| P.W. Michael |
Plants annual or perennial; with or
without rhizomes. Culms 10-460 cm, prostrate, decumbent or erect, distal
portions sometimes floating, sometimes rooting at the lower nodes; nodes
usually glabrous; internodes hollow or solid. Sheaths open, compressed;
auricles absent; ligules usually absent but, if present, of hairs;
blades linear to linear-lanceolate, usually more than 10 times longer
than wide, flat, with a prominent midrib. Inflorescences terminal, panicles
of simple or compound spikelike branches attached to elongate rachises, axes
not terminating in a bristle, spikelets subsessile, densely packed on the angular
branches; disarticulation below the glumes (cultivated taxa not or tardily
disarticulating). Spikelets plano-convex, with 2(3) florets; lower
florets sterile or staminate; upper florets bisexual, dorsally compressed.
Glumes membranous; lower glumes usually 1/4-2/5 as long as the
spikelets (varying to more than 1/2 as long), unawned to minutely awn-tipped;
upper glumes unawned or shortly awned; lower lemmas similar to
the upper glumes in length and texture, unawned or awned, awns to 60 mm; lower
paleas vestigial to well-developed; upper lemmas coriaceous, dorsally
rounded, mostly smooth, apices short or elongate, firm or membranous, unawned;
upper paleas free from the lemmas at the apices; lodicules absent
or minute; anthers 3. Caryopses ellipsoid, broadly ovoid or spheroid;
embryos usually 0.7-0.9 times as long as the caryopses. x = 9.
Name from the Greek echinos, hedgehog, and chloa, grass, in reference
to the bristly or often awned spikelets.
Echinochloa is a tropical to warm-temperate genus of 40-50 species that
are usually associated with wet or damp places. Many of the species are difficult
to distinguish because they tend to intergrade. Some of the characters traditionally
used for distinguishing taxa, e.g., awn length, are affected by the amount of
moisture available; others reflect selection by cultivation, e.g., non-disarticulation
in grain taxa, mimicry of rice as weeds of rice fields. There are 13 species
in the Flora region: five native and one possibly native, four established,
two grown as commercial crops, and one in research.
In North America, the most abundant species appears to be the
introduced, weedy Echinochloa crus-galli,
which closely resembles the native E. muricata.
The confusion between the two species has caused them to be treated as the same
species. This confusion is likely reflected in the mapping of both Echinochloa
crus-galli and E. muricata. Echinochloa
frumentacea and E. esculenta are grown for grain in India and in China and Japan, respectively, but not
in North America. Echinochloa oryzoides and E. oryzicola are weeds
whose success and distribution reflects their adaptation to the periodic inundations
of commercial rice fields.
Cytogenetic data suggest that E. frumentacea and E.
esculenta are domesticated derivatives of E.
colona and E. crus-galli, respectively (Yabuno 1962) and that E. oryzoides is
very closely related to E. crus-galli (Yabuno 1984). Molecular work (Yamaguchi et al. (2005) are consisten with the cytogenetic daa with respect to the origin of the two domesticated taxa and places E. oryzoides in the unresolved clade that includes E. crus-galli and E. esculenta. Yabuno
(1966) suggested that E. crus-galli is an allohexaploid
produced by natural hybridization between the tetraploid E.
oryzicola with a not-yet-discovered diploid
species of Echinochloa and subsequent chromosome doubling. Studies
using seed protein electrophoresis and isozyme analyses (Kim et al.
1989; González-Andrés
et al. 1996; Asíns et al. 1999), and molecular studies involving
RAPD markers and DNA sequences (Hilu 1994; Roy et al. 2000) or PCR-RFLP
techniques (Yasuda et al. 2001), will help in clarifying the phylogenetic
problems in Echinochloa,
providing that proper attention is paid to the morphological characterization
of the plant materials used and voucher specimens are made.
The map for Echinochloa polystachya has been amended (see comment), the correct varietal epithet for Echinochloa crus-pavonis should be var. macera [for additional comments on the correct varietal epithet, click here], and a reference supporting use of 'colona' for the epithet of E. colona added.
SELECTED REFERENCES Asíns, M.J., J.L. Carretero, A. Del Busto, E.A. Carbonell, and D. Gomez de Barreda. 1999. Morphologic and isozyme variation in barnyard grass (Echinochloa) weed species. Weed Technol. 13:209-215; Barrett, S.C.H. and D.E. Seaman. 1980. The weed flora of California rice fields. Aquatic Bot. 9:351-376; Carretero, J.L. 1981. El género Echinochloa Beauv. en el sudeste de Europa. Anales Jard. Bot. Madrid 38:91-108; Fishbein, M.1995. Noteworthy collections: Arizona. Madroño 42:83; González-Andrés, F., J.M. Pita, and J.M. Ortiz. Caryopsis isoenzymes of Echinocloa [sic] weed species as an aid for taxonomic discrimination. J. Hort. Sci. 71:187-193; Gould, F.W., M.A. Ali, and D.E. Fairbrothers. 1972. A revision of Echinochloa in the United States. Amer. Midl. Naturalist 87:36-59; Hilu, K.W. 1994. Evidence from RAPD markers in the evolution of Echinochloa millets (Poaceae). Pl. Syst. Evol. 189:247-257; Hitchcock, A.S. 1913. Mexican grasses in the United States National Herbarium. Contr. U.S. Natl. Herb. 173:181-389; Jauzein, P. 1993. Le genre Echinochloa en Camargue. Monde Pl. 88, no. 446:1-5; Kim, K.V., J.H. Kim, and I.J. Lee. 1989. Biochemical identification of Echinochloa species collected in Korea. Proc. Conf. Asian-Pacific Weed Sci. Soc. 12th [Proc. II]:519-531; Kobayashi, H. and S. Sakamoto. 1990. Weed-crop complexes in cereal cultivation. Pp. 67-80 inS. Kawano (ed.). Biological Approaches and Evolutionary Trends in Plants. Academic Press, London, England and San Diego, California, U.S.A. 417 pp.; Michael, P.W. 1983. Taxonomy and distribution of Echinochloa species with a special reference to their occurrence as weeds of rice. Pp. 291-306 in International Rice Research Institute. Proc. Conf. Weed Control in Rice (31 Aug.-4 Sept. 1981). International Rice Research Institute, Los Baños, Laguna, Philippines. 422 pp.; Michael, P.W. 2001. The taxonomy and distribution of Echinochloa species (barnyard grasses) in the Asian-Pacific region, with a review of pertinent biological studies. Proc. Conf. Asian-Pacific Weed Sci. Soc. 18th [Proc. I]:57-66; Nