & L. Watson. Taxonomic significance of microhair morphology in the genus
Eragrostis Beauv. (Poaceae). Taxon 39: 59-65. 1990.
Atkins, M. D., & Longley, A. J. Sand lovegrass (Eragrostis trichodes) makes a come-back. Soil
Conserv. 15: 138-141. 1950. *
Ball, C. R. An anatomical study of the leaves of Eragrostis. Proc. Iowa Acad. Sci. 4: 138-146. pl.
16-18. 1897.*
Beal, W. J. Some monstrosities in spikelets of Eragrostis and Setaria. Bull. Torr. Bot. Club 27:
85, 86. 1900.
Bekele, E., & R. Lester. Biochemical assessment of the relationships of Eragrostis tef (Zucc.)
Trotter with some wild Eragrostis species (Gramineae) Ann. Bot., II, 48: 717-725. 1981.*
Berhe, T., L.A. Nelson, M.R. Morriss, & J.W. Schmidt. Inheritance of phenotypic traits in tef. I.
Lemma color. Jour. Hered. 80: 62-65. 1989; II. Seed color. Ibid, 65-67; III, Panicle form,
Ibid, 67-70.
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zone. Geobios 14: 131-133. 1987.*
Leigh, J. H. Leaf anatomy in certain strains of Eragrostis (Beauv.). Jour. S. Afr. Bot. 27(3): 141146. 1961.*
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Mattei, G.E., & C. Tropea. Ricerche e studi sul genere Eragrostis, in rapporto ai nettarii estanuzali.
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McDaniel, B., & A. Boe. A new host record for Eurytomocharis eragrostidis Howard
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-----, R.C. Pickett, & R.L. Davis. Genetic variability and interrelationship of characters in teff,
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Eragrostis tef (Zucc.) Trotter. Crop Sci. 5: 155-157. 1965.
Mimeur, G. Sur l'origine phylétique du genre Eragrostis. Mus. Natl. d'Hist. Nat. B. (sér. 2) 24:
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Morris, E.J. The cell wall of Eragrostis tef: variations in chemical composition and digestibility.
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Nicora, E. G. Contribución al estudio histológico de las glandulas epidermicas de algunas especies
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Nieto Feliner, G. Eragrostis curvula (Schrader) Nees: a new grass for the Spanish flora. An. Jardin
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(Poaceae). Pl. Syst. Evol. 166(3-4): 173-182. 1989.
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Rubio, H.O., M.K. Wood, M. Cardenas, & B.A. Buchanan. Effect of polacrylamide on seedling
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emergence of three grass. species. Soil Sci. 148: 355-360. 1989.
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1963.*
Stover, E. L. The embryo sac of Eragrostis cilianensis (All.) Link. Ohio Jour. Sci. 37: 172-184. pl.
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Toole, V.K., & H.A. Borthwick. The photoreaction controlling seed germination in Eragrsotis
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1971.
-----. Induced seed dormancy in weeping lovegrass, Eragrostis curvula. Crop Sci. 13: 76-79. 1973.
----- & E. C. Bashaw. Apomixis and sexuality in Eragrostis curvula. Crop Sci. 12(6): 843-847.
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_____ & _____. Facultative apomixis in Eragrostis curvula. Crop Sci. 16: 803-806. 1976.*
-----, B.L. Burson, & R.A. Sherman. Mode of reproduction in cytoypes of lehmann lovegrass.
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Wang, S., & B. Sun. New taxa of Eragrostis in Yunnan, China. Acta bot. Yunnanica 11(3): 303307. 1989.
Wester, D.B., B.E. Dahl, & P.F. Cotter. Effects of pattern and amount of simulated rainfall on
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Wright, L.N. Seed dormancy, germination environment, and seed structure of Lehmann lovegrass,
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Eragrostis lehmanniana Nees. Crop Sci. 13: 432-435. 1973.
9. Neeragrostis Bush, Trans. Acad. Sci. St. Louis 13: 178. 1903.
Low annuals of damp sunny shorelines. Rhizomes and stolons absent. Stems decumbent
(repent, rooting at the nodes), mat-forming, simple; leaves cauline and basal, few; sheath shorter
than internode, glandular; ligule a fringe of short white hairs; blade lance-linear, flat or
conduplicate. Inflorescence a panicle, somewhat contracted with the branches ascendant-appressed.
Spikelets ovate to linear, strongly laterally compressed, 15-60 flowered; glumes 2, unequal,
lanceolate to ovate-lanceolate, thin , translucent, the first 1-nerved, the second 1-3 nerved; rachilla
disarticulating above glumes; flowers unisexual, the plants dioecious; lemmas elliptic to oblonglanceolate, awned or mucronate, 3-nerved, glabrous or hairy; paleas elliptic (lacking or vestigial in
carpellate spikelets), a little shorter than the lemmas, 2-keeled, the keels often ciliolate. Lodicules
2, truncate, fleshy. Stamens 3; anthers narrowly ellipsoid. Ovaries ovoid to ellipsoid, glabrous;
styles 2, separate, short, terminal stigmas plumose, persistent on the maturing achene. Fruit a
caryopsis, terete, oblong; hilum linear, nearly as long as grain; embryo large, 1/3 –1/2 the length
of the grain. Base chromosome number 10. Type species: Poa reptans Michaux = N. reptans
(Michx.) Nees. (Name from Eragrostis, q.v.). – Ponygrass, Creeping Lovegrass.
A genus of two species, native to North and Central America Neeragrostis contrerasii
(Pohl) Peterson, is found in Central America. N. reptans occurs from Kentucky, western
Tennessee, Arkansas, and Louisiana west to Texas and South Dakota. Disjunct populations are
known from Florida and northeastern Mexico (Gould).
Neeragrostis is believed to be closely related to Monanthochloë and Distichlis (Nicora). It
has long been included in Eragrostis, by Hitchcock and Chase and others. Neeragrostis was
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 66
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accepted by Gould and by Peterson et al., although synonymized in Eragrostis without comment by
Clayton & Renvoize.
References:
Under subfamily references see Gould; Peterson et al.
Nicora, Revista Argent. Agron. 29: 1-11. 1962.*
10. Eleusine Gaertner, Fruct. 1: 7. 1788*.
Annuals [perennials] of disturbed open places, typically in somewhat moist rather than
well-drained soils. Rhizomes absent [present]; stolons absent [present, indurate]. Stems erect to
spreading, unbranched, terete to compressed; leaves cauline, few; sheaths longer or shorter than the
internodes, flattened, sharply keeled, glabrous [lanate basally]; ligule membranous, with lacerateciliate margin; blade linear, flat or slightly plicate, with strong midrib, keeled basally. Inflorescence
of 1-several secund racemose branches, these branches ± digitately or spicately arranged at the top
of the stem; rachis triquetrous or flattened. Spikelets strongly laterally compressed, ovate to
ellipsoid, 3-6 [15] flowered, arranged in two rows on opposite sides of the rachis, sessile or shortpedicellate; glumes 2, unequal, lanceolate, the first 1-nerved, about as long as the first lemma, the
second 3-7 nerved, slightly longer; disarticulation above glumes; flowers bisexual (the upper ones
sometimes staminate, or sterile); lemmas oblong-ovate, 3-5 (7) nerved, stongly keeled, the nerves
and margins ciliate; paleas elliptic, about ¾ as long as the lemmas, acute, strongly 2-nerved,
glabrous. Lodicules 2, lobed or truncate (acuminate?), fleshy. Stamens 3; anthers oblongellipsoid. Ovaries obconical to ellipsoid, glabrous; styles 2, separate, short, terminal stigmas
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 67
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plumose. Fruit an achene (pericarp thin, free, membranous or papery), ellipsoid [globose],
transversely ridged and rugose; hilum linear, nearly as long as grain; embryo large, typically1/3
–3/4 the length of the grain. Base chromosome number 9. (synonym). Type species: Type species:
E. coracana (L.) Gaertner was designated by ). (Named for the ancient Greek town of Eleusis,
where the goddess of harvests was worshipped.) – Goosegrass.
A genus of about six species, one native to South America, the others to tropical regions of
the Old World.
E. indica is widely naturalized in North America.
E. tristachya (Lam.) Lam., native of-----, has been collected as a ballast plant during the
Nineteenth Centruy in New Jersey, Pennsylvanina, Florida, and Alabama7. There is no evidence to
suggest it has ever persisted or naturalized anywhere in North America.
The foliage and seeds of E. indica are reported to contain cyanogenic glycosides (Bor; Morton).
Under some conditions, the plant may accumulate toxic levels of nitrates (Morton). the seeds are
reportedly eaten by humans, or fed to poultry. when young, the plants provide good forage for
cattle. The tender young plants are eaten raw or cooked in Java. The plant has provided a number of
folk remedies. A decoction is reported to counteract dysentery, diarrhea and convulsions, or to
function as a diuretic. A water extract of the whole plant has been used as shampoo or hair tonic
(Morton).
Eleusine coracana (L.) Gaertn., finger millet, 2n=36, is a species widely cultivated in tropical
7
The following serve as vouchers for these states: New Jersey, Communipaw Ferry, 2 Sept. 1880, A.
Brown (MO); Pennsylvania: below Philadelphia, Sept. 1874, C.F. Parker 9400 (MO); Alabama: Mobile,
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 68
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Africa and southern Asia. It is closely related to E. africana and E. indica. It is perhaps derived
from E. africana, having originated in the region of Uganda (Purseglove). E. coracana is a shortday plant, a 12-hour photoperiod being optimum. Flowering takes place over a period of 8-10 days,
with the maximum number of flowers open on the third day. flowering commences at the top of the
spike and proceeds toward the base; in a spikelet, flowering proceeds acropetally, the lowest floret
opening first. The plants are highly self-compatible and only about 1-3 % cross-pollination occurs
(Fryxell).
References
Under subfamily references see: Bor; Clayton & Renvoize; Fryxell; Harlan (1986); Hitchcock &
Chase; Morton; Purseglove; Wanous.
Gasser M, Vegetti AC, 1997., Inflorescence typology in Eleusine indica and Eleusine tristachya
(Poaceae). Flora (Germany) 192. (1): 17-20 (1997) - illus.
Hilu KW, 1980., Noteworthy collections: Eleusine tristachya (Lam.) Lam. Madroคo 27. (4): 177 178 (19
Hilu KW, Johnson JL, 1997., Systematics of Eleusine Gaertn. (Poaceae: Chloridoideae):
chloroplast DNA and total evidence. Ann. Missouri Bot. Gard. 84. (4): 841-847 (1997) –
Hilu, K.W. Validation of the combination Eleusine coracana subspecies africana (KennedyO'Byrne) Hilu et Dewet. Phytologia 76. (5): 410-411. 1994.
Hilu KW, 1981., Taxonomic status of the disputable Eleusine compressa (Gramineae). Kew Bull.
36. (3): 559 - 563 (1981) - illus.
Hilu KW, De Wet JMJ, 1976., Domestication of Eleusine coracana. Econ. Bot. 30. (3): 199 - 208
(1976) - Illus., maps. Hilu KW, De Wet JMJ, 1976., Racial evolution in Eleusine coracana ssp.
coracana (finger millet). Amer. J. Bot. 63. (10): 1311 - 1318 (1976) –
Sept. 1897, C. Mohr 6379 (MO); Florida: Lake City, 1893, A.W. Bittung 780 (MO).
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 69
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Hiremath, S.C.,& S.S. Salimath. Poaceae: on the origin of Eleusine africana. Bothalia 21: 161162. 1991.
Hiremath SC, Chennaueeraiah MS, 1982., Cytogenetical studies in wild and cultivated species of
Eleusine (Gramineae). Caryologia 35. (1): 57-69 (1982)- illus., chrom. nos.
Gupta RK, Saxena SK, 1980., Ecological studies on Eleusine compressa: a potential grass for
sheep pasturage in the arid zone. Ann. Arid Zone 19. (1 - 2): 1 - 13 (1980) –
Bisht MS, Mukai Y, 2002., Genome organization and polyploid evolution in the genus Eleusine
(Poaceae). Pl. Syst. Evol. 233. (3-4): 243-258 (2002) - illus., col. illus.
Chavan, V. M., N. Gopalkrishna, & B. T. Khadilkar. Blooming and anthesis in nagil (Eleusine
coracana (Linn.) Gaertn.). Poona Agr. Col. Mag. 46: 175-179. 1955.
Clifford, H. T. Vivipary in Eleusine indica (L.) Gaertn. Nature 184(4702, sup. 24): 1888. 1959.
Cummins, M. P. Development of the integument and germination of the seed of Eleusine indica.
Bull. Torrey Club 56: 155-162. f. 1-7. 1929.
Divakaran, K. Inheritance of toppling in Eleusine coracana (Gaertn.). Madras Agr. Jour. 47(2):
74, 75. 1960.
Divakaran, K. Leaf arrangement in ragi (Eleusine coracana Gaertn.) and its significance. Madras
Agr. Jour. 46(7): 260-263. 1959.
Figliola, S.S., N.D Camper, & W.H. Ridings. Potential biological control agents for goosegrass
(Eleusine indica). Weed sci. 36: 830-835. 1988. Note Includes references. eleusine indica.
cochliobolus setariae. pyricularia. fungal diseases. weed control. biological control.
pathogenicity. OtherSubject piricularia grisea Other Author Abstract Two leaf-spotting
pathogens, [Bipolaris setariae (Saw.)] and [Piricularia grisea (Cke.) Sacc.], were isolated from
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 70
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severely infected goosegrass plants. Pathogenicity tests conducted at 28 C showed that both fungi
were 100% effective in infecting goosegrass when given a 72-h dew period. Dew period
temperature and duration requirements were tested by inoculating 2-week-old plants with
conidial suspensions of each fungus and incubating them in dew chambers (100% relative
humidity). Disease index increased as dew period duration increased for both fungi and at all
temperatures tested. Infection occurred at all temperatures with an optimum of 24 to 28 C for B.
setariae and 28 C for P. grisea. In host range tests, representative plants of the Fabaceae,
Malvaceae, Poaceae, and Solanaceae were inoculated with suspensions of either 20000 or 60000
spores/ml of each fungus and placed in growth chambers at 28 C. Infection was limited to
members of the Poaceae. Sorghum, showed a hypersensitive response to B. setariae. Both
cultivars of corn developed light symptoms in response to both fungi at 20000 and 60000
spores/ml.
Fulwider, J. R., & R. E. Engel. Pre-emergence control of goosegrass [Eleusine indica] in turf areas.
Northeast. Weed Control Conf. Proc. 12: 138-139. 1958. [CIPC and monuron]
_____ & _____. The effect of temperature and light conditions on the germination of goosegrass
(Eleusine indica). Weed Sci. Soc. Amer. Abs. 1958: 45.
_____ & _____. . The effect of temperature and light on germination of seed of goosegrass,
Eleusine indica. Weeds 7(3): 359-361. 1959.
Ganeshaiah, K.N., & R. Uma Shaanker. Evolution of reproductive behaviour in the genus
Eleusine. Euphytica 31: 397-404. 1982. [Compatibility.]
Gasser, M., & A.C. Vegetti. Inflorescence typology in Eleusine indica and Eleusine tristachya
(Poaceae). Flora (Jena) 192(1): 17-20. 1997.
Gray, R. A. Breaking the dormancy of peach seeds and crab grass seeds with gibberellins. (Abs.)
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 71
06/27/17
Plant Physiol. 33(suppl.): xl-xli. 1958.
Hawton, D. Temperature effects on Eleusine indica and Setaria anceps grown in association. II
Weed Res 20(5): 261-266. 1980. . [Competitive abilities].
Hilu, K. W. & J. M. J. deWet. Origin and taxonomy of Eleusine coracana (finger millet). Bot.
Soc. Am. Abstr. Pap. Meetings Oregon State Univ. Aug. 1975: 55. 1975.
----- & -----. Domestication of Eleusine coracana. Econ. Bot. 30: 199-208. 1976.
----- & -----. Racial evolution in Eleusine coracana ssp. coracana (finger millet). Am. Jour. Bot.
63: 1311-1318. 1976.
-----, -----, & J. R. Harlan. Archaeobotanical studies of Eleusine coracana ssp. coracana (finger
millet). Am. Jour. Bot. 66: 330-333. 1979.
Hilu, K.W. Evolution of finger millet: evidence from random amplified polymorphic DNA.
Genome. v. 38: 232-238. 1995. Abstract Finger millet (Eleusine coracana ssp. coracana) is an
annual tetraploid member of a predominantly African genus. The crop is believed to have been
domesticated from the tetraploid E. coracana ssp. africana. Cytogenetic and isozyme data point to
the allopolyploid nature of the species and molecular information has shown E. indica to be one
of the genomic donors. A recent isozyme study questioned the proposed phylogenetic
relationship between finger millet and its direct ancestor subspecies africana. An approach using
random amplified polymorphic DNA (RAPD) was employed in this study to examine genetic
diversity and to evaluate hypotheses concerning the evolution of domesticated and wild annual
species of Eleusine. Unlike previous molecular approaches, the RAPD study revealed genetic
diversity in the crop. The pattern of genetic variation was loosely correlated to geographic
distribution. The allotetraploid nature of the crop was confirmed and molecular markers that can
possibly identify the other genomic donor were proposed. Genotypes of subspecies africana did
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 72
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not group closely with those of the crop but showed higher affinities to E. indica, reflecting the
pattern of similarity revealed by the isozyme study. The multiple origin of subspecies africana
could explain the discrepancy between the isozyme-RAPD evidence and previous information.
The RAPD study showed the close genetic affinity of E. tristachya to the E. coracana-E. indica
group and underscored the distinctness of E. multiflora.
Hilu, K.W. Identification of the "A" genome of finger millet using chloroplast DNA. Genetics.
Jan 1988. v. 118 (1) :163-167. ill.
Hilu, K.W. Ribosomal DNA variation in finger millet and wild species of Eleusine (Poaceae).
Theoretical and applied genetics 83(6/7): 895-902. 1992. Abstract Finger millet is an
important cereal crop in the semi-arid regions of Africa and India. The crop belongs to the
grass genus Eleusine, which includes nine annual and perennial species native to Africa
except for the New World species E. tristachya. Ribosomal DNA (rDNA) variation in
finger millet and related wild species was used to provide information on the origin of the
genomes of this tetraploid crop and point out genetic relationships of the crop to other
species in the genus. The restriction endonucleases used revealed a lack of variability in
the rDNA spacer region in domesticated finger millet. All the rDNA variants of the crop
were found in the proposed direct tetraploid ancestor, E. coracana subsp. africana. Wild
and domesticated finger millet displayed the phenotypes found in diploid E. indica.
Diploid Eleusine tristachya showed some similarity to the crop in some restriction sites.
The remaining species were quite distinct in rDNA fragment patterns. The study supports
the direct origin of finger millet from subspecies africana, shows E. indica to be one of
the genome donors of the crop, and demonstrates that none of the other species examined
could have donated the second genome of the crop. The rDNA data raise the possibility
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 73
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that wild and domesticated finger millet could have originated as infraspecific polyploid
hybrids from different varieties of E. indica.
Hiremath, S.C., & M.S. Chennaveeraiah. Cytogenetical studies in wild and cultivated species of
eleusine (Gramineae). Caryologia 35: 57-69. 1982.*
----- & S.S. Salimath. Quantitative nuclear DNA changes in Eleusine (Gramineae). Pl. syst.
evol. 178(3/4): 225-233. 1991. [Incl. interspecific and intraspecific hybridization.]
----- & -----. The 'A' genome donor of Eleusine coracana (L.) Gaertn. (Gramineae). Theoretical
and applied genetics. 84 (5/6): 747-754. 1992. In an attempt to discover 'A' and 'B'
genome donor(s) to finger millet, Eleusine coracana, or its progenitor species, E. africana
(both allotetraploid 2n = 4x = 36), five diploid species, E. indica, E. floccifolia, E.
multiflora, E. tristachya and E. intermedia, were crossed to finger millet and its progenitor
taxon. Crosses were successful only with E. coracana. Three combinations of triploid
hybrids E. coracana X E. indica, E. coracana X E. floccifolia, and E. coracana X E.
multiflora were obtained and analysed. Meiotic behaviour was perfectly normal in
parental species. The regular number of 18 bivalents in E. coracana, 9 bivalents in E.
indica, E. intermedia, E. tristachya and E. floccifolia and 8 bivalents in E. multiflora were
invariably noticed. In E. coracana X E. indica hybrids a mean chromosome pairing of
8.84(I) + 8.80(II) + 0.03(III) + 0.10(IV) per cell was found. About 86.5% of the cells
showed the typical 9(I) + 9(II) configuration, suggesting that E. indica (AA) is one of the
diploid genome donors to cultivated species E. coracana. A mean chromosome pairing of
11.08(I) + 7.63(II) + 0.16(III) + 0.04(IV) per cell was found in E. coracana X E.
floccifolia hybrids. Two to ten bivalents and varying numbers of univalents were seen in
55% of the cells. About 45% of the cells showed the 9(I) + 9(II) configuration. Various
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 74
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evidence suggests that perennial E. floccifolia is a primitive member of the 'A' genome
group of Eleusine species, and it may not be a genome donor to E. coracana. In E.
coracana X E. multiflora hybrids (2n = 26) mean chromosome pairing of 21.45(I) +
1.97(II) + 0.13(III) + 0.04(IV) per cell was found. About 91% of the cells were observed
to have 20-26 univalents. Only a small percentage of the cells contained bivalents or
multivalents. This pairing behaviour indicates that E. multiflora lacks genomic Abstract
coracana. Genomically E. multiflora is a distinct species and a genomic symbol of 'C' is
assigned to it. Identification of the 'B' genome donor species to cultivated millet E.
coracana remains elusive.
Johnson, R.M., & W.D. Raymond. The chemical composition of some tropical food plants: 1.
Finger millet and bulrush millet. Trop. Sci. 6: 6-11. 1964.*
Kennedy-O’Byrne, J. Notes on African grasses: XXIX: a new species of Eleusine from Africa and
south Africa. Kew Bull. 1: 65-72. 1957.
Kimata, M., M. Kanoda, & A. Seetharam. Traditional and modern utilizations of millets in Japan.
Environm. Ed. Stud. Tokyo Gakugei Univ. 8: 21-29. 1998. [E. coracana used for mochi
(cakes) and porridge.]
-----., & S. Sakamoto. Utilization of several species of millet in Eurasia. Bull. Field Studies Inst.
Tokyo Gakugei Univ. 3: 1-12. 1992. [E. coracana an important grain in India utilized in 8
different cooking styles.]
Koba H, Matsumoto M, 2003., (Eleusine tristachya (Lam.) Lam. (Gramineae), newly naturalized
in Japan.) Bull. Kanagawa Prefect. Mus., Nat. Sci. no.32. 69-70 (2003) - illus.
Krishnaswamy, N., & G. N. R. Ayyangar. Cytological studies in Eleusine coracana Gaertn. Beih.
Bot. Centralbl. 57: 297-318. 1937.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 75
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Lye, K.A. Nomenclature of finger millet (Poaceae). Lidia 4(5): 149-152. 1999. –
Mann, S.K., & I. Singh. Status of finger millet in the mountain agricultural system of Himachal
Pradesh, India. Pp. 159-163 in K.W. Riley et al., eds. Mountain Agrigulture and crop
genetic resources. Oxford and Delhi. 1990.
Mehra, K. L. Natural hybridization between Eleusine coracana and E. africana in Uganda. Jour.
Indian Bot. Soc. 41: 531-539. 1962. 450 J821.
-----. Differentiation of the cultivated and wild Eleusine species. Phyton 20: 189-198. 1963.*
Mehra, K.L. Considerations on the African origin of Eleusine coracana (L.) Gaertn. Current
science (Bangalore) 32(7): 300, 301. 1963. Note Includes references. eleusine coracana.
origin. centers of origin or diversity. chromosome number. plant morphology.
Geographic africa. Geographic india
Mysore, K.S., & W.M.V. Baird. Molecular characterization of the tubulin-related gene families
in herbicide resistant and susceptible goosegrass (Eleusine indica). Weed science 43(1): 28-33.
1995. Abstract Goosegrass, wide spread throughout the tropics and subtropics, is one of the most
noxious weeds known. Recently, biotypes of goosegrass have been found resistant to the
dinitroaniline herbicides. An alteration in the structure/composition of a tubulin protein has been
postulated as an explanation for the hyperstable microtubules and the resistant phenotype. Our
study was initiated to investigate the structure of the alpha (alpha)-, beta (beta)- and gamma
(gamma)-tubulin related gene sequences in resistant, intermediately resistant, and susceptible
biotypes. Heterologous tubulin gene clones were used as probes of restriction endonucleasedigested genomic DNA from each biotype, to determine gene size and copy number and to screen
for restriction fragment length polymorphisms. The tubulin genes are organized into gene
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 76
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families. There are three to five alpha-tubulin genes, four to seven beta-tubulin genes, and four to
eight gamma-tubulin genes. There was no evidence of multiple copies or tandem repeats of any
individual gene sequence. Although RFLPs were observed, no significant difference in the
banding pattern between the resistant and the susceptible biotypes was found for either alpha-,
beta-, or gamma-tubulin gene families. Therefore, it is unlikely that the difference between the
herbicide-response phenotypes can be attributed to large deletions or insertions in a tubulin gene.
Narayanaswami, S. Microsporogenesis and male gametophyte in Eleusine coracana Gaert. Cur.
Sci. 21: 19-21. 1952.
-----. The structure and development of the caryopsis in some Indian millets V. Eleusine coracana
Gaertn. Pap. Mich. Acad. 40: 33-46. pl. 1-3. 1955.
Neves SS, Swire Clark G, Hilu KW, Baird WV, 2005., Phylogeny of Eleusine (Poaceae:
Chloridoideae) based on nuclear ITS and plastid trnT-trnF sequences. Molec. Phylogenet. Evol.
35. 395-419
Philips. -------------------. Kew Bull. 27: 251-270. 1972.
Pohl, R.W. Nardus stricta and Eleusine multiflora (Gramineae), new to Mesoamerica. Rev. Biol.
Trop. 35: 147-149. 1987.*
Portères, R. Eleusine coracana Gaertner, Céréale des humanités pauvres des pays tropicaux. Bull.
Inst. Franç. Afr. Noire 13: 1-78.*
Purseglove, J. W. Millets. Eleusine coracana, Pennisetum americanum (Gramineae), pp. 91-93.
In: N. W. Simmonds, Evolution of crop plants. xii + 339 pp. London & New York. 1976.
Ramachandran, K. R. A note on the occurrence of monostachous inflorescence in Eleusine indica,
Gaertn. Cur. Sci. 20: 189. 1951.
Rao, V. R., & K. V. R. Rao. Some observations on the panicle types in ragi (Eleusine coracana).
Andhra Agr. Jour. 9(4): 229-231. 1962.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 77
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Ravindran, G. Seed protein of millets: amino acid composition, proteinase inhibitors and in-vitro
protein digestibility. Food chemistry 44(1): 13-17. 1992. Abstract Six varieties of
common millet (Panicum miliaceum, three varieties of finger millet (Eleusine coracana)
and four varieties of foxtail millet (Setaria italica) were analyzed to determine the
nitrogen constituents, amino acid composition, proteinase inhibitors and in-vitro protein
digestibility (IVPD). The non-protein N accounted for 17.3, 12.5 and 17.0% of the total N
found in common millet, finger millet and foxtail millet, respectively. Millet proteins
were deficient in lysine, but contained adequate levels of the other essential amino acids.
The proteins in finger millets were better balanced compared to those in common millet
and foxtail millet. Little varietal differences were observed within millet types in terms of
amino acid concentrations. The anti-tryptic activities of millets were high compared to
their anti-chymotryptic activities. Foxtail millet had no detectable anti-chymotryptic
activity. The IVPD values of raw, uncooked millets were low, but were improved by
cooking.
Reddy, C.N. & M.P. Shree. Differential effects of seedcoat leachates on seed-borne pathogens
associated with fingermillet (Eleusine coracana Gaertn.). Plant physiology &
biochemistry 14 (1):103-107. 1987. Note Includes references. eleusine coracana.
varieties. drechslera. drechslera tetramera. seed coats. plant extracts. antifungal agents.
phenolic compounds. chemical analysis. disease resistance. OtherSubject drechslera
nodulosa
Rachie, K.O.,& L.V. Peters LV, The Eleusines: a review of the world literature. Hyderabad :
International Crops Research Institute for the Semiarid Tropics vii, 129p. 1977.,
Phillips, S.M. 1972. A survey of the genus Eleusine Gaertn. (Gramineae) in Africa. Kew Bull.
27: 251-270.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 78
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Salimath, S.S., A.C. De Oliveira, I.D. Godwin, & J.L. Bennetzen. Assessment of genome origins
and genetic diversity in the genus Eleusine with DNA markers. Genome 38(4): 757-763.
1995. Abstract Finger millet (Eleusine coracana), an allotetraploid cereal, is widely
cultivated in the arid and semiarid regions of the world. Three DNA marker techniques,
restriction fragment length polymorphism (RFLP), randomly amplified polymorphic
DNA (RAPD), and inter simple sequence repeat amplification (ISSR), were employed to
analyze 22 accessions belonging to 5 species of Eleusine. An 8 probe-3 enzyme RFLP
combination, 18 RAPD primers, and 6 ISSR primers, respectively, revealed 14, 10, and
26% polymorphism in 17 accessions of E. coracana from Africa and Asia. These results
indicated a very low level of DNA sequence variability in the finger millets but did allow
each line to be distinguished. The different Eleusine species could be easily identified by
DNA marker technology and the 16% intraspecific polymorphism exhibited by the two
analyzed accessions of E. floccifolia suggested a much higher level of diversity in this
species than in E. coracana. Between species, E. coracana and E. indica shared the most
markers, while E. indica and E. tristachya shared a considerable number of markers,
indicating that these three species form a close genetic assemblage within the Eleusine.
Eleusine floccifolia and E. compressa were found to be the most divergent among the
species examined. Comparison of RFLP, RAPD, and ISSR technologies, in terms of the
quantity and quality of data output, indicated that ISSRs are particularly promising for the
analysis of plant genome diversity.
Samathuvam, K. Studies on the relationship between grain yield and tiller sequence in ragi
(Eleusine coracana Gaertn.). Madras Agr. Jour. 49: 222-224. 1962.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 79
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Singh, J. S., & R. Misra. Influence of the direction of slope and reduced light intensities on the
growth of Eleusine indica. Trop. Ecol. 10: 27-33. 1969.
Sivagnanam, L. Extra-axillary digitate spike in Eleusine coracana Gaertn. Madras Agr. Jour.
47(7): 329. 1960.
Sridhar, R., & G. Lakshminarayana. Contents of total lipids and lipid classes and composition of
fatty acids in small millets: foxtail (Setaria italica), proso (Panicum miliaceum), and
finger (Eleusine coracana). Cereal chemistry. July/Aug 1994. v. 71 (4) :355-359. Note
Includes references setaria italica. panicum miliaceum. eleusine coracana. lipids. fatty
acids. phospholipids. classification. food composition. nutrient content.
Thomas, D.G. Finger millet (Eleusine coracana (L.) Gaertn.), pp. – in J.D. Jameson, ed.
Agriculture in Uganda. London. 1970.*
Toole, E.H., & V.K. Toole. Germination of seed of goosegrass, Eleusine indica. Jour. Am. Soc.
Agron. 32: 320, 321. 1940*
Ujwal, M.L., P.S. Reddy, & J.D. Cherayil. Nucleotide sequence of a nuclear tRNA(Gly) (GCC)
gene of a higher plant, ragi (Eleusine coracana). Plant physiology 106(3): 1217-1218.
1994. Note Includes references eleusine coracana. registration. genes. rna. nucleotide
sequences. OtherSubject molecular sequence data OtherSubject genbank/u02636 Other
Author
Upadhyaya, M.N., S.V. Hegde, P.V. Rai, & S.P. Wani. Root-associated fixation in finger millet.
Cereal nitrogen fixation : proceedings of the Working Group Meeting held at ICRISAT
Center, India, 9-12 October 1984. :93-101. Note Includes references. eleusine coracana.
roots. nitrogen fixation. nitrogenase. enzyme activity. varietal effects. nitrogen-fixing
bacteria.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 80
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Vaughn, K.C., M.A. Vaughan, & B.J. Gossett. A biotype of goosegrass (Eleusine indica) with an
intermediate level of dinitroaniline herbicide resistance. Weed technology 4: 157-162.
1990. ill.
Wakizuka, T. & Yamaguchi, T. Multiple floral bud formation in vitro on the giant dome of
Eleusine coracana Gaertn.
Current plant science and biotechnology in agriculture ; 22
:405-410. 1994
Werth, C.R. Isozymes of Eleusine (Gramineae) and the origin of finger millet. Am. Jour. Bot..
Sept 1994. v. 81 (9) :1186-1197. Note Includes references eleusine. genetic analysis. loci.
isoenzymes. genetic variation. ploidy. alleles. phylogeny. Other Author Hilu, K.W. Other
Author Langner, C.A. Abstract The predominantly African grass genus Eleusine comprises nine
species, including diploids and tetraploids based on n = 8, 9, and 10. Among the polyploids are
the important crop finger millet, Eleusine coracana subsp. coracana, and its putative wild
ancestor, E. coracana subsp. africana. Eleusine coracana is believed to be an allotetraploid
derived by hybridization between E. indica and an unknown diploid. To evaluate this hypothesis,
16 isozyme loci coding nine enzymes were compared among seven of the nine Eleusine species
(E. intermedia and E. semisterilis were unavailable). Genetic variability differed substantially
among diploid species, ranging from P = 0.563, A = 1.6, H = 0.208 in E. indica to P = 0.188, A =
1.2, H = 0.042 in E. jaegeri. The diploids tended to be genetically distinct, with values of Rogers'
Similanty ranging from S = 0.294 (E. jaegeri/floccifolia) to S = 0.794 (E. indica/tristachya). Both
subspecies of the tetraploid E. coracana exhibited fixed heterozygosity at several loci, verifying
their hypothesized allotetraploid status. Both tetraploids also possessed E. indica marker alleles
at all loci, corroborating ancestry by this taxon. Genotypes of the non-indica ancestor, inferred
separately for each tetraploid, differed substantially from all candidate diploids and also from
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 81
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each other. These data indicate that 1) none of the candidate diploids investigated is likely to
have been the non-indica ancestor of E. coracana, and 2) the non-indica ancestor of the wild
tetraploid may differ from that of the crop. The latter conclusion is inconsistent with the
complete chromosomal homology exhibited between the two tetraploid subspecies, indicating the
need for additional evidence bearing on their relationships.
Zeng, L. Identification of a molecular marker linked to dinitroaniline herbicide resistance in
goosegrass. Proceedings, Southern Weed Science Society. Southern Weed Science Society
(U.S.) 1996. v. 49 :141-145. Note Meeting held January 15-17, 1996, Charlotte, North Carolina.
Note Includes references eleusine indica. herbicide resistance. herbicide resistant weeds. Other
Author Baird, W.V.
11. Dactyloctenium Willd., Enum. Hort. Berol. 1029. 1809.
Annuals [perennials or of indefinite duration] of disturbed open places, typically in
somewhat moist soils. Rhizomes absent; short stolons sometimes present. Stems spreading to
erect, unbranched; leaves cauline and basal, few; sheaths shorter than the internodes, compressed,
sharply keeled, glabrous, ligule membranous, truncate, often ciliate margined, the hairs about as
long as the membrane; blade linear, flat or somewhat plicate, hispid with papillose based hairs.
Inflorescence of 2-several secund racemose branches, these branches ± digitately arranged at the top
of the stem; rachis flattened, the tip projecting a few mm beyond the last spikelet. Spikelets
strongly laterally compressed, orbiculate-ovate, 2-3 flowered, arranged in two rows on opposite
sides of the rachis, sessile; glumes 2, subequal, ovate to ovate-lanceolate, about half as long as the
first lemma, 1-nerved, keeled, glabrous, the first mucronulate, the second awned from the apex, the
awn about as long as the glume, scabrous, incurved; rachilla articulated between or above the
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 82
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glumes; flowers bisexual; lemmas ovate, 3 nerved (the midvein conspicuous, the lateral nerves
inconspicuous), mucronate; paleas elliptic, as long as the lemmas, acute, strongly 2-nerved, the
nerves minutely scabrous or ciliate. Lodicules 2, cuneate, truncate, fleshy. Stamens 3; anthers
ellipsoid. Ovaries ovoid to ellipsoid, glabrous; styles 2, separate, short, terminal stigmas plumose.
Fruit an achene (pericarp free and easily detached), ovoid-orbiculate, terete, transversely ridged and
conspicuously rugose; hilum linear, nearly as long as grain; embryo large, typically 1/3 –3/4 the
length of the grain. Base chromosome number 9. Type species: D. aegyptium Willd., the only
species treated fully in the text of the protologue (D. mucronatum and D. prostratum were included
in a footnote). (Named from the Greek, dactylon, finger, and ctenium, comb, from the digitate
arrangement of the racemes, and the awned spikelets.). – Crowfootgrass.
A small genus of about 10 species, native to warm regions of the Old World. . Dactyloctenium is
closely related to Eleusine, sharing free pericarp and general aspect, including digitate clustering of
racemose branches.
D. aegyptium 2n=20, 36, 40, 45, 46, 48, is a common weed, established throughout the world. In
our area known from Arkansas (Clark Co.; Smith); statewide in Louisiana (MacRoberts)
Note check spelling of D. aegypticum vs. aegyptium
References
Under subfamily see; Jordan et al.
Fischer & Schweickerdt. --------------- Ann. Natal Museum 10: 47-77. 1941. at MCZ ??
Garg, S.P., R. Bhushan, R.C. Kapoor, & B.K. Dutta. Free amino acids and sugars in
Dactyloctenium species growing on different soil types in arid regions.. Transactions of
Indian Society of Desert Technology and University Centre of Desert Studies 5(1): 125128. 1980. [D. aegyptium, D. scindicum.]
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 83
06/27/17
Hatch, S.
Koteswari, M.V.&, T.N. Mary. Reproductive effort in crowfoot grass, Dactyloctenum aegyptium
(L.)Beauv. Indian bot. reporter 5(1): 10-13. 1986. [Ecotypes; genotypes; chromosome
number].
Pringle, J.S. Dactyloctenium aegyptium (Gramineae) new to Belize. Sida 11 (2): 245.1985.
Mary, T.N. & M.V. Koteswari. Some ecological and anatomical studies of the crowfoot grass,
Dactyloctenium aegyptium (L.) Beauv. Geobios New Reports. 3(1): 17-20. 1984. [Ill.;
seed production; seed germination; reproductive ability.]*
Sharma, B.M, & Chivinge, A.O. Contribution to the ecology of Dactyloctenium aegyptium (L.)
P. Beauv., a nutritious fodder, Nigeria. Jour. range management.. 35: 326-331. 1982
[ill.]
Sharma, M.L., et al. Leaf anatomy of north-west Indian species of Dactyloctenium Willd.
(Gramineae) and its allies. Jour. Pl. Sci. Res. 3: 83-88. 1987.*
Sachdeva, S.K. & R. Kals. Cytologic, morphologic and chemotaxonomic studies in
Dactyloctenium aegyptium (L.) Beauv. complex. Proceedings. Plant sciences - Indian
Academy of Sciences 90(3): 217-225. 1981.*
Sharma, M.L., R.K. Bhanwra, & S. Kaur. Seed sterility in Dactyloctenium sindicum Boiss.
(Poaceae). Current Science (Bangalore) 50: 771-772. 1981.*
12. Sporobolus R. Brown, Prodr. 169. 1810.8
Small to large perennials or annuals of grasslands, open woods, and disturbed open places.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 84
06/27/17
Rhizomes present or not; stolons present or not. Stems erect, unbranched; leaves cauline, few;
sheaths longer or shorter than the internodes, rounded, glabrous, or puberulent near the collar,
ligule a fringe of hairs; blade linear, flat, involute, or terete [solid, cylindrical in S. rigens].
Inflorescence a narrow to ovoid panicle, with ascending or spreading branches (s9ometimes partly
included in the uppermost sheath). Spikelets scarcely laterally compressed, 1 flowered, shortpedicellate in clusters at the branch tips; disarticulation above glumes; callus glabrous; glumes 2,
unequal, ovate to ovate-lanceolate, firm, glabrous, the first much shorter than the first lemma,
nerveless or 1-nerved, the second nearly as long as the first lemma, 1-nerved; rachilla not prolonged
[except in S. subtilis]; lemmas ovate-lanceolate, acute, 1-nerved [3-nerved in S. palmeri],
sometimes keeled, hyaline, or chartaceous, glabrous, pubescent, or pilose; paleas elliptic, a little
longer than the lemmas, the apex acute, awnless, 2-nerved, glabrous, not enclosing the mature fruit.
Lodicules 0 or 2, cuneate, truncate, fleshy. Stamens 2 or 3; anthers ellipsoid. Ovaries ovoid to
ellipsoid, glabrous; styles 2, separate, short, terminal stigmas plumose. Fruit an achene (pericarp
free), terete , oblong, ellipsoid, or ovoid; hilum linear, nearly as long as grain; embryo large,
typically1/3 –3/4 the length of the grain. Base chromosome number 9. (synonyms). Type species:
S. indicus (L.) R.Br.; see Nash in Britton & Brown, Illus. Fl. No. U.S. ed. 2, 1: 194. 1913) (Name
from the Greek spora, seed, and ballein, to throw, referring to the seed that falls free from the
lemmas.). – Dropseed, Rushgrass, Sacaton.
A genus of about 100 species, worldwide in warm-temperate and tropical regions, but most
diverse in the New world. About 40 species occur in the U.S., and about 12 species in the
Southeast. Gould recognized 23 species in Texas, many of which also occur in the southeast. In
8
Brown included the following species: S. indicus (Agrostis indica); S. eleongatus; S. pulchellus.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 85
06/27/17
general, the one-flowered spikelets distinguish this genus from other similar ones in the subfamily,
the pericarp is mucilaginous when moistened. There is general agreement that Sporobolus is
closely related to Muhlenbergia. Several species have been moved from one genus to the other. The
distinction between Sporobolus and Eragrostis is also problematic. Sporobolus is generally
distinguished by its one-flowered, spikelets and 1-nerved lemmas from eragrostis. Sporobolus
asper and its close relatives show some similarity to Calamovilfa in their chartaceous, keeled
lemmas, but lack the distinctive bearded callus of Calamovilfa. Molecular approaches may be
valuable in resolving the relationships and origins of these genera.
S. airoides (Torr.) Torr., alkali sacaton, finetop saltgrass, 2n = ca. 80, ca. 90, ca. 108, ca. 126, is a
western and southwestern species similar in appearance to S. heterolepis. It occurs from South
Dakota to British Columbia (Darbyshire), south to Arkansas (Pulaski co.; Smith), Texas, northern
Mexico and southern California. In much of its range it occurs in alkaline or saline places. It has
recently reported as naturalized in India.
S. asper (Michx.) Kunth, harsh dropseed, 2n = 54, ca. 88, is a wide ranging species. Three varieties
have been recognized. The typical variety, var. asper, has narrow inflorescences and lacks scaly
rhizomes occurs from Vermont to eastern Washington, south to Alabama (not florida ??), and
Texas. Var. macer, has elongate scaly rhizomes and narrow inflorescences, and occurs over a much
narrower range, Mississippi, Louisiana, Arkansas, and east Texas. Var. drummondii (Trin.) Vasey,
also lacks rhizomes, and has open inflorescences. It occurs from Alabama and Missouri to Texas.
S. clandestinus (Biehler) Hitchc., dropseed (chromosome number not reported), ranges from New
england to Wisconsin, south to florida and Texas. It is a common species of dry fields, roadsides,
and edges of woods throughout the southeast. . It is a perennial with narrrow, terminal and axillary,
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 86
06/27/17
partly enclosed inflorescences.
S. cryptandrus (Torr.) a.Gray, Sand Dropseed, 2n = 18, 36, 38, occurs from Maine to Ontario and
Washington State, south to Arkansas, Texas, and and Mexico. It is apparently lacking in the
southeast, being reported only from Arkansas, where it is found throughout the state (Smith). It is a
grass with narrow inflorescence, typically found in well drained soils of roadsides and railroads
Infraspecific variation see Jones & Fassett.
S. heterolepis (A.Gray) A.Gray, northern dropseed, prairie dropseed, 2n = 72, is a wide ranging
species found from Quebec to Saskatchewan, south to Texas and Colorado. It barely enters our area
in northwestern Arkansas only (Smith).
S. neglectus Nash, poverty-grass, puffsheath dropseed, 2n=36, is a wide-ranging species primarily
to our north. It ranges from New England to Montana and Washington, south to Virginia,
Tennessee, Arkansas (Smith), and Arizona. It has a narrow inflorescence partly enclosed in the
uppermost leaf.
S. ozarkanus Fern., Ozark dropseed, (chromosome number apparently unknown), is endemic to
Missouri, Arkansas, and north central Texas. It has inflorescences that are narrow and partly
enclosed with in the uppermost leaf.
S. indicus (L.) R.Br. [S. poiretii (Roem. & Schult.) Hitch.], smutgrass, blackseed, 2n=18, 24, 36,
occurs in all the southeastern states (???), and ranges from Virginia, Tennesse, and Oklahoma south
to Argentina. It is believed to be native to the tropical regions of the Old world. The species is often
infected with the smut fungus _____, hence the common name.
S. pyramidatus (Lam.) Hitchc., Kansas to Colorado, south to northwestern Arkansas (Franklin Co.,
Smith), Louisiana, Texas, and Arizona. It has been found as an adventive further east in several
statees. It is a tufted perennial, with an open inflorescence and conspicuous verticillate branches.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 87
06/27/17
S. vaginiflorus (Torr. ex A.Gray) A. Wood, is a wide-ranging species, from New England to ___
west to Minnesota, Nebraska, and Arizona. It is typically found in droughty sites, thin soils over
rock outcrops, parking areas, railroads. It occurs in Arkansas only in the northwestern part of the
state (Smith)
S. junceus (Michx.) Kunth, Pineywoods Dropseed, occurs from Virginia to florida west to east
Texas. A chromosome report for this species would be desirable.
S. virginicus (L.) Kunth, seashore dropseed, 2n=20, 30
Several species have ethnobotanical significance. Sporobolus pyramidatus, for example, is
medicinally versatile (Morton). A decoction is taken for urinary irritation, as a diuretic, and for
severe fevers. A hot infusion is taken for nasal and chest congestion in Curaçao, and it is drunk as a
laxative in Aruba. Sporobolus virginicus is used for urinary and kidney problems in Venezuela
(Morton). The seeds of S. cryptandrus and S. flexuosus (Thurber) Rydb. have been used as a food
by some of the indigenous tribes of the American Southwest (Holmgren & Holmgren).
References
Under subfamily references see: Austin; Eastman et al.; Morton; Rasmussen & Rice.
Abrams, M.D. Effects of burning regime on buried seed banks and canopy coverage in a Kansas
tallgrass prairie, U.S.A. Southwest. Nat. 33: 65-70. 1988.
Astegiano, M.E. Cleistogamy and chasmogamy in Sporobolus indicus (Poaceae). Kurtziana 18:
69-76. 1986.
Baskin, J. M., & C. Caudle. Germination and dormancy in cedar glade plants. I. Aristida
longespica and Sporobolus vaginiflorus. Tenn. Acad. Sci. Jour. 42(4): 132, 133.. 1967.
[Cold treatment breaks physiological dormancy; best germination obtained at 20°C.].
-----, -----, & E. Quarterman. Germination and dormancy in two cedar glade grasses. (Abstr.) ASB
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 88
06/27/17
Bull. 14: 22. 1967. [Aristida longispica Poir. and Sporobolus vaginiflorus (Torr.) Wood.]
Batanouny, K.H., A.H. Hassan, & K.M. Zayed. Proline accumulation in plants of different
ecological groups as a response to water deficit. Qatar Univ. Sci. Bull. 5: 131-144. 1985.*
[In Sporobolus virginicus, proline content increased during the day and decreased at night,
just as in Zea mays. Seasonal changes corresponding to relative humidity were also noted.]
Beal, W. J. Expulsion of the seeds of Sporobolus cryptandrus. Bot. Gaz. 11: 247. year.
Bir, S.S. , M. Sahni, & C.P. Singh. Cytology of genus Sporobolus R.Br. from North India (Punjab
Plain). Cytologia (Tokyo) 53: 53-57. 1988.
Böcher, T.W. Leaf anatomy in Sporobolus rigens (Tr.) Desv. Bot. Not. 125: 344-360. 1972.
Boe, A. Variability for seed size and yield in two tall dropseed populations. Jour. Range
Managem. 43: 195-197. 1990.
Boechat, S.C. Quatro espécies novas de Sporobolus R. Br. (Poaceae, Chloridoideae) do Brasil.
Iheringia, Sér. Bot. 51(2): 33-44. 1994.*
Bor, N. L. Sporobolus capillaris Miq. Kew Bull. 1949: 224. 451.
Boyle, R. V. Sacaton [Sporobolus wrighti] its value to ranchers. Ariz. Stockman 22(12) 20, 85.
1956. *.
Brecke, B.J. Smutgrass (Sporobolus poiretii) control in bahiagrass (Paspalum notatum) pastures.
Weed Sci. 29: 553-555. 1981.
Canales, M.J., & J.F. Silva. Effect of fire on growth and demography of Sporobolus cubensis.
Acta Oecol. Oecol. Gen. 8(3): 391-402. 1987.*
Cavalheiro, E.M., & H.M. Longhi-Wagner. As epecies do gênero Sporobolus R.Br. (Gramineae,
Chloridoideae) no Rio Grande do Sul, Brasil. (in Portuguese, English summary.) Iheringia,
Bot. 41: 101-139. 1991. [7 spp., keys, descriptions.]
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 89
06/27/17
Clayton, W. D. Studies in Gramineae. XXIII. Sporoboleae. Kew Bull. 25(2): 247-251. 1971.
Colbry, V. L. Diagnostic characteristics of the fruits and florets of economic species of North
America Sporobolus. Contrib. U. S. Natl. Herb. 34(1), 24 p. 1957.*
Cormack, R. G. H. The effect of environmental factors on the development of root hairs in Phleum
pratense and Sporobolus cryptandrus. Am. Jour. Bot. 31: 443-449. f. 1-11. 1944.
Danckwerts, J.E. Growth and desiccation of Themeda triandra and sporobolus fimbriatus in
relation to diminishing moisture availability. Jour. Grassl. Soc. So. Africa 5(2): 96-101.
1988.
Darbyshire, S.J. Sporobolus airoides (Eragrosteae: Poaceae) in British Columbia. Syesis 17: 11,
12. 1984 [1985].*
Donovan, L. Intraspecific genetic variability in Sporobolus virginicus in response to anaerobic
substrates. (Abstr.) Bot. Soc. Am. Misc. Publ. 162: 68.1982.
-----, & J.L. Gallagher. Anaerobic substrate tolerance in Sporobolus virginicus (L.) Kunth. Amer.
Jour. Bot. 71: 1424-1431. 1984.
Doyon, D., & W. G. Dore. Notes on the distribution of two grasses, Sporobolus neglectus and
Leersia virginica, in Quebec. Can. Field-Natur. 81(1): 30-32, map. 1967.
Eastman, P.A.K., N.G. Dengler, & C.A. Peterson. Suberized bundle sheaths in grasses (Poaceae)
of different photosynthetic types. I. Anatomy, ultrastructure, and histochemistry.
Protoplasma 142(2-3): 92-11. 1988. [Incl. Sporobolus sp.]*
Ebbers, B.C., R.C. Anderson, & A.E. Liberta. Aspects of mycorrhizal ecology of prairie dropseed,
Sporobolus heterolepis (Poaceae). Amer. Jour. Bot. 74: 564-573. 1987.
Ehara, K., H. Ikeda, & A. Fujii. Studies on ecological & growth characteristics of warm-season
native grasses. 5. On salt tolerance of Sporobolus virginicus Kuntze. (Japanese; Eng.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 90
06/27/17
Sum.). Kyushu Univ. Fac. Agr. Sci. Bull. 26(1/4): 441-444. 1972.
Fernald, M. L. Two segregates in Sporobolus. Rhodora 35: 108-110. 1933.
Freeman, O. M. Notes on flora of Polk Co., N.C. Castanea 20: 37-57. 1955. [S. clandestinus, S.
poireti.]
Frith, J. L. The germination of Sporobolus virginicus. Austral. Inst. Agr. Sci. J. 23(1): 69-75. Mar.
1957.
Hajduk, J. Plants in fissures of and spaces between rocks and concrete as an ecological
phenomenon. Biologia (Bratislava) 433 : 811-820. 1988.*
Hall, D.W. Is it wiregrass? Nat. Areas Jour. 9: 219-222. 1989.*
Harper, R. M. Some new or otherwise noteworthy plants from the coastal plain of Georgia. Bull.
Torrey Club 33: 229-245. fig. 1, 2. 1906. [New species described in Sporobolus.]
Jacobs, S.W.L. Taxonomy of Paramatta Grass and related species. Pp. 11-23 in R. Dyason, ed.,
Paramatta Grass and its control. New South Wales. 1985.*
Johnson, R.G., & R.C. Anderson. The seed bank of a tallgrass prairie in Illinois. Amer. Midland
nat. 115: 123-130. 1986. [S. heterolepis a dominant, relative cover 13.8 %, but only 5% of
germinated seeds.]
Jones, E. K. & N. C. Fassett. Subspecific variation in Sporobolus cryptandrus. Rhodora 52: 125,
126. 1950.
Judziewicz, E.J., & P.M. Peterson. Sporobolus temomairensis, new species (Poaceae,
Eragrostideae) from northern South America. Syst. Bot. 14: 525-528. 1989.
Mahmood, K., K.A. Malik, K.H. Sheikh, & M.A.K. Lodhi. Allelopathy in saline agricultural land:
vegetation successional changes and patch dynamics. Jour. Chem. Ecol. 15: 565-580.
1989.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 91
06/27/17
McCormick, J., & M. F. Buell. Sporobolus cryptandrus (Torr.) A. Gray in the New Jersey pine
barrens. Torrey. Bot. Club. Bull. 83(6): 439.. 1956.
McGregor, R.L. Seed dormancy and germination in the annual cleistogamous species of
Sporobolus (Poaceae). Trans. Kansas Acad. Sci. 93(1-2): 8-11. 1990.
Moore, D. M. Additional new records for the Arkansas flora, II. Proc. Ark. Acad. Sci. 5: 91-93.
1952. [S. pulvinatus Swallen].
Nugent, G., & D.F. Gaff. Electrofusion of protoplasts from desiccation tolerant and desiccation
sensitive species of grasses. Biochem. Physiol Pflanz. 185: (1-2): 93-97. 1989.*
Potvin, M.A. Seed rain on a Nebraska, U.S.A., sandhills prairie. Prairie Nat. 20: 810-90. 1988.
Quinn, J. A. Ecotypic variation in switch grass (Panicum virgatum) and sand dropseed (Sporobolus
cryptandrus). Diss. Abstr., Sect. B, 27(12): 4261B-4262B. 1967.
Riggins, R. A biosystematic study of the Sporobolus asper complex (Gramineae). Iowa State jour.
Res. 51(3): 287-321. 1977.*
Rolfsmeier, S.B., R.B. Kaul, M.M. Garabrandt, & D.M. Sutherland. New and corrected floristic
records for Nebraska, U.S.A. Trans. Nebr. Acad. Sci. 16: 115-122. 1988.*
Roy, G.P. &, Singh, V. Vicia monantha Retz. and Sporobolus airoides (Torr.) Torr., new to
Indian flora. Jour. the Bombay Natural History Society 77 (3): 532-534. 1980.
Scribner, F. L. Notes on Sporobolus. Bot. Gaz. 21: 14-16. 1896.
Sheikh, K.H., & K. Mahmood. Some studies on field distribution and seed germination of Suaeda
fruticosa and Sporobolus arabicus with reference to salinity and sodicity of the medium.
Pl. & Soil 94: 333-340. 1986.
Sirrine, E. & E. Pammel. Some anatomical Studies of the Leaves of Sporobolus and Panicum.
Proc. Iowa Acad. Sci. 3: 148-159. pl. 6. 1896.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 92
06/27/17
Smith-White, A.R. Sporobolus virginicus (L.) Kunth in coastal Australia: the reproductive
behaviour and distribution of morphological types and chromosome races. Austral. Jour.
Bot. 36: 23-39. 1988.
-----, & P. Adam. An unusual form of the salt marsh grass Sporobolus virginicus (L.) Kunth. West.
Austral. Nat. 17: 118-120. 1988.
Straub, P.F., & J.L. Gallagher. Cold tolerance in Sporobolus virginicus (L.) Kunth, robust form,
tissue cultures and whole plants. Pl. Cell Environm. 12: 503-510. 1989.
Sundell, E. Two additions to the Arkansas flora from Warren Prairie (Aster pratensis, Sporobolus
junceus). Sida 10: 188, 189. 1983.
Swallen, J. R. New United States grasses. Jour. Wash. Acad. Sci. 31: 348-355. f. 1-8. 1941.
[Sporobolus silveanus Swallen, sp. nov., n.e. of Orange, Tex.]
Thieret, J. W. Twenty-five species of vascular plants new to Louisiana. Proc. Louisiana Acad. Sci.
32: 78-82. 1969. [S. heterolepis Gray.]
Toole, V.K. Factors affecting the germination of various dropseed grasses (Sporobolus spp.) Jour.
Agr. Res. 62: 691-715. 1941.
Veldkamp, J.F. The true identity of Sporobolus poiretii (Gramineae). Taxon 39: 327, 328. 1990.
Winter, K., M.R. Schmitt, & G.E. Edwards. Microstegium vimineum, a shade adapted C4 (carbon
pathway) grass. Pl. Sci. Lett. 24: 311-318. 1982. [Comparison of growth with Sporobolus
airoides.]
Wipff, J.K., & S.D. Jones. Nomenclatural combination in Poaceae. Phytologia 78: 244, 245.
1995. [S. compositus var. clandestinus.]*
Wood, J.N., & D.F. Gaff. Salinity studies with drought-resistant species of Sporobolus. Oecologia
78: 559-564. 1989.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 93
06/27/17
Zacharias, P.J.K., N.M. Tainton, & C. Oberholster. The effect of fir e on germination of five
common veld grasses. Jour. Grassl. Soc. South Afr. 5(4): 229-230. 1988.
16?????. Crypsis Aiton, Hort. Kew. 1: 48. 1789, nom. cons
Type species: C. aculeata (L.) Aiton, the only species included by Aiton..name from Greek krupsis,
concealment, because the inflorescence is partly hidden by the subtending leaf).
C. schoenoides known in U.S. from Illinois, Indiana, New York
Are we sure any occur in the southeast? Hamel cites none from SE U.S.
References:
Hammel, B. Syst. Bot. 1979.
13. Calamovilfa (Gray) Scribner in Hackel, True Grasses 113. 1890.
Large perennials of disturbed open places, typically in sandy well drained soils. Rhizomes
present or not; horizontal, with conspicuous lance-ovate scales. Stems erect, unbranched; leaves
cauline, few; sheaths rounded, glabrous, or puberulent near the collar, ligule a fringe of hairs; blade
linear, flat near the base, becoming involute distally, tapering to a long narrow involute apex.
Inflorescence a panicle, with ascending or spreading branches. Spikelets weakly laterally
compressed, 1 flowered, short-pedicellate in clusters at the branch tips; disarticulation above
glumes; callus abundantly pubescent with long silky hairs; glumes 2, subequal, lanceolate, firm,
glabrous, 1-nerved; rachilla not prolonged; lemmas ovate-lanceolate, acute, 1-nerved, pubescent on
the lower half abaxially; paleas narrowly elliptic, a little longer than the lemmas, the apex acute,
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 94
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awnless, 2-nerved, the nerves ciliate. Lodicules 2, cuneate, truncate, and distinctly assymetrical,
fleshy. Stamens 3; anthers ellipsoid. Ovaries ovoid to ellipsoid, glabrous; styles 2, separate, short,
terminal stigmas plumose. Fruit an achene (pericarp free), subterete to cylindric, oblong, to broadly
ellipsoid; hilum linear, nearly as long as grain; embryo large, typically1/3 –3/4 the length of the
grain. Base chromosome number 10. (Calamagrostis sect. Calamovilfa Gray, Man. Bot. 582.
1848; Scribner refered to "Calamovilfa (Gray as a sect. of Calamagrostis)" without reference to
edition or page number). Type species (Named--------------.). – Sandreed.
A small genus of only five species, endemic to North America. four of the five species
occur in the southeast. C. curtissii (Vase) Scribn., pine barrens of northern Florida, a tufted
perennial with slender, virgate inflorescence (see Johnson & blyth). C. brevipilis (Torr.) Scribn.,
coastal wetlands from New Jersey to South Carolina, tufted perennial with open inflorescence. C.
longifolia (Hooker) Scribn., long creeping rhizomes, single stems, glabrous lemmas, Great Lakes
region from western New York to Alberta, Idaho, and Colorado
Calamovilfa gigantea (Nutt.) Scribn. & Merr. [USDA Div. Agrost. Circ. 35: 2. 1901], big
sandreed, 2n=60, sand dunes from Kansas to Utah, Texas, and Arizona, long creeping rhizomes,
lemma villous. It is closely related to the preceding, but is more robust, and has pubescent lemmas
and palea. It is considered to be a valuable sand binder in dune areas Holmgren & Holmgren).
Calamovilfa arcuata Rogers Oklahoma: McCurtain Co., 15 Aug 1984, Taylor & Taylor 32540
(MO). Also specimens from Pushmataha and Atoka cos.
Calamovilfa resembles Calamagrostis and Ammophila of the Pooideae in general habit. The
affinities of Calamovilfa with the Chloridoideae were recognized by Reeder & Ellington, noting
similarities in embryo, lodicules, leaf anatomy, microhairs, and chromosome number. Calamovilfa
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 95
06/27/17
is apparently closely related to Sporobolus, with which it shares several features, including 1-nerved
lemmas (scarce in this subfamily) and fruits with the pericarp free from the seed coats (Clayton &
Renvoize)
The seeds of C. longifolia are able to germinate under 10-12 cm of sand (Maun & Lapierre)
References
Under subfamily references see Clayton & Renvoize, Gould; Hitchcock, Hitchcock &
Chase, Holmgren & Holmgren; Tyndall et al.
Johnson, A.E., && A. Blyth. Re-discovery of Calamovilfa curtissii (Gramineae) in the Florida
Panhandle. Sida 13: 137-140. 1988.
Maun, M.A., & J. Lapierre. Effects of burial by sand on seed germination and seedling emergence
of four dune species. Am. Jour. Bot. 73: 450-455. 1986.*
Reeder & Ellington. Brittonia 12: 71-77. 1960.*
Thieret. Castanea 31: 145-152. 1966.*
13. Muhlenbergia Schreber, Gen. Pl. ed. 8, 1: 74. 1789.
Perennials (or annuals) of disturbed open places, woods, and wetlands. Rhizomes often
present, horizontal, scaly. Stems erect (or decumbent and mat-forming), simple or branched,
glabrous or shortly pubescent at or below the nodes; leaves cauline and basal, few; sheaths rounded
or keeled, glabrous [scabrous], ligule a membrane with a fringe of short white hairs; blade linear to
linear-lanceolate, flat, plicate, or involute. Inflorescence a panicle, terminal, or both terminal and
lateral, sometimes partly included within the subtending sheaths, narrow and spike-like, or open, in
length ranging from less than a tenth to nearly as long as the entire plant. Spikelets scarcely
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 96
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laterally compressed, 1 (-2-3) flowered, sessile or short-pedicellate; glumes 2, subequal or unequal
(the first often rudimentary in M. schreberi), generally shorter than the lemma, lanceolate to elliptic
(ovate in M. schreberi), acute to aristate [bifid], glabrous [pubescent]; rachilla short, persistent;
articulated above the glumes; lemmas oblong-ovate, 3 (rarely 5) nerved, acute to acuminate,
sometimes with an awn 2-4 [-10] times as long as the lemma body, variously glabrous or
pubescent; paleas elliptic or ovate, shorter than the lemmas, the apex acute, 2-nerved, variously
glabrous or pubescent on or between the nerves. Lodicules 2, cuneate, truncate or rounded, fleshy.
Stamens 3 [sometimes 2 in some South American species]; anthers ellipsoid to linear. Ovaries
ellipsoid, glabrous; styles 2, separate, short, terminal stigmas plumose. Fruit a caryopsis, ellipsoid,
subterete (dorsiventrally compressed) to cylindrical, generally remaining clasped by lemma and
palea; hilum linear, nearly as long as grain; embryo about ½ as long as grain. Base chromosome
number 10. (synonym??). Type species: M. schreberi, designated by --------. (Named for Gotthilf
Henry Ernest Muhlenberg, 1753-1815, Lutheran minister of Pennsylvania, early American botanist
and first American agrostologist). – Muhly.
A genus of about 125 species. Most of these occur in the New World, but a few are found in Africa,
eastern Asia, and the Himalayas.. Mexico and the southwestern U.S. form the center of diversity.
About 70 species occur in the U.S. (Hitchcok & Chase), with about 14 in the southeast, making it
the second largest chloridoid genus in our area, exceeded only by Eragrostis. About 35 species
occur in Texas, but most of these are southwestern and Mexican species not extending in to the
southeast. About 45 species occur in Arizona, 16 species in the Great Basin Region, and only 9 in
the Pacific northwest. Some 115 species occur in Mexico, with about 50 percent endemic (Peterson
et al.)
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 97
06/27/17
Divisions of the genus
List of Species
M. bushii Pohl (M. brachyphylla Bush), nodding muhly, 2n=40 has a two-parted range, from
Maryland to North Carolina, and from Indiana to Wisconsin to Nebraska south to northwestern
Arkansas (Smith) and Texas.
M. capillaris (Lam.) Trin., hairyawn muhly, hairgrass, occurs in rocky or sandy woods from New
Hampshire, Indiana and Kansas, south to Florida and eastern Texas; disjunct in the West Indies and
eastern Mexico. the chromosome number is apparently not reported.
M. cuspidata (Torr.) Rydb., Plains Muhly, occurs from Ohio to Montana south to Ky. and New
Mexico; it is known in our area from northwestern Arkansas (Smith, but not known to Chase).
M. torreyana (Schult.) Hitchc., Torrey’s Muhly, occurs interruptedly from New Jersey to Kentucky,
Georgia, and Tennessee. A plant of pine barrens and meadows, it has an open inflorescence, scaly
rhizomes and awnless lemmas and glumes. Like several similar species, including the northeastern
M. uniflora (Muhl.) Fern. and the western M. asperifolia (Nees & Meyen) Parodi, it has also been
placed in Sporobolus.
M. expansa (DC.) Trin., Virginia to Texas, including all the Southeastern states except Tennessee
and Arkansas, western Louisiana (MacR). It occurs primarily in moist coastal pine barrens. the old
leaf sheaths from a fibrous mass around the base of the stems. the chromosome number is
apparently unknown.
M. filipes M.A. Curtis, occurs in sand dunes and open coastal woodlands from North Carolina to
Florida, west along the coastal plain to southeastern Texas. It has been treated as a synonym of M.
expansa (Correll & Johnston) and of M. capillaris (Pinson & Batson).
M. frondosa (Poir.) Fern., Wirestem Muhly, 2n= 40 (42), occurs from New Brunswick to North
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 98
06/27/17
Dakota, south to northern Georgia, Louisiana (McKenzie & Urbatsch), and Texas (Gould). A
somewhat weedy species, it occurs in riverine thickets, disturbed soils, and makes trouble of itself
in flower and vegetable gardens.
M. glabriflora Scribn., Inland Muhly, 2n=40, occurs from Maryland to Illinois, south to North
Carolina, western Arkansas (Smith), western Louisiana, and Texas. It is a species of moist
woodlands.
M. glomerata (Willd.) Trin., Marsh “Timothy,” spike Muhly, 2n=20, 40, is a transcontinental,
wetland species from newfoundland to British Columbia, occurs south tin the mountains to North
Carolina. It has scaley rhizomes and densely cylindric panicles with a bristly appearance from the
awned glumesThis species has been included in M. racemosa (Michx.) BSP., 2n=40, a tetraploid of
the Rocky Mountain and Great Lakes region with one genome derived from M. glomerata, the
other from an unknown species (Pohl & Mitchell). the two species are distinct in several features
(Holmgren & Holmgren).
M. mexicana (L.) Trin., Satin-Grass, 2n=40, ranges from Maine to Washington, south to North
Carolina, Arkansas (only four counties; Smith), and Arizona (Hitchcock & Chase). 9 A species of
partly shaded damp soils.
M. schreberi Gmel., nimble-will, 2n=40 (42), New Hampshire to Nebraska, south to northern
Florida and Texas, eastern Mexico, and Arizona. Nimble will, a weedy little grass that does well in
part shade, of damp lawns, gardens, and woodland paths can be distinguished from our other
muhlies by its reduced glumes (the first is obsolete, the second only 0.1-0.2 mm long) and lack of
rhizomes. The somewhat decumbent lower stems may creep and root at the nodes, but no true
9
The epithet mexicana is a misnomer; the species does not extend into Mexico. Evidently Linnaeus was
misinformed by Jacquin about the source of the seed from which he grew plants in Uppsala on which he
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 99
06/27/17
rhizomes are produced.
M. sobolifera (Muhl.) Trin., New England to Nebraska, south to Virginia, Tennessee, northern
Arkansas (Smith), and Texas. It occurs in well drained woods. Hitchcock & Chase recognized var.
setigera Scribner from Texas and Arkansas, differing in having awned lemmas.
M. sylvatica (Torr.) Torr. ex A.Gray, forest muhly, 2n=40, occurs from Maine to South Dakota
south to North Carolina, Tennessee, Mississippi, northern Arkansas (Smith), Texas, and Arizona
(Gould). It occurs in woods, generally in mesic soils.
M. tenuiflora (Willd.) BSP, occurs from New England to Iowa, south to North Carolina, Georgia
(Duncan), Tennessee, northwestern Arkansas (Smith), and Oklahoma. Like M. sylvatica, with
which it is easy to confuse (believe me! or cf Dore & McNeill), it occurs in well drained woods.
References
Under subfamily see: Gould (Muhlenbergia treatment by C.G. Reeder); Laegaard & Sanchez Vega..
Allred, K.W., & J.T. Columbus. Additions to the flora of New Mexico, U.S.A. Phytologia 67:
361-365. 1989.*
Andariese, S.W., & W.W. Covington. Biomass estimation for four common grass species in
norhtern Arizona, U.S.A, Ponderosa pine (Pinus ponderosa). Jour. Range Managem. 39:
472-473. 1986.*
----- & -----. Changes in understory production for three prescribed burns of different ages in
Ponderosa pine (Pinus ponderosa). For. Ecol. Managem. 14: 193-204. 1986.*
Baskin, J.M., & C.C. Baskin. Dormancy breaking and germination requirements for nimble will
based the name Agrostis mexicana L. (Mantissa Pl. 1: 31. 1767).
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 100
06/27/17
(Muhlenbergia schreberi Gmel.) seeds. Jour. Range Managem. 38: 513-515. 1985.*
Beal, W. J. Rootstocks of Leersia and Muhlenbergia. Am. Nat. 22: 351, 352. pl. IV.
Bowyer, R.T., & V.C. Bleich. Effects of cattle grazing on selected habitats of southern mule deer
(Odocoileus hemionus fuliginatus) Calif. Fish Game 70: 240-247. 1984. [Cattle grazing
may limit deer numbers by reducing dense patches of M. rigens used for cover during
fawning period..]
Bryant, F.C., & R.D. Farfan. Dry season forage selection by alpaca (Lama pacos) in southern Peru.
Jour. Range Managem. 37: 330-333. 1984.*
Bush, B. F. The Missouri Muhlenbergias. Am. Mid. Nat. 6: 17-20. Mr 1919.
Caprio, A.C., & D.L. Taylor. Effect of frost on a subtropical Muhlenbergia prairie in south Florida.
Fla. Sci. 47: 27-32. 1984.*
Daniel, W. H., & J. W. Herron. Use of Zytron for controlling nimblewill [Muhlenbergia schreberi]
in bluegrass turf. Down Earth 17(4): 21-23. 1962.
Dorr, L.J., & P.M. Peterson. Typification of two Buckley grass names revisited: Muhlenbergia
texana and M. monticola (Poaceae). Sida 15: 589-591. 1993. [Typified with specimens
from PH; M. monticola a synonym of M. tenuifolia (H.B.K.) Kunth.]
Duncan, W. H. Preliminary reports on the flora of Georgia. 1. The distribution in Georgia of
spermatophytes new and rare to the state. Castanea 13: 70-83. 1948. [M. tenuiflora.]
Eddy, T.L. Muhlenbergia richardsonis in Wisconsin. Mich. Bot. 31 (1): 39, 40. 1992
Edwards, S. Saving the sweetgrass. Soil & water conservation news. U.S. Dept. Agr. 12(5): 15.
1992. [Muhlenbergia filipes in S. Carolina. ]
Fernald, M. L. Muhlenbergia uniflora. Rhodora 29: 10-14. 1927. [Bog sp. occurring in the
Northeast.]
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 101
06/27/17
-----. Five common rhizomatous species of Muhlenbergia. Rhodora 45: 221-239. 1943.
Fowler, N.L., & D.W. Dunlap. Grassland vegetation of the eastern Edwards Plateau, Texas, U.S.A.
Am. Midl. Nat. 115: 146-155. 1986.*
Freeman, O. M. New or noteworthy plants from Polk Co., North Carolina or vicinity. Castanea 21:
41-43. 1956. [M. capillaris, M. sobolifera.]
Freeman, O. M. Notes on the flora of Polk Co., North Carolina Castanea 20: 37-57. 1955. [M.
schreberi, M. sylvatica.]
Glenn Lewin, D.C., & J.M. Ver Hoef. Prairies and grasslands of the St. Croix National Scenic
Riverway, Wisconsin and Minnesota, U.S.A. Prairie Nat. 20: 65-80. 1988.*
Goodding, C. O. Two new species of Muhlenbergia. Jour. Wash. Acad. Sci. 30: 19, 20. 15 Ja
1940.
-----. Three new species of Muhlenbergia. Jour. Wash. Acad. Sci. 31: 504-506. 15 D 1941.
Harper, W. G. Ring growth of plants. Proc Soil Sci. Soc. Amer.. 21(4): 425-428. 1957. [Chiefly
M. torreyi.]
Haygood, R.A., &, O. Barnett. Widespread occurrence of centipedegrass mosaic in South
Carolina. Plant Disease 76: 46-49. 1992. [ Incl. M. schreberi.]
Herrera A., Y., & J.F. Bain. Flavonoid profiles in the Muhlenbergia montana complex (Poaceae).
Biochem. Syst. Ecol. 19: 665-672. 1991.*
& P.M. Peterson. Muhlenbergia cualensis and M. michisensis (Poaceae: Eragrostidea): two new
species from Mexico. Novon 2: 114-118. 1992. [From Durango, Mexico.]
Hudson, J.H. Foxtail muhly new to Saskatchewan, Canada; rigid sedge reaffirmed; additional
range extensions. Blue Jay 46(2): 71-74. 1988. [M. andina (Nutt.) Hitch., a western sp.]*
Lamson-Scribner, F. Notes on Muhlenbergia. Rhodora 9: 17-23. 1907.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 102
06/27/17
Layser, E. F. New distributional records for plants in the Pacific Northwest. Madroño 21: 491,
492. 1972.
Lura, C.L., W.T. Barker, & P.E. Nyren. Range plant communies of the Central Grasslands
Research Station in south central North Dakota, U.S.A. Prairie nat. 20: 177-192. 1988.*
Mackenzie, K. K. &, B. F. Rush. New Plants from Missouri. Trans. Acad. Sci. St. Louis 12: 7889. pl. 12-17. 1902.
Matthei, O. The genus Muhlenbergia Schreber (Poaceae) in Chile. Gayana Bot. 41 (1/2): 53-60.
1984.*
McCarty, E. C. Seasonal march of carbohydrates in Elymus ambiguus and Muhlenbergia gracilis,
and their reaction under moderate grazing use. Plant Physiol. 10: 727-738. f. 1-3. 1935.
McKenzie, P.M., & L.E. Urbatsch. Muhlenbergia frondosa (Poaceae) new to Louisiana. Sida 11:
486-488. 1986.*
Meyer, S.E., & E. Garcia Moya. Plant community patterns and soil moisture regime in gypsum
grasslands of northern central Mexico. Jour. Arid Environm. 16: 147-156. 1989.
Miller, R.F. & Donart, G.B. .Response of Muhlenbergia porteri Scribn. to season of defoliation.
Jour. Range Managem. 34: 91-94. 1981
Mitchell, W.W., & R.W. Pohl. Variation and aneuploidy in Muhlenbergia glomerata. Am. Midl.
Nat. 76: 211-221. 1966.
Morden, C. W., & S. L. Hatch. Numerical analysis of Muhlenbergia capillaris (Poaceae) and its
allies in the southeastern United States. (Abstr.) Am. Jour. Bot. 71 (5-part 2): 179. 1984.
---- & -----. Cleistogamy in Muhlenbergia cuspidata (Poaceae): structure. Sida 10: 254- 255.
1984.*
---- & -----. Vegetative apomixis in Muhlenbergia repens (Poaceae: Eragrostideae). Sida 11: 282Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 103
06/27/17
285. 1986.* [Produces bulbils.]
----- & -----. Anatomical study of the Muhlenbergia repens complex (Poaceae: Chloridoideae:
Eragrostideae). Sida 12: 347-359. 1987.*
---- & -----. An analysis of morphological variation in Muhlenbergia capillaris (Poaceae) and its
allies in the southeastern United States. Sida. 13: 303-314. 1989. Note Includes.
muhlenbergia expansa muhlenbergia filipes
Peterson, P.M. Chromosome numbers in the annual Muhlenbergia (Poaceae). Madroño 35: 320324. 1988.*
-----. Muhlenbergia majalcensis, new species (Poaceae Eragrostideae) from Chihuahua, Mexico.
Syst. Bot. 14: 316-319. 1989.
-----. Leaf blade anatomical survey of Muhlenbergia (Poaceae: Muhlenbergiinae). Sida 19: 469506. 2001.*
_____, C. R. Annable, & V.R. Franceschi. Comparative leaf anatomy of the annual Muhlenbergia
(Poaceae). Nordic Jour. Bot. 8: 575-583. 1989.
-----. & L. H. Rieseberg. Flavonoids of the annual Mulhenbergia. Biochem. Syst. Ecol. 15: 647652. 1987.*
Peterson, P.M., M.R. Duvall &, A.H. Christensen.Allozyme differentiation among Bealia
mexicana, Muhlenbergia argentea, and M. lucida (Poaceae: Eragrostideae). Madroño 40:
148-160. 1993.
Peterson, P.M., J.K. Wipff & S.D. Jones. Muhlenbergia pilosa (Poaceae: Eragrostideae), a new
species from Mexico. Madroño 39: 150-154. 1992.
----- & J.J. Ortiz-Diaz. Allelic variation in the amphitropical disjunct Muhlenbergia torreyi
(Poaceae: Muhlenbergiinae). Brittonia 50: 381-391. 1998.*
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 104
06/27/17
Pinson & Batson. Jour. Elisha Mitchell Sci. Soc. 87(4): 188-191. 1971.*
Pizzolato, T.D. Patterns of procambial development in rachilla and floret of Muhlenbergia
torreyana (Gramineae, Chloridoideae). International Jour. plant sciences.. 156: 603-614.
1995
Pohl, R. W. Muhlenbergia, subgenus Muhlenbergia (Gramineae) in North America. Am. Midl.
Nat. 82: 512-542. 1969. [M. glomerata transcontinental in glaciated areas to B.C., s. to W.
Va., one sta. in mts. of NC.; rejects Koyama & Kawano's union of M. frondosa and M.
ramosa.]
Pohl, R. W., & W. W. Mitchell. Cytogeography of the rhizomatous American species of
Muhlenbergia. Brittonia 17: 107-112. 1965.
Reeder, C. G. Muhlenbergia minutissima (Steud.) Swallen and its allies. Jour. Wash. Acad. Sci.
39: 363-367. 1949.
Reeder, J.R., & C.G. Reeder. Aneuploidy in the Muhlenbergia subbiflora complex (Gramineae).
Phytologia 65: 155-157. 1988.*
----- & -----. The resurrection of a species: Muhlenbergia straminea (Gramineae). Phytologia 78:
417-427. 1995.
Salzman, F. & J. Kells. Controlling wirestem muhly. Extension bulletin E - Cooperative
Extension Service, Michigan State University. 1992. (2256, rev.) 2 p. [Muhlenbergia
frondosa.]
Scholz, H. Eine neue Muhlenbergia (Gramineae) aus den bolivianischen Anden. Willdenowia
14: 393-395. 1994. [M. nimuscula; ill.]
Scott, W. O., & F. W. Slife. A new cornfield weed menace. Crops & Soils 12(8): 19, 20. 1960.
[Muhlenbergia frondosa and its control; not really that scary.]
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 105
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Smith, M. Effect of temperature on rhizome regrowth and biomass in Muhlenbergia sobolifera,
a shade-tolerant C4 grass. Trans. Illinois State Acad. Sci. 85: 19-28. 1992. [. ]
----- &, C.E. Martin.. Growth and morphological responses to irradiance in three forest
understory species of the C4 grass genus Muhlenbergia. Bot. Gaz.. 148: 141-148. 1987.
[ill. Note Includes references. muhlenbergia. muhlenbergia schreberi. forest trees.
carbon pathways. shade. solar radiation. growth. plant morphology. muhlenbergia
frondosa muhlenbergia sobilifera muhlenbergia cuspidata a sun tolerant plant Other
----- & -----. Field studies of photosynthetic and growth responses to irradiance in three forest
understory species of the C4 grass genus Muhlenbergia. Bot. Gaz. 148: 456-462. 1987.*
-----. & Y. Wu. Photosynthetic characteristics of the shade-adapted C4 grass Muhlenbergia
sobolifera (Muhl.) Trin.: control of development of photorespiration by growth
temperature. Plant, cell and environment 17: 763-769. 1994 . [The most shade adapted
C4 plant documented to date.]
Soderstrom, T.R. Taxonomic study of subgenus Podosemum and section Epicampes of
Muhlenbergia (Gramineae). Contr. U.S. Natl. Herb. 34 (4): 75-189. 1967.
Swallen, J. R. New United States grasses. Jour. Wash. Acad. Sci. 31: 348-355. f. 1-8. 1941.
[Muhlenbergia villosa Swallen, sp. nov., from Stanton, Tex.]
Swallen, J. R. The awnless annual species of Muhlenbergia. Contr. U.S. Nat. Herb., 29: 203-208,
1947.
Uttal, L. J., & R. S. Mitchell. Amendments to the flora of Virginia--II. Castanea 37: 96-118. 1972.
[Muhlenbergia frondosa var. *commutata, M. glabriflora, M. glomerata.]
14. Lycurus, s US
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 106
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attributed to southeastern U.S. by Peterson et al.; chloroplast DNA analysis supports close
relationship with Muhlenbergia (Duvall et al. 1994)
15. Chloris Swartz, Prodr. Veg. Ind. Occ. 25, 1788.*
Perennials or annuals of disturbed open places, typically in somewhat moist rather than well
drained soils. Rhizomes often present; stolons sometimes also present. Stems erect, simple or
branched near basal leaves cauline and basal, few; sheaths sharply keeled, glabrous, ligule a fringe
of white hairs; blade linear, flat or plicate. Inflorescence a panicle, with a few spicate or subspicate
branches, these branches ± digitately or spicately arranged at the top of the stem; rachis 3-sided.
Spikelets laterally compressed, 1 (-2) (several) flowered, arranged in two rows on two sides of the
rachis, sessile or short-pedicellate, the pedicel often pilose; glumes 2, unequal, lanceolate, the
second about twice as long as the first, and about as long as the first lemma; rachilla short,
persistent; flowers of two sorts, the basal bisexual, those above it staminate, rudimentary, or sterile;
lemmas oblong-ovate, slightly emarginate, awnless or with an awn (arising from the notch at apex
of lemma) as much as 2 (3) times as long as the spikelet, 3-nerved (the lateral nerves marginal,
sometimes ciliate distally); paleas narrowly elliptic, a little shorter than the lemmas, the apex bifid,
strongly 2-nerved, the nerves glabrous or strongly ciliate. Lodicules 2, cuneate, truncate, fleshy.
Stamens 3; anthers ellipsoid. Ovaries ovoid to ellipsoid, glabrous; styles 2, separate, short, terminal
stigmas plumose. Fruit an achene (the pericarp ± free), subterete to cylindric, ellipsoid to
lanceolate; hilum linear, nearly as long as grain; embryo large, typically1/3 –3/4 the length of
the grain. Base chromosome number 10. (synonym). Type species: Agrostis cruciata L., [ = C. ---], was designated by Hitchcock (1920). (Named for Chloris, in Greek mythology the mother of
Nestor, goddess of flowers.). – Fingergrass.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 107
06/27/17
A genus of about 55 to 60 species, worldwide in tropical and warm temperate regions.
Several species are significant weeds, and Chloris gayana, Rhodes grass, is an important forage
species in many tropical countries.
C. virgata Sw., feeatehr windmill grass, 2n=20, 26, 30, 40, a pantropic species occurring in all the
southeastern states, as well as the southern plains and southwest.
C. verticillata, Arkansas (Smith) (Attributed to Louisiana by Hitchcock & chase, but not confirmed
by either Allen or MacRoberts.)
C. cucullata from Mississippi (Harrison Co., 1978, MO) to New Mexico southward.
C. barbata Sw., swollen windmill grass, 2n=20, 40, occurs in southern Florida (Monroe and
Miami-Dade counties), south trough the Caribbean and easteren Mexico to South Amreica. It is a
weedy species, often found in calcareous soils.
C. canterae Arech., 2n = 36, Louisiana (Acadia, Beauregard, Caldwell parishes; Allen; but only
Acadia, fide MacRoberts), South Carolina, and Texas; native to south America
C. subdolichostachya 2n = 72 (Gould 7699, MO, Lampasas Co., Texas).
C. truncata R.Br. is an Australian species once found on wool waste in South Carolina (Shinners)
C. gayana Kunth, Rhodes grass, 2n=20, 30, 40, occurs in florida, Mississippi, Louisiana
(MacRoberts, Thieret) ; Texas to California. A native of Africa, it is widely cultivated as a pasture
grass in tropical and warm temperate areas worldwide.
Chloris elata Desv., tall windmill grass, 2n=72, occurs in peninsular Florida, and southward
through the Caribbean to Argentina and Peru.
C. ciliata Texas
C. andropogonoides Fourn. Texas
C. verticillata Nutt., Tumble windmill grass, 2n= ca. 28, 40, 63, is widespread in the souther plains
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 108
06/27/17
and southwest. In our area it has only been found in northwest Arkansas and Tennessee. It was
reported from Louisiana by Hitchcock & Chase, but neither Allen or MacRoberts could find a
supporting specimen. In recent decades it has been spreading in the Midwestern states, especially
Illinois (Tucker).
C. virgata Sw. is widespread in Louisiana (Macr 1989).
Chloris ventricosa R.Br., plump windmill grass, is an Australian species that has been found in the
U.S. only once, around woolen mills in South Carolina. Chloris truncate R.Br., 2n=40, is another
Australian species, similarly collected in SouthCarolina (as well as southern California).
Chloris divaricata R.Br., spreading windmill grass, is yet another Australian grass found around
woolen mills in South Caroliina; it has since become established in the Southwest (New Mexico,
Texas).
Chloris pectinata Bisch., hooded windmill grass, 2n=40, is a species onf Mexico, Texas, New
Meixoc, Oklahoma, and Kansas; it has also been found as a wool waif in South Carolina.
A number of species have been investigated for germination characterisitics. Chloris inflata
produces seeds without dormancy ( Van Rooden et al. 1970). The optimum temperature for
germination shows a wide range: 31° in C. pilosa (Elberse & Brenen 1989), 32° in C. virgata (
Lodge & Whalley 1992), 20-30° in C. truncata (Jurado & Westoby 1992) and only 20 ° in C.
scariosa (Hacker 1989)
REFERENCES
Under subfamily references see: Bor; Clayton & Renvoize; Hitchcock & chase; Jurado & Wesoby;
Purseglove.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 109
06/27/17
Anderson, D. Introgression in Texas species of Chloris (Gramineae). (Abstr.) Am. Jour. Bot. 52:
6747. 1965.
-----. The grass genus Chloris and its relatives. XI. Int. Bot. Congr. Abstr. p. 3. 1969.
-----. Taxonomy of the genus Chloris (Gramineae). Brigham Young Univ. Sci. Bull., Biol. Ser.
19(2): 1-133. 1974. [peterson et al. 2001 give vol # as 29]
Barkworth, M.E. Chloris. Pp. 204--218 in Barkworth, M.E., K.M. Capels, S. Long, & M.B. Piep,
eds. Flora of North America, north of Mexico. Magnoliophyta: Commelinidae (in part): Poaceae,
part 2. New York and Oxford. 2003.
Brown, L. E. A biosystematic study of a Texas Chloris complex. (Abstr.) Am. Jour. Bot. 54: 657.
1967.
Elberse, W.T., & H. Breman. Germination and establishment of Sahelian rangeland species. I.
Seed properties. Oecologia 80: 477-484. 1989.*
Gould, F. W. Parapholis incurva and Chloris polydactyla in Texas. Field & Lab. 23: 83-84.
June/Oct. 1955. [not in Grass manual]
Cowan, R.T., R.J. Moss, & D.V. Kerr. ----------------------------------Trop. Grassl. 27: 150-161.
1994.*
Ehrlich, W. K, R.T. Cowan, R.L. Romano, D.A. Crouch, & K.F. Lowe. Management of Rhodes
grass (Chloris gayana cv. Callide) during autumn. Animal Production in Australia: Proc.
Austral. Soc. Animal Production 20: 370. 1994 [useful in subtropical pastures.]
Kartesz, J.T., & K.N. Gandhi. Chloris barbata Sw. and C. elata Desvaux (Poaceae), the earlier
names for C. inflata and C. dandeyana Adams. Rhodora 94: 135-140. 1992.*
Mauldin, M. P. The occurrence of two fertile florets in the spikelets of Chloris. Science 105: 336Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 110
06/27/17
337. 1947.*
Nash, G. V. A Revision of the Genera Chloris and Eustachys in North America. Bull. Torrey Bot.
Club 25: 432-450. 1898. [Chloris tenuispica and C. brevispica, sp. nov., and Chloris
Nealleyi, C. latisquamea, C. Texana (Vasey) and Eustachys neglecta, nom. nov.]
Parodi, L. R. Una nueva especie de Graminea del genero Chloris y sus relaciones con los
Gymnopogon. Rev. Argent. Agron. 12: 45-50. f. 1. 20 Mr 1945.
Prakash, K.S., A. Mani, & T. Al Zidgali. Effect of nitrogen, phopohorus and potassium fertilisation
on herbage yield and quality and plant parasitic nematode populations in an irrigated
Rhodes grass (Chloris gayana) pasture in Oman. Trop. Grassl. 28: 164-169. 1994.
Shinners, L. H. Chloris truncata R. Br. (Gramineae), an Australian wool-waste adventive in South
Carolina. Sida 5: 182. 1973.
Thieret, J. W. Additions to the flora of Louisiana. Castanea 28: 169, 170. 1963. [C. gayana, 169].
16. Eustachys Desv., Nouv. Bull. Sci. Soc. Philom. 2: 188. 1810.* Check to see if at Harvard
Perennials or annuals of disturbed open places, typically in somewhat moist rather than well
drained soils. Rhizomes often present; stolons sometimes also present. Stems erect, simple or
branched near basal leaves cauline and basal, few; sheaths sharply keeled, glabrous, ligule a fringe
of short white hairs; blade linear, flat or plicate. Inflorescence a digitate group of 2-many racemes;
rachis 3-sided. Spikelets laterally compressed, 1 (-2)-flowered, arranged in two rows on two sides
of the rachis, short-pedicellate, the pedicel often pilose; flowers bisexual (occasionally a second
upper floret has a staminate flower only); glumes 2, unequal, lanceolate, the second about as long as
the lemma; lemmas oblong-ovate, acute to emarginate, awnless or mucronulate, 1 nerved, keeled;
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 111
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paleas elliptic, a little shorter than the lemma, the apex bifid, awnless or with an awn about as long
as the spikelet, strongly 2-nerved, the nerves glabrous or strongly ciliate. Lodicules 2, cuneate,
truncate, fleshy. Stamens 3; anthers ellipsoid. Ovaries ovoid to ellipsoid, glabrous; styles 2,
separate, short, terminal stigmas plumose. Fruit a caryopsis, subterete to cylindric, oblong, or to
broadly ellipsoid; hilum linear, nearly as long as grain; embryo large, typically1/3 –3/4 the
length of the grain. Base chromosome number 10. (synonym). Type species: Eustachys L., [ = C.
----], was designated by Hitchcock (1920). (Named for from Greek, eu-, true, and stachys, spike.).
– Fingergrass.
Close-knit group barely separable from Chloris
E. floridana: GA, FL, AL (MO)
E. glauca: Florida
E. neglecta: Florida
E. petraea: ALA, MS, GA, FL, LA (fide MacRoberts), TX
References
Under Chloris see Nash.
Jones, S.D., & J.K. Wipff. Eustachys retusa (Poaceae), the first report in Florida and a key to
Eustachys in Florida. Phytologia 73: 274-276. 1992.
Molina, A.M. Revisión taxonómica del género Eustachys Desv. (Poaceae: Chloridoideae,
cynodonteae) de Sudamérica. Candollea 51: 225-272. 1996.*
17. Trichloris Bentham, Jour. Linn. Soc. Bot. 19: 102. 1881.* [check at GH....
Description:
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 112
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(Name from...
Type species:
Two species, tropical south America.
None in Smith or MacR; unlikely to be present based on Texas distribution
Questionable whether either species native or naturalized in SE.
T. crinita: Texas to Arizona (MO) but not in smith
T. pleurifolia: MS, Biloxi, cult. ?; Texas to Arizona to bolivia (Gould)
Texas: Cameron Co., Kenedy Co., Laredo Co.
18. Schedonnardus Steudel, Syn. Pl. Glum. 1:146. 1854.
Low perennials of grasslands. Rhizomes short, erect. Stems erect with decumbent base;
leaves cauline and basal, few; sheaths compressed, keeled, glabrous, ligule membranous, ovatelanceolate (1-3 mm long); blade linear, plicate, with conspicuous midrib, scabrous on the margins.
Inflorescence a panicle, terminal comprising about half or more of the height of the plant, the
branches widely spaced, unbranched, breaking off from the plant as a unit, dispersing as a
“tumbleweed.” Spikelets widely spaced on the inflorescence branches, slightly laterally
compressed, 1 (-2) flowered, sessile or short-pedicellate; glumes 2, unequal (the second about twice
as long as the first), shorter than the lemma, lanceolate, acute, 1-nerved, glabrous; rachilla short,
persistent; articulated below the glumes; lemmas lanceolate, 3-nerved, acute to mucronulate,
glabrous or scabridulous; paleas lanceolate, scarcely shorter than the lemmas, the apex acute, 2nerved, glabrous. Lodicules 2, cuneate, truncate, oblong, fleshy. Stamens 3; anthers ellipsoid.
Ovaries ellipsoid, glabrous; styles 2, separate, short, terminal stigmas plumose. Fruit a caryopsis,
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 113
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scarcely clasped by lemma and palea, subterete to cylindrical, narrowly ellipsoid; hilum linear,
nearly as long as grain; embryo about 1/3 the length of the grain. Base chromosome number 10.
Type species: S. texanus Steudel [=S. paniculatus (Nutt.) Trel.], the only species included in the
prologue. (Name from Greek schedon, near and Nardus, a genus of Pooideae that Steudel thought
was a close relative.) – Tumblegrass.
A monotypic genus endemic to the prairies of North America. Introduced in Argentina.
Steudel included this genus in his tribe Agrostideae, because of the one-flowered spikelets. It is an
isolated genus. S. paniculatus (Nutt.) Trel., tumblegrass, 2n = 20, 30 occurs from Saskatchewan and
Montana south to Texas, northeastern Mexico, and Arizona; it is also introduced Argentina. In our
area it is known from Arkansas (Franklin, Marion, and Washington cos. in northwest part of the
state) and Louisiana (Caddo parish in NW corner of the state; MacRoberts 1989). The spikelets
show considerable similarity to those of Sporobolus and Muhlenbergia, but the genus differs greatly
in the form of the inflorescence which is more like Cynodon.
Gould noted that although S. paniculatus is widespread throughout the grasslands of central
North America, it contributes little forage for cattle and horses, because the stems have few leaves
and the blades are narrow. Tumblegrass tends to increase in abundance with overgrazing, as do
many annual grass species.
19. Gymnopogon Palisot de Beauvois, Essai Agrost. 41. 1812.
Perennials [annuals]of typically wooded areas. Rhizomes short, knotty. Stems erect, often
branched fom the upper nodes; leaves numerous, cauline, distichous; sheaths sub-terete, glabrous
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 114
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except for a few hairs around the collar, ligule a short membranous structure; blade linearlanceolate, abruptly narrowed at the base, flat or slightly plicate, without midrib, glabrous.
Inflorescence a panicle, with many spicate branches, these branches solitary or two together at
lower nodes, bearing spikelets their entire length; rachis slender. Spikelets slightly laterally
compressed, narrowly lanceolate, 2-flowered, short-pedicelled, the pedicel slightly scabrous;
glumes 2, unequal, the second longer than the first lemma, narrowly lanceolate, 1-nerved, apex
acute, awned; disarticulation above the glumes; rachilla short, persistent; flowers of two sorts, the
basal bisexual, the one above it sterile, reduced to a narrow awned lemma; lemmas narrowly
lanceolate, bidentate, with apical awn slightly shorter than to two times longer than lemma, 3nerved, ciliate with a few long hairs near the base; paleas narrowly elliptic, a little shorter than the
lemmas, the apex acute, awnless, the nerves glabrous (??). Lodicules 2, cuneate, truncate, fleshy.
Stamens 3; anthers ellipsoid. Ovaries ovoid to ellipsoid, glabrous; styles 2, separate, short, terminal
stigmas plumose. Fruit a caryopsis, subterete to cylindric, narrowly ellipsoid; hilum linear, nearly
as long as grain; embryo large, typically 1/3 –3/4 the length of the grain. Base chromosome
number 10. (Anthopogon Nutt., Alloiantheros Elliot ex Raf., Biatherium Desv., Sciadonardus
Steud., Monochaete Dell., Doellochloa Kuntze). Type species: G. racemosus Beauv. L., [ = G.
ambiguus (Michx.) BSP.]. (Name from Greek gymnos, naked, and pogon, head, the application
here uncertain.) – Skeletongrass.
A genus of about 15 species, in the warm temperate and tropical regions of the New World.
A single species, G. delicatulus (C.B. Clarke ex Hooker f.) N.L.Bor occurs in India and Burma
(Bor). The only southeastern species, G. ambiguus (Michx.) BSP. [G. brevifolius Trin.], 2n=40,
occurs from Pennsylvania, Indiana and Kansas south to Florida and eastern Texas.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 115
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References
Under subfamily references see: Bor, Clayton & Renvoize, Gould
Smith. ------- Iowa St. Jour. Sci. 45: 319-385. 1971.
20. Ctenium Panzer, Id. Rev. Gräser 36, 59. 1813, nom. cons.
Perennials or annuals of disturbed open places, typically in somewhat moist rather than well
drained soils. Rhizomes often present; stolons sometimes also present. Stems erect, simple or
branched near basal leaves cauline and basal, few; sheaths sharply keeled, glabrous, ligule a fringe
of short white hairs; blade linear, flat or plicate. Inflorescence a panicle, with a few spicate or
subspicate branches, these branches ± digitately or spicately arranged at the top of the stem; rachis
3-sided. Spikelets laterally compressed, 2-4 flowered, arranged in two rows on two sides of the
rachis, sessile or short-pedicellate, the pedicel often pilose; glumes 2, unequal, lanceolate, the
second about as long as the first lemma; rachilla short, persistent; flowers of two sorts, the basal
bisexual, those above it staminate or sterile; lemmas oblong-ovate, 1-5 (most often 3) nerved;
paleas elliptic, a little shorter than the lemmas, the apex bifid, awnless or with an awn about as long
as the spikelet, strongly 2-nerved, the nerves glabrous or strongly ciliate. Lodicules 2, cuneate,
truncate, fleshy. Stamens 3; anthers ellipsoid. Ovaries ovoid to ellipsoid, glabrous; styles 2,
separate, short, terminal stigmas plumose. Fruit a caryopsis, subterete to cylindric, oblong, or to
broadly ellipsoid; hilum linear, nearly as long as grain; embryo large, typically1/3 –3/4 the
length of the grain. Base chromosome number 10. (Monocera Elliot; Triantherus Raf.;
Monanthera Raf.; Aplocera Raf.). Type species: C. carolinianum Panzer [=C. aromaticum], was
designated by ?. (Named from Greek, ctenion, comb.). – combgrass
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 116
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southern parishes of Louisiana (MacRoberts)
Reference
Clayton, D. Kew. Bull. 16: 471-476. 1963.
Longhi-Wagner, H.M. Circunscriçao de algunas especies brasilieras do gênero Ctenium Panzer
(Gramineae). Acta Bot. Bras. 1, Suppl.: 53-62. 1988* [6 spp., illus.]
21. Cynodon L.C. Richard in Persoon, Syn. Pl. 1: 85. 1805, nom. cons.
Low mat or sward forming perennials of lawns, roadsides, pastures [savannas]. Rhizomes
present, scaly, horizontal, extensive; stolons present, arching, with sub-opposite leaves because of
the alternating long and short internodes. Stems erect, cespitose, unbranched; leaves cauline, few;
sheath glabrous or ciliate; ligule a ciliate membrane; blade linear, flat or plicate, glabrous except
for some long soft hairs near base. Inflorescence of 2-7 digitately arranged branches, each branch
spicate. Spikelets laterally compressed, 1-(2) flowered; glumes 2, subequal, ovate-lanceolate, 1nerved; rachilla disarticulating above glumes; flowers bisexual (the upper one sterile or
rudimenttary); lemmas broadly elliptic, 3-nerved (the lateral nerves marginal), acute, smooth,
usually villous along midvein; paleas elliptic, a little shorter than the lemmas, 2-nerved. Lodicules
2, cuneate, truncate, fleshy. Stamens 3; anthers narrowly ellipsoid. Ovaries ovoid to ellipsoid,
glabrous; styles 2, separate, short, terminal stigmas plumose. Fruit a caryopsis, terete, oblong;
hilum linear, nearly as long as grain; embryo large, typically1/3 –3/4 the length of the grain.
Base chromosome number 9. [Capriola Adans.] (Name from...Type species: C. dactylon (L.)
Kuntze). – Bermuda Grass, Star Grass, Dhub..
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 117
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A small genus of about 8 spp. The only America species, Cynodon dactylon, Bermuda grass, 2n=
36 (18 and 40 also reported), native to tropical Africa, is now worldwide in tropoical and warm
temperate regions. In the U.S., it ranges from Connecticut to Washington, south to Florida, Texas,
and California, but is uncommon or sporadic in the northern half of the country. I have seen
specimens from all the southeastern states. It is an important forage grass in the southeastern states,
and a common lawn grass in warm humid regions, and is also used for pasturage in tropical and
subtropical climates..
A number of medicinal uses have been ascribed to C. dactylon. A decoction of the roots is widely
used as a diuretic in the west Indies and latin America (Morton). Under some conditions, the plant
has a high concentration of hydrocyanic acid and can cause intoxication in cows and horses.
Cynodon nlimfuensis Vanderyst, African Stargrass, 2n=18, a large, often sterile species, is
cultivated and escaped in Costa Rica (Pohl). Also naturalizedin Texas (Jones & Jones) It is native
to Africa.
References:
Under subfamily see Ashraf & Yasmin; Bor; Clayton & Renvoize; Hitchcock & Chase; Marcum &
Murdoch; Morton; Pohl; Purseglove; Rasmussen & Rice.
de Wet & Harlan. --- Taxon 19: 565-569. 1970.
Dudeck, A.E., S. Singh, C.E. Giordano, T.A. Tell, & D.B. McConnell. Effects of sodium chloride
on Cynodon turfgrasses. Agron. Jour. 75: 927-930. 1983.*
Gatschet, M.J., C.M. Taliaferro, D.R. Porter, M.P. Anderson, J.A. Anderson, & K.W. Jackson. A
cold-regulated protein from bermudagrass is a chitinase. Crop Sci. 36: 712-718. 1996.*
Harlan, de Wet, & Richardson. --- Am. Jour. bot. 56: 944-950. 1969.
Hill, G.M., R.N. Gates, & G.W. Burton. Forage quality and grazing steer performance from Tifton
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 118
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85 and Tifton 78 Bermudagrass pastures. Jour. Animal Sci. 71: 3219-3225. 1993.*
Jones, S.D., & G.D. Jones. Cynodon nlemfuensis (Poaceae: Chlorideae) previously unreported for
Texas. Phytologia 72: 93-95. 1992.*
Miller, G.L., & R. Dickens. Bermudagrass carbohydrate levels as influenced by potassium fertilizer
and cultivar. Crop Sci. 36: 1283-1289. 1996.*
_____ & _____. Potassium fertilization related to cold resistance in bermudagrass. Crop Sci. 36:
1290-1295. 1996.*
Ramakrishnan, P.S., & R. Nagpal. Adaptation to excess salts in an alkaline soil population of
Cynodon dactylon (L.) Pers. Jour. Ecol. 61: 369-381. 1973.*
Shashikumar, K., & J.L. Nus. Cultivar and winter cover effects on bermudagrass cold aclimation
and crown moisture content. Crop Sci. 33: 813-818. 1993.*
Youngner, V.B., & O.R. Lunt. Salinity effects on roots and tops of bermuda grass. Jour. brit.
Grassland Soc. 22: 257-259. 1967.*
Geng SL, Zhao S, Wu H, 2002., (Anatomical characters of stems and leaves of three lawn
grasses.) J. Trop. Subtrop. Bot. 10. (2): 145-151 (2002) - illus.
Roodt R, Spies JJ, 2002., Chromosome studies on African plants. 18. The subfamily
Chloridoideae. Bothalia 32. (2): 240-249 (2002) - illus.
Gorski P, 1999., Cynodon dactylon (Poaceae) in Poland. Fragm. Flor. Geobot. suppl.7. 65-71
(1999) - illus. Liu JX, He SA, Liu YD, Chen SL, 1996., (Taxonomy of Cynodon dactylon types
in east China and their turfgrass quality.) J. Pl. Resourc. Environ. 5. (3): 18-22 (1996) - Speranza
M, 1995., Morphology and phenology of Cynodon dactylon (L.) Pers. (Gramineae) in Italy.
Webbia 49. (2): 225-237 (1995) - Santos AMPV dos, Boechat S de C, 1994., (Cynodon
(Poaceae, Chloridoideae) in Rio Grande do Sul, Brazil.) (In Portuguese). Iheringia, Bot. no.44.
85-102 (1994) - illus.
Jones SD, Jones GD, 1992., Cynodon nlemfuensis (Poaceae : Chlorideae) previously unreported
in Texas. Phytologia 72. (2): 93-95 (1992) –
Nowack R, 1992., What is Cynodon radiatus Roth ex R. and S. (Poaceae)? Blumea 36. (2): 477478 (1992) - Kumar P, Sachdeva SK, 1988., Karyomorphological observations and biochemical
variability in triploids of Cynodon dactylon (L.) Pers. complex. Acta Bot. Indica 16. (2): 143-152
(1988) –
Romero Zarco C, 1986., Notas taxonomicas y corologicas sobre la flora de Andalucia
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 119
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Occidental: 141 - 257. 160 - 257. Notas breves: 244. Cynodon dactylon var. affinis. (Caro &
Sanchez) Romero Zarco, comb. et stat. nov. Lagascalia 14. (1): 171 (1986) –
Vegetti AC, 1986., Contribucion al conocimiento de las inflorescencias en Chloris y Cynodon
(Poaceae). Kurtziana 18. 109-120 (1986) - illus.
Moreira I, 1984., Biology of bermudagrass (Cynodon dactylon (L.) Pers). Rev. Biol., 12. (3-4):
519-528 (1983-1984) - illus.
Caro JA, 1983., Cuatro especies y una variedad nuevas de Cynodon. Dominguezia, no.6. 5-20
(1983) - illus.
Oross JW, Thomson WW, 1982., The ultrastructure of the salt glands of Cynodon and Distichlis
(Poaceae). Amer.J.Bot. 69. (6): 939-949 (1982)- illus.
Aldous DE, 1980., Know your turfgrass: 6. Hort. New Zealand, No.14. 16 - 17 (1980)
Chheda HR, KM, Rawal. Phylogenetic relationships in Cynodon: 1. C. aethiopicus, C.
nlemfuensis and C. transvaalensis. Fyton 28(1): 15-21. 1971.
De Wet JMJ, Harlan JR, 1971., South African species of Cynodon (Gramineae). (SuidAfrikaanse soorte van Cynodon (Gramineae)). J.S. Afr. Bot. 37. (1): 53-56 (1971)
Rawal KM, Chheda HR, 1971., Phylogenetic relationships in Cynodon: 2. C. nlemfuensis, C.
dactylon vars. afghanicus and aridus. Fyton 28. (2): 121-130
Rawal KM, Harlan JR, 1971., The evolution of growth habit in Cynodon L. C. Rich
(Gramineae). Trans. Illin. Acad. Sci. 64. (2): 110-118 (1971)
21. Spartina Schreber, Gen. Pl. ed. 8., 43, 1789.
Perennials of freshwater marshes, ditches, and tidal marshes. Rhizomes usually present.
Stems single or ceaspitose, erect or decumbent, simple; leaves cauline, few; sheaths sharply keeled,
glabrous, ligule a fringe of short white hairs; blade linear, flat or involute. Inflorescence a narrow
panicle, with a few secund spicate branches, these branches spicately arranged at the top of the
stem; rachis 3-sided. Spikelets strongly laterally compressed, narrowly oblong, 1-flowered,
arranged on one side of the rachis, sessile; glumes 2, unequal, lanceolate, acute or obtuse,
mucronate or aristate, the second about as twice as long as the first; rachilla short, persistent;
disarticulation at base of glumes; lemmas oblong, 1-3 (-5) nerved; paleas narrowly elliptic, a
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 120
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equalling or a little longer than the lemmas, the apex blunt or emarginate, awnless, strongly 2nerved, the nerves glabrous or strongly ciliate. Lodicules 2, cuneate, truncate, fleshy. Stamens 3;
anthers linear. Ovaries ovoid to ellipsoid, glabrous; styles 2, separate, short, terminal stigmas
plumose. Fruit a caryopsis (the pericarp “reluctantly free” –Clayton & Renvoize), subterete,
fusiform; hilum linear, nearly as long as grain; embryo large, typically1/3 –3/4 the length of
the grain. Base chromosome number 10. Type species: S. cynosuroides (L.) Roth. (Trachynotia
Michx.) (Named for ) – Cordgrass.
Most of the specimens at MO checked by Mobberley in 1953
S. alterniflora
S. bakeri
S. cynosuroides
S. patens
S. spartinae
S. cespitosa
S. foliosa
S. pectinata
References:
See Johnson-Green et al., Smart & Barko;
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 121
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Akers, S. W., C. E. Anderson, & U. Blum. Characterization of vacuolar bodies in Spartina
alterniflora: I. Formation, development, morphology, and ultra-structure. Amer. J. Bot.
64(6): 635-640. 1977. ------, ----- & -----. Characterization of vacuolar bodies in Spartina
alterniflora: II. Some physical and chemical properties. Amer. J. Bot. 64(6): 641-648.
1977.
Anderson, R.C., B.C. Ebbers, & A.E. Liberta. Soil moisture influences colonization of prairie
cordgrass (Spartina pectinata Lind.) by vesicular-arbuscular mycorrhizal fungi. New Phytol. 102:
523-527. 1986.*
Angell, R.F., J.W. Stuth, & D.L. Drawe. Diets and liveweight changes of cattle grazing burned gulf
cordgrass. Jour. Range Managem. 39: 233-236. 1986.*
Bertness, M.D. Peat accumulation and the success of marsh plants. Ecol. Publ. Ecol. Soc. Am. 69:
703-713. 1988.
Blum, U. Photosynthesis and respiration of Spartina alterniflora Loisel. (Poaceae) in North
Carolina salt marshes: model validation (Abstr.) ASB Bull. 24: 37. 1977.
Booker, F.L., E.D. Seneca, & E. blum. Effects of weathered fuel oil on solute leakage, respiration
and plasmolysis in exsiced root sections of Spartina alterniflora Loisel. and Limonium
carolinianum (Walt.) Britt. Envir. Exp. Bot. 29: 249-259. 1989.
Bradley, P.M, & E.L. Dunn. Effects of sulfide on the growth of three salt marsh halophytes of the
southeastern united States. Am. Jour. bot. 76: 1707-1713. 1989. *[Spartina spp.]
Broome, S.W., E.D. Seneca, & W.W. Woodhouse, Jr. Long term growth and development of
transplants of the salt marsh grass Spartina alterniflora. Estuaries 9: 63-74. 1986.*
Broughton, W. S., & K. L. Webb. Investigations on certain factors affecting growth and
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productivity of Spartina alterniflora Loisel. (Abs.) ASB Bull. 10(2): 24. 1963.
Brown, S. D., & W. H. Queen. Root inundation as a factor affecting the vertical position of
Spartina alterniflora in coastal marshes. (Abstr.) ASB Bull. 18: 28. 1971.
Burdick, D. M., & I. A. Mendelssohn. Waterlogging responses in dune, swale and marsh
populations of Spartina patens under field conditions. Oecologia 74: 321-329. 1987.*
Buschsbaum, R., & I. Valiela. Variability in the chemistry of estuarine plants and its effect on
feeding by Canada geese. Oecologia 73: 146-153. 1987.*
Caldwell, P. A. The spatial development of Spartina colonies growing without competetion. Ann.
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Chalmers, A.G. The effects of fertilization on nitrogen distribution in a spartina alterniflora salt
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Chater, E. H., & H. Jones. New forms of Spartina townsendii (Groves). Nature (London) 168:
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Chevalier, A. La distribution géographique et le nomenclature des Spartina des vases salées dans
l'Ancien et dans le Nouveau-Monde. Dernière opinion. Rev. Internatl. de Bot. Appl. et
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Church, G. L. Cytotaxonomic studies in the Gramineae: Spartina, Andropogon and Panicum. Am.
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Jour. Bot. 27: 263-271. f. 1-33. 15 My 1940.
Cranford, P.J., P. Schwinghamer, & D.C. Gordon, Jr. Identification of microdetrius derived from
spartina and its occurrence in the water column and intertidal sediments of Cumberland
Basin, Bay of Fundy. Estuaries 10: 108-117. 1987.*
Cruz, A. A. de la, C. T. Hackney, & J. P. Stout. Aboveground net primary productivity of three
gulf coast marsh macrophytes in artificially fertilized plots [Juncus roemerianus, Spartina
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47: 7-8. [059917].
Cunningham, B. Grass in a turtle egg. Am. Nat. 65: 478-479. illust. S-0 1931.
Curtis, P.S., B.G. Drake, P.W. Leadley, W.J. Arp, & D.F. Whigham. Growth and senescence in
plant communities exposed to elevated CO2 concentraions on an esturine salt marsh.
Oecologia 78: 20-26. 1989.
Dai, T. & R.G. Wiegert. Ramet population dynamics and net aerial primary production of Spartina
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06/27/17
25a. Chondrosum ??? none at ILL from SE; pLANTS USDA treats all as synonyms of
Bouteloa
22. Bouteloua Lagasca, Varied. Ci. 2, 4: 134. 1805, emend. Lagasca, Gen. Sp. Nov. 5. 1805
[1816?], nom. cons. [GH: Tax L13.3 E]
Perennials [annuals] of prairies and open woodlands, generally in well drained soils.
Rhizomes often present; stolons sometimes present. Stems single or ceaspitose, erect or
decumbent, simple; leaves mostly basal, few; sheaths keeled, glabrous, ligule a fringe of short
white hairs; blade linear, flat or folded. Inflorescence a simple or compound raceme; branches if
present erect or arcuate, sometimes reflexed. Spikelets strongly laterally compressed, narrowly or
broadly oblong, 2-4 flowered, arrangedin two rows on one side of the rachis, stipitate; glumes 2,
unequal to nearly equal, lanceolate, acute, awnless or short-awned, 1-nerved; rachilla short,
disarticulating at base of branch or above glumes; flowers of two sorts: the lowermost bisexual, the
upper 1-3 staminate or sterile; lemmas oblong, 3-nerved, acute or shortly 3-awned; paleas narrowly
elliptic, equalling or a little longer than the lemmas, the apex acute, awnless or short awned,
strongly 2-nerved, the nerves glabrous or strongly ciliate. Lodicules 2, cuneate, truncate, fleshy.
Stamens 3; anthers linear. Ovaries ovoid to ellipsoid, glabrous; styles 2, separate, short, terminal
stigmas plumose. Fruit a caryopsis, subterete, fusiform; hilum linear, nearly as long as grain;
embryo large, typically1/3 –3/4 the length of the grain. Base chromosome number 10. Type
species: B. racemosa Lag. [=B. curtipendula (Michx.) Torr.]. (Named for the Boutelou brothers) –
Grama-Grass.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 134
06/27/17
A genus of about 25 species occurring fromwestern Canada south to Argentina. The greatest
diversity is in Mexico. About 12 species occcur in the western and southwestern states, but only
two reach our area.
B. curtipendula, side-oats grama, 2n = 40, occcurs in the northern and western counties of
Arkansas (Smith). B. rigidiseta (Steud.) Hitchc., texas grama, 2n = 28, 35, 40, occurs in Little
River Co., southwestern Arkansas (Smith) and in Vernon parish, west-central Louisiana (Allen et
al.).
B. hirsuta Lag. occurs in Calcasieu Parish in southwestern Louisiana .
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membranous; blade linear, flat, glabrous or sparsely hispid. Staminate and carpellate inflorescences
dimorphic; staminate inflorescence a simple or compound raceme; branches if present erect,
flatened, scabrous on margins, bearing ??? spikelets; staminate spikelets 1-flowered, lance-ovate;
glumes unequal, lance-ovate, shorter than lemmas, 1-2 nerved, occasionally awned; paleas 2nerved, lanceolate; stamens 3; carpellate inflorescence of 1-2 short racemes, partly enclosed by the
upper leaf sheath, deciduous as a unit; carpellate spikelets 1-flowered (an awned rudiment present
above fertile floret), ovate, the glumes and lemmas strongly nerved; laterally compressed, narrowly
or broadly oblong; glumes 2, unequal, the first reduced, the second ovate, lanceolate; lemmas
narrowly oblong, 3-nerved, 4-lobed, with three awns between lobes; paleas narrowly elliptic,
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 136
06/27/17
equalling the lemmas, the apex acute, awnless, 2-nerved, the nerves broadly winged. Lodicules 2,
cuneate, truncate, fleshy. Ovaries narrowly ellipsoid, glabrous; styles 2, separate, short, terminal
stigmas plumose. Fruit a caryopsis, compressed, ellipsoid (enclosed by indurated glume); hilum
linear, nearly as long as grain; embryo large, typically 1/3 –3/4 the length of the grain. Base
chromosome number 10. Type species: O. stolonifera J.S. Presl, the only species included. (Named
for ---------------------) –
Attributed to florida by USDA PLANTS database. No material from U.S. at MO; doubtfully in
U.S. clayton & Renvoize say it occurs in Mexico and West Indies.
[The generic name was attributed to Rafinesque by Campbell, but the name does not appear in
Merrill's Index Rafinesquianum or any other index I have checked.]
23. Buchloë Engelman, Trans. Acad. Sci. St. Louis 1: 432. 1859, nom. cons.
Dioecious (occasionally monoecious) low mat-forming perennial of prairies [semidesert
scrub], generally in well drained soils. Rhizomes absent; stolons present. Stems single or
caespitose, erect, simple; leaves mostly cauline, few; sheaths rounded, glabrous, ligule
membranous, with a fringe of short white hairs; blade linear, flat, glabrous or sparsely hispid.
Staminate and carpellate inflorescences dimorphic; staminate inflorescence a simple or compound
raceme; branches if present erect, flatened, scabrous on margins, bearing 6-12 spikelets; staminate
spikelets 2-flowered, lance-ovate; glumes unequal, lance-ovate, shorter than lemmas, 1-2 nerved,
occasionally awned; palesa 2-nerved, lanceolate; stamens 3; carpellate inflorescence of two short
racemes, partly enclosed by the upper leaf sheath, deciduous as a bur-like unit; carpellate spikelets
1-flowered, broadly ovate, the glumes and lemmas strongly nerved; laterally compressed, narrowly
or broadly oblong; glumes 2, unequal, the first reduced, the second ovate, indurate, 3-5 lobed at the
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 137
06/27/17
apex; lemmas narrowly oblong, 3-nerved, 3-lobed; paleas narrowly elliptic, equalling the lemmas,
the apex acute, awnless, 2-nerved, the nerves scabrous. Lodicules 2, cuneate, truncate, fleshy.
Ovaries narrowly ellipsoid, glabrous; styles 2, separate, short, terminal stigmas plumose. Fruit a
caryopsis, compressed, ovoid or oblong (enclosed by indurated glume); hilum linear, nearly as
long as grain; embryo large, typically 1/3 –3/4 the length of the grain. Base chromosome
number 10. Type species: B. dactyloides (Nutt.) Engelm. (Sesleria dactyloides Nutt.), the
conserved type. (Named for ---------------------) – Buffalo-Grass.
A monotypic genus of the Great Plains of North America. Buchloe dactyloides, 2n = 20,
40, 56, ranges from Minnesota to Montana, south to Louisiana, Northern Mexico, Arizona and
nevada. In our area it is known from Arkansas (Fulton and Sharp counties; Smith) and Louisiana
(northwestern and southwestern parishes; Allen).
Buffalo grass is one of the most important grazing grasses of the short grass plains of the
U.S.
References
Under subfamily references see:
Ahring, R.M., & G.W. Todd. The bur enclosure of the caryopses of buffalograss as a factor
affecting germination. Agron. Jour. 69: 15-17. 1977.*
Anderson, K., & A.E. Aldous. Monoecious buffalo grass, Buchloe dactyloides. Jour.Am. Soc.
Agron. 29(8): 709-710. 1937. [From Kansas.]
Ferrara, M. Plant a low-maintenance lawn. Organic gardening. Feb 1992. v. 39 (2) :46-51.
lawns and turf. zoysia. festuca. buchloe dactyloides. pest resistance. endophytes
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 138
06/27/17
Frank, K.W. Gaussoin, R.E., Riordan, T.P., Miltner, E.D.Date of planting effects on seeded turftype buffalograss. Crop science 38: 1201-1213. 1998. buchloe dactyloides. sowing
date. establishment. weeds. heat sums. cultivars. sowing. ground cover. plant
density. genotype environment interaction. survival. winter kill. Geographic nebraska.
Geographic utah. Other
Abstract Establishing seeded buffalograss is often difficult
because improper timing can result in weed interference or stand failure. Research was
conducted to determine optimal buffalograss seeding dates and growing degree day
(GDD) requirements for 'Cody' and 'Texoka' buffalograss establishmemt. Both cultivars
were seeded monthly at 10 g burs m-2 from April through October in 1995 at the John
Seaton Anderson Turfgrass Research Facility near Mead, NE, and at the Greenville
Research Farm at Logan, UT. The soil types at the Nebraska and Utah sites were
Sharpsburg silty clay loam (fine, montmorillonitic, mesic Typic Argiudoll) and Millville
silt loam (coarse-silty, carbonatic, mesic Typic Rendoll), respectively. The percentages of
buffalograss and weed cover were rated visually on a 0 to 100% scale. Buffalograss
density was determined by tossing either a 0.1- or 0.2-m2 quadrat into the plot and
counting the number of plants in the quadrat. The experimental design was a randomized
incomplete block design with three replications. Location, buffalograss cultivar, and
planting date were treatment factors. The optimal seeding time for buffalograss at the
Nebraska site was late April through June and at the Utah site, late April through July,
August, September, and October planting dates at both sites did not result in successful
stands. The negative relationship between the percentage of weed cover and the
percentage of buffalograss cover implicated weed interference as a factor limiting
establishment. Our results indicate that to ensure survival and establishment of
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 139
06/27/17
buffalograss in regions where soils freeze during the winter, at least 1000 postplanting
GDD are required before cessation of growth in the fall.
Fry, J., Upham, W., & Leuthold, L. Seeding month and seed soaking affect buffalograss
establishment. HortScience 28: 902-903. 1993. buchloe dactyloides. seed treatment.
soaking. imbibition. sowing date. seedling emergence. seasonal variation.
environmental temperature. stand establishment. lawns and turf. Geographic kansas.
Other Abstract Field studies were conducted in consecutive years to evaluate the
influence of seeding month and seed soaking on buffalograss [Buchloe dactyloides (Nutt.)
Engelm.] establishment, as measured by percentage of coverage and seedling emergence.
In 1991, plots where 'Sharp's Improved' buffalograss burrs were seeded in May, June, or
July exhibited complete coverage 7 weeks after seeding (WAS). Between Apr. and Sept.
1992, mean high and low temperatures were approximately 3C cooler than in 1991, and
seeding in June or July resulted in >95% coverage 9 WAS. In the same year, seeding in
April or May required 12 to 13 weeks for complete coverage. Buffalograss seeded in
August exhibited 30% more seedlings 2 WAS compared with nonsoaked burrs and
increased coverage by up to 18% on selected rating dates 3 to 13 WAS. However,
complete coverage occurred only approximately 1 week sooner where soaked vs.
nonsoaked burrs were planted.
Gaitan-Gaitan, F., C.B. McKenney, &, D.L. Auld. Effect of burrs vs. caryopses and planting date
on establishment of seeded buffalograss in the Southern Great Plains. Crop science.
May/June 1998. v. 38 (3) :795-800.
buchloe dactyloides. establishment. planting
date. grass seeds. semiarid zones. seedling emergence. plant density. cultivars.
quality. Geographic texas Abstract Successful establishment of buffalograss [Buchole
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 140
06/27/17
dactyloides (Nutt.) Engelm.] by seed is the greatest limitation of this native species as a
turfgrass. There has been limited research to determine the impact of planting date and
use of untreated burrs vs. caryopses on establishment in the semiarid environments of the
Southern Great Plains. During the 1992, 1993, and 1994 growing seasons, buffalograss
was established by seed at 2-wk intervals from mid-May to mid-September at the Texas
Tech Univ. Plant Stress Lab. in Lubbock, TX. The test site was located on an Amarillo
fine, sandy soil (fine-loamy, mixed, thermic Aridic Paleustalf). Seedling emergence at 2
and 4 wk after planting, and stand density and turf quality at the end of the growing
season were used as indices of stand establishment. The cultivars Texoka and Comanche
differed only slightly in stand establishment and final turfgrass quality ratings in these
studies. Caryopses had an average of 34 and 28% higher seedling establishment than
burrs at 2 and 4 wk after planting, respectively. The improved establishment of caryopses
was significant in 1992 and 1994. Neither caryopses nor burrs produced good quality turf
by the end of the growing season when planted after mid-July (Day of the Year 193-197).
Based on these studies, both commercial turfgrass growers and homeowners in the
Southern Great Plains should be encouraged to plant buffalograss caryopses in the late
spring or early summer to ensure successful establishment of high quality turf.
Gernert, W.B. Variation in buffalo grass. Jour.Am. Soc. Agron. 29: 242-246. 1937.*
Harlan, J.R. The development of buffalo grass seed.. Jour.Am. Soc. Agron. 38:135-141. 1946
[Details of embryology; comparison with Triticum.]
Huff, D.R.
Wu, L. Sex expression in buffalograss under different environments. Crop science.
1987. v. 27 (4) :623-626. . buchloe dactyloides. populations. sex expression.
environmental factors. genotype environment interaction. light. nitrogen. Geographic
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 141
06/27/17
texas. Geographic colorado.
Huff, D.R. Sex ratios and inheritance of anther and stigma color in diploid buffalograss. Crop
science. Mar/Apr 1991. v. 31 (2) :328-332. . buchloe dactyloides. inheritance.
diploidy. sex ratio. sex determination. stigma. anthers. color. cytology. alleles.
recessive genes. chromosome pairing. segregation. Abstract The lack of genetic
information on buffalograss [Buchloe dactyloides (Nutt.) Engelm.] may be due to the
high ploidy level (2n = 40 and 2n = 60) of all reported U.S. germplasm. This study was
conducted to confirm the existence of a Mexican diploid (2n = 20) race and to initiate
genetic analysis in buffalograss by examining male-to-female sex ratios and inheritance
of anther and stigma color. Cytological investigation confirmed the existence of the
diploid race (2n = 20). Sex ratios were 1:1 at the population level but were significantly
(P < 0.01) male biased for particular parents. A polygenic sex determining mechanism
was hypothesized. Anther color was either red or blue and stigma color was either purple
or clear (absence of pigmentation), and were found to be readily observable and
consistently expressed. Blue anther color was controlled by a single recessive allele that
was assigned the symbol b. Clear stigmas were inherited as a bifactorial double recessive
trait.
Huff, D.R., Peakall, R. & Smouse, P.E. RAPD variation within and among natural populations of
outcrossing buffalograss [Buchloe dactyloides (Nutt.) Engelm.]. Theoretical and applied
genetics.. 86 (8) :927-934 1993. Abstract RAPD markers provide a powerful tool for the
investigation of genetic variation in natural and domesticated populations. Recent studies
of strain/cultivar identification have shown extensive RAPD divergence among, but little
variation within, inbred species or cultivars. In contrast, little is known about the pattern
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 142
06/27/17
and extent of RAPD variation in heterogeneous, outcrossing species. We describe the
population genetic variation of RAPD markers in natural, diploid sources of dioecious
buffalograss [Buchloe dactyloides (Nutt.) Engelm.]. Buffalograss is native to the semiarid regions of the Great Plains of North America, where it is important for rangeland
forage, soil conservation, and as turfgrass. Most sources of buffalograss germplasm are
polyploid; diploid populations are previously known only from semi-arid Central Mexico.
This is the first report of diploids from humid Gulf Coastal Texas. These two diploid
sources represent divergent adaptive ecotypes. Seven 10-mer primers produced 98
polymorphic banding sites. Based on the presence/ absence of bands, a genetic distance
matrix was calculated. The new Analysis of Molecular Variance (AMOVA) technique
was used to apportion the variation among individuals within populations, among
populations within adaptive regions, and among regions. There was considerable
variation within each of the four populations, and every individual was genetically
distinct. Even so, genetic divergence was found among local populations. Withinpopulation variation was larger and among-population variation smaller in Mexico than
in Texas. The largest observed genetic differences were those between the two regional
eco- types. These patterns of genetic variation Abstract were very different from those
reported for inbred species and provide important baseline data for cultivar identification
and continuing studies of the evolution of polyploid races in this species.
Johnson, P.G. Riordan, T.P. Other Arumuganathan, K. Ploidy level determinations in
buffalograss clones and populations. Crop science.. v. 38 (2) :478-482. 1998. buchloe
dactyloides. plant breeding. plant morphology. flow cytometry. clones. populations.
dna. chromosome number. aneuploidy. polyploidy. Other Abstract Buffalograss
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 143
06/27/17
[Buchloe dactyloides (Nutt.) Engelm.] is the subject of breeding programs for turfgrass
use because of its drought and heat tolerance, low growing habit, and low management
input. Three ploidy levels are known to exist (x = n = 10), but plants at each level are
nearly morphologically indistinguishable. In this paper, flow cytometry was evaluated as
a means of determining ploidy levels of 31 buffalograss clones and 34 seeded
populations. Six clones were analyzed for nuclear DNA base-pair composition by flow
cytometry. Based on DNA contents, buffalograss clones can be grouped into four sets
corresponding to chromosome number or ploidy level. Mean DNA contents were 0.93,
1.80, 2.15, and 2.63 pg DNA/nucleus for diploid, tetraploid, pentaploid, and hexaploids,
respectively. Chromosome numbers explained most of the DNA content variation as
shown by the strong linear relationship between DNA content and chromosome number.
Cultivar 315 is a pentaploid and is the first report of a pentaploid among buffalograss.
Seeded populations analyzed include those exclusively tetraploid, mixture of tetraploid
through hexaploid, and those exclusively hexaploid. Aneuploids between pentaploid and
hexaploid levels were noted. No differences were observed between ploidy levels or
genders with regard to base-pair composition.
Jones, M.D. Newell, L.C. Longevity of pollen and stigmas of grasses: buffalograss, Buchloe
dactyloides (Nutt.) engelm., and corn, Zea mays L. Jour.American Society of
Agronomy. Mar 1948. v. 40 (3) :195-204 Abstract Studies to determine the longevity of
pollen and stigmas of buffalograss and corn were conducted at Lincoln, Neb., during
1944 and 1945. Data on longevity of pollen were obtained from daily pollinations with
pollen stored under different conditions, while data on the duration of receptiveness of
stigmas were obtained from pollinations made with fresh pollen to stigmas of different
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 144
06/27/17
ages. Buffalograss pollen stored in the spike under a temperature of 40 degrees F and a
relative humidity of 90% effected fertilization for 7 days in both years. Free pollen stored
in a beaker under the same temperature and humidity remained viable for 6 days in 1944
and 8 days in 1945. These same treatments were applied to corn pollen in 1944 and 1945.
Corn pollen stored in the tassel effected fertilization for 9 days in 1944 and 8 days in
1945. No pollinations were made after 8 days in 1945 as a result of heavy rainfall. Corn
pollen stored in a beaker remained viable in part for 8 days. Under this treatment a very
good set of kernels was obtained the first 2 days with the viability decreasing rapidly
thereafter. Pollen refrigerated in the inflorescence with both buffalograss and corn was
more effective over a period of several days than pollen refrigerated in a beaker. Three
additional treatments involving temperatures of 77 degrees to 113 degrees and relative
humidities of 30 to 70% were ineffective in extending life of buffalograss or corn pollen.
Corn pollen stored in pollinating bags in direct sunlight under a maximum temperature of
96 degrees F was effective in fertilizing silks for only 3 hours, while pollen stored in the
shade of the plants under a maximum temperature of 86 degrees F remained viable for 30
hours. Abstract Cool temperature and high relative humidity appear to be important
factors in extending the life of grass pollen. Receptivity of buffalograss stigmas remained
fairly high for 13 days, after which time it decreased rapidly until no seed was produced
on the twenty-first day. Duration of receptiveness in corn silks extended over a period of
24 days. Two- to 12-day-old-silks set seed rather uniformly, averaging 60%. Seed setting
decreased from 41% on silks 14 days old to only 2% on silks 24 days old. The duration of
receptivity of stigmas was 5 days more in corn than in buffalograss, The peak of
receptivity occurred on the fifth day in buffalograss and on the eighth day in corn. It was
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 145
06/27/17
found that with both buffalograss and corn the stigmas continued to grow in length until
pollen was applied. Stigmas were observed to grow to lengths of 1 inch in buffalograss
and 12 inches in corn.
Leuthold, L.D. Buffalograss lawns. - Cooperative Extension Service, Kansas State University,
Manhattan.. (658) 4 pp. 1982. buchloe dactyloides. lawns and turf. site selection. soil
types. irrigation. mowing. weed control. seeding. pest control.
Lowe, A.E. Viability of buffalo grass seeds found in the walls of a sod house. Jour. Am. Soc.
Agron. Nov 1940. v. 32 (11) :891-893. buchloe dactyloides. seed germination. grass
seeds. viability. kansas.
Peakall, R. Smouse, P.E. Huff, D.R. Evolutionary implications of allozyme and RAPD
variation in diploid populations of dioecious buffalograss Buchloe dactyloides.
Molecular ecology. Apr 1995. v. 4 (2) :135-147.
buchloe dactyloides. genetic
markers. alloenzymes. loci. alleles. genetic variation. genetic polymorphism.
genetic variance. geographical variation. population genetics. OtherSubject random
amplified polymorphic dna OtherSubject genetic diversity OtherSubject analysis of
molecular variance Geographic texas. Geographic mexico.
Quinn, J.A. Evolution of dioecy in Buchloe dactyloides (Gramineae): tests for sex-specific
vegetative characters, ecological differences, and sexual niche-partitioning. American
journal of botany. Apr 1991. v. 78 (4) :481-488. . buchloe dactyloides. dioecy. sex
expression. plant morphology. leaves. sex differences. plant ecology. growth.
plant competition. evolution. OtherSubject sexual dimorphism Geographic oklahoma.
Geographic kansas. Abstract Buchloe dactyloides is a perennial dioecious grass in which
male and female inflorescences are so strikingly dimorphic that they were originally
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 146
06/27/17
assigned to different genera. The objective of this paper is to present the results of tests
for sex-specific vegetative characters, ecological differences, and sexual nichepartitioning, combining them with prior information on the reproductive biology of
Buchloe for an evaluation of the key factors leading to the evolution of dioecy and sexual
dimorphism. Field and greenhouse data were collected from Oklahoma and Kansas
populations on vegetative characters, allocation to reproduction, and relative growth and
competitive success along resource gradients. Except for greater susceptibility to leaf rust
by males, there were no significant differences between males and females in vegetative
characters, total biomass, or reproductive effort. Field studies of spatial distributions of
males and females failed to show any relation to soil, topograph, or soil moisture. In a 45month greenhouse experiment starting at the seedling stage, the relative growth and
competitive success of randomly paired individuals showed no evidence for differential
competitive success or for niche-partitioning of males and females. The "outcrossing
advantage" and subsequent sexual specialization of the female inflorescence appear to be
the major factors underlying this dimorphic system.
Quinn, J.A. Natural expansion of Buchloë dactyloides at a disturbed site in New Jersey and its
implications for turf and conservation uses. Jour. Torrey Bot. Soc. 125: 319-323.
1998. buchloe dactyloides. disturbed land. erosion control. establishment. growth.
Geographic new jersey.
Quinn, J.A. Relationship between synaptospermy and dioecy in the life history strategies of
Buchloe dactyloides (Gramineae). American journal of botany. Aug 1987. v. 74 (8)
:1167-1172. ill. . buchloe dactyloides. life history. dioecy. seed dispersal. seed
germination. soil moisture. plant density. plant establishment. survival.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 147
06/27/17
Quinn, J.A. Engel, J.L. Life-history strategies and sex ratios for a cultivar and a wild population
of Buchloe dactyloides (Gramineae). American journal of botany. June 1986. v. 73 (6)
:874-881. ill. . buchloe dactyloides. cultivars. wild plants. life history. sex ratio.
sex expression. reproduction. Geographic oklahoma. Geographic kansas. Geographic
grasslands.
Ramsey, G.W., & C.R. Brooks. Buchloë dactyloides (Nutt.) Engelm. (Poaceae) new to Virginia in
Bedford County. Virg. Jour. Sci. 38: 249-252. 1987.
Remington, K.K.
Bonham, C.D. Other Reich, R.M. Blue grama-buffalograss responses to
grazing: a Weibull distribution. Journal of range management. May 1992. v. 45 (3) :272276. . bouteloua gracilis. buchloe dactyloides. mixed pastures. biomass. prediction.
cattle. grazing effects. plant height. frequency distribution. mathematical models.
grazing intensity. spatial variation. temporal variation. Other Abstract
Characterization of standing herbaceous biomass on rangeland is complicated by both
temporal and spatial variability that results from patchiness in vegetation. These patches
often cause nonuniform levels of grazing by livestock. Currently accepted methods for
estimation of forage, and its utilization, assume a normal distribution. This assumption
may not be appropriate if the frequency distribution of amount of biomass becomes
skewed as grazing occurs. We evaluated the 3 parameter Weibull distribution as an
alternative to the normal distribution in modeling the frequency distributions of plant
height and biomass as a function of grazing intensity over time in a shortgrass steppe.
Weibull distributions, estimated by probability weighted moments, fit all observed plant
height and biomass data distributions at the alpha = 0.05 level of significance. In contrast,
the normal distribution fit only 25% of the data sets.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 148
06/27/17
Riordan, T. Buffalograss. Grounds maintenance. Feb 1991. v. 26 (2) :12-14. ill., maps.
buchloe dactyloides. cultural methods.
Schaffner, J.H. The dioecious nature of buffalo-grass. Bull. Torrey bot Club 47: 119-124. 1920*
Shaw, R.B. Bern, C.M. Other Winkler, G.L Sex ratios of Buchloe dactyloides (Nutt.) Engelm.
along catenas on the shortgrass steppe. Bot. Gaz.. Mar 1987. v. 148 (1) :85-89. .
buchloe dactyloides. sex ratio. catenas. steppes. dioecy. plant colonization.
Wenger, L.E. Inflorescence variations in buffalo grass, Buchloe dactyloides. Jour.American
Society of Agronomy. Apr 1940. v. 32 (4) :274-277. buchloe dactyloides.
inflorescences. variation.
Whitcomb, R.F. Other Hicks, A.L. Other Blocker, H.D. Other Lynn, D.E. Biogeography of
leafhopper specalists of the shortgrass prairie: evidence for the roles of phenology and
phylogeny in determination of biological diversity. American entomologist.40: 19-35.
1994. buchloe dactyloides. bouteloua gracilis.
Wu, L. Allelopathic effects of phenolic acids detected in buffalograss (Buchloe dactyloides)
clippings on growth of annual bluegrass (Poa annua) and buffalograss seedlings.
Environmental and experimental botany. Apr 1998. v. 39 (2) :159-167.
buchloe
dactyloides. poa annua. seedlings. allelopathins. phenolic acids. crop residues.
plant extracts. p-coumaric acid. ferulic acid. vanillic acid. plant composition.
growth. varieties. genetic variation. seed germination. roots. inhibition.
establishment. shoots. length. OtherSubject gentisic acid OtherSubject homoveratric
acid OtherSubject p-hydroxybenzoic acid
Wu, L., & H. Lin. Salt tolerance and salt uptake in diploid and polyploid buffalograsses (Buchloe
dactyloides). Jour. Pl. Nutr. 17: 1905-1928. 1994..
variation. roots. Abstract Seed
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 149
06/27/17
samples of diploid, tetraploid, and hexaploid buffalograss (Buchloe dactyloides) were
collected from locations over a geographical latitudinal gradient from San Luis Potosí,
Mexico to Lincoln, Nebraska in the United States. Seed samples and samples of
vegetatively propagated clones were tested for salt tolerance and salt uptake. Under
nutrient solution culture, young shoots separated from the established buffalograss clones
were found to be more tolerant to salt than the seedlings. Significant difference was found
in the percentage of seedling survivorship among the populations. Substantial genetic
variation of salt tolerance was detected among the vegetatively propagated clones within
buffalograss populations. Over all, the buffalograss can be considered to be a moderately
salt sensitive species. Its wealth of genetic variation of salt tolerance represents a
potential for rapid salt tolerance selection response. A salt exclusion mechanism was
found in the tolerance mechanism of the buffalograss. A greater sodium (Na)
concentration was found in the root tissue than in the shoot tissue, suggesting a
preferential exclusion of Na taken up by the shoots. The negative correlations between the
plant tissue potassium (K) concentrations and Na/K ratios indicate a partial substitution
of K by Na, and there was a less substitution of K by the more salt tolerant plants than the
less tolerant plants. Differential susceptibility to calcium (Ca) disorders at high Na/Ca
levels was detected between the salt tolerant and salt sensitive buffalograss genotypes.
Wu, L., Z.Z. Huang, & R.G. Burau. Selenium accumulation and selenium-salt cotolerance in five
grass species. Crop sci. 1988. 28: 517-522. [buchloe dactyloides cynodon dactylon]
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 150
06/27/17
27a. Cathestecum s. U.S.
27b. Aegopogon same
24. Tragus [Haller, Hist. Stirp. Helv. 2: 203. 1768] Scopoli, Intr. 73. 1777, nom. cons.
Annuals [perennials] of disturbed open places, typically in temporarily wet soils. Rhizomes
absent. Stems erect, simple or branched near the base; leaves cauline and basal, few; sheath shorter
than the internode, glabrous, sparsely pubescent at the throat; ligule membranous, ciliate-margined;
blade linear, flat, or plicate. Inflorescence a false raceme, with spaced, spreading spikelets in
clusters of 2-5; short secondary branches sometimes present. Spikelets pedicellate, strongly
laterally compressed, 2-4 flowered; glumes 2, unequal, the first only about ¼ as long as the second,
sometimes obsolete; second glume lanceolate, acute to mucronate, about as long as the spikelet, 3nerved, coriaceous, covered with stout, uncinate hairs especially along the midvein; rachilla
disarticulating at base of each spikelet; flowers bisexual, the uppermost 2 or 3 sterile or
rudimentary; lemmas shorter than second glume, narrowly elliptic, acute, 1(-3) nerved, keeled,
pubescent on the nerves, or glabrous; paleas elliptic, a little shorter than the lemmas, 2keeled, the keels often ciliolate. Lodicules 2, cuneate, truncate, fleshy. Stamens 3; anthers
ellipsoid. Ovaries ovoid to ellipsoid, glabrous; styles 2, separate, short, terminal stigmas
plumose. Fruit a caryopsis, trigonous, ellipsoid; hilum linear, nearly as long as grain; embryo
large, 1/2 the length of the grain. Base chromosome number ? Type species: T. racemosus (L.)
All.. (Nazia Adans., Lappago Schreb.) (Name from ----.) -- Burgrass
A small genus of cool looking grasses, thoroughly unmistakable. Seven species are
generally recognized. Tragus racemosus Willd. [or (L.) Desf.???] is naturalized in the U.S., but
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 151
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records from the Southeast are very few, including only North Carolina (Hitchcock & Chase) and
Mississippi (Starkville, "introduced," Tracy, 8-6-[18]91, MO).
References
Anton, A.M. The genus Tragus (Gramineae). Kew Bull. 36: 55-61. 1981
25. Zoysia Willdenow., Ges. Nat. Freunde Berlin Neue Schr. or Ser.? 3: 440. 1801, nom. cons.
Low mat or sward forming perennials of lawns, roadsides, pastures [savannas]. Rhizomes
horizontal, extensive, clothed with lanceolate scales; stolons present, arching. Stems oblique to
erect, cespitose, unbranched; leaves cauline, few; sheath glabrous or ciliate; ligule a ciliate
membrane; blade linear, flat or plicate. Inflorescence a spike-like raceme [a single spikelet in Z.
minima], the individual pedicellate spikelets appressed. Spikelets laterally compressed, 1-flowered;
glumes 1 (the first absent), ovate-lanceolate, acute [awned], 1-nerved; rachilla disarticulating below
the glume (the pedicel persistent on stem); flowers bisexual; lemmas ovate-lanceolate, [1-] 3nerved, acute [emarginate], smooth, scabrous along the midvein; paleas elliptic, a little shorter than
the lemmas, 2-nerved [obsolete]. Lodicules 2, cuneate, truncate, fleshy. Stamens 3; anthers
ellipsoid. Ovaries ovoid to ellipsoid, glabrous; styles 2, separate, short, terminal stigmas plumose.
Fruit a caryopsis, laterally compressed, ellipsoid, hilum linear, nearly as long as grain; embryo
large, typically1/3 –3/4 the length of the grain. Base chromosome number 9. [Matrella Pers.;
Brousemichea Bal.; Osterdamia Necker ex Kuntze.] (Named for Karl von Zois, ) Type species: Z.
pungens Willd. [=Z. matrella (L.) Merr.] – Zoysia.
A genus of about 10 species, native to southern and eastern Asia, the Mascarene Islands of
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 152
06/27/17
the Indian Ocean, and Australia, and New Zealand. Several species have been introduced as lawn
grasses in tropical and warm-temperate regions worldwide (Roshevits). Several species have been
used for stabiling coastal sands. In the United States, Z. matrella (L.) Merr. is found from …..
MO: St. Louis, "cultivated," Davidse 30832 (MO)
NC: Orange Co., Chapel Hill, Boufford 15962 (MO)
IL: Jackson Co., "not believed to be planted," 31 May 1979, Voigt (MO).
Z. tenuifolia Willd. ex Trin. said to introduced in Florida and California (Bor)
Z. japonica Steud., native to Japan, used as lawn grass in U.S.
References
Under subfamily references see Bor; Clayton & Renvoize; Hitchcock & Chase.
Akiyoshi, M., N. Endo, M. Yaneshita, R. Nagasawa, K. Yoshida, & T. Yagura. Sea water tolerance
in plants. (4). Zoysiagrass in japan.. Journal 30: 267-272. 1997. have reprint
Fukuoka, H. Breeding of Zoysia spp. Jour. Jap. Soc. Turfgrass Sci. 17: 183-190. 1989
Hondo, H. Morphology and structure of zoysiagrass. Pp. 26-43 in N. Nakamura, ed. Science and
management for golf course. Tokyo. 1993.*
Ikeda, H. & M. Hoshino. Studies on the ecotypes of Zoysia japonica collected from all over Japan.
Jour. Japan. Turfgrass Res. Assoc. 7(1): 27-34. 1978.
Inokuma, C., K. Sugiura, N. Imaizumi, & C. Cho. Transgenic Japanese lawngrass (Zoysia japonica
Steud.) plants regenerated from protoplasts. Pl. Cell Rep. 17: 334-338. 1998.
Kitamuraa, F. Studies on the salt tolerance of lawn grasses. 4. On the salt tolerance of Japanese
lawn grasses (Zoysia grasses). Jour. Jap. Inst. Landscape Architects 33: 28-33. 1970.
-----. Studies on the horticultural classification and development of Japanese lawn grasses. Bull.
Kemigawa Arboretum, Fac. Agr. Univ. Tokyo 3: 1-60. 1970.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 153
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Lee, G.J., Y.Y. Kweon, & K.K. Sun. comparative salt tolerance study in zoysia grasses. II.
Interspecific comparison among eight zoysia grasses (Zoysia spp.). Jour. Kor. Soc. Hort.
Sci. 35 (2): 178-185. 1994.*
Shoji, S. Species ecology on Zoysia grass. Jour. Jap. Turfgrass Res. Assoc. 12 (2): 105-110. 1983.
Yamada, T. & H. Fukuoka. Variations in peroxidase isozyme of Japanese lawn grass (Zoysia
japonica Steud.) populations in Japan. Japan. Jour. Breed. 34: 431-438. 1984.*
Yaneshita, M., M. Akiyoshi, N. Endo, & S. Kaneko. Identification of isozyme loci in Zoysia. Jour.
Jap. Soc. Turfgrass Sci. 26: 114, 115. 1997.
Yaneshita, M., R. Nagasawa, R. Engelke, & T. Sasakuma. Genetic variation and interspecific
hybridization among natural populations of zoysiagrasses detected by RFLP analyses of
chloroplast and nuclear DNA. Genes Genet. Syst. 72: 173-179. 1997.
-----, -----, S. Kaneko, Y. Ogihara, & T. Sasakuma. Genetic charactrization of zoysiagrasses by
RFLP analysis of nuclear DNA. Intern. Turfgrass Soc. Res. Jour. 7: 786-792. 1993.*
-----, T. Ohmura, T. Sasakuma, & Y. Ogihara. Phylogenetic relationships of turfgrasses as revealed
by the restriction fragment analysis of chloroplast DNA. Theor. Appl. Genet. 87: 129-135.
1993.
Gordon C.Tucker, Chloridoideae of the S.E. U.S., manuscript, p. 154
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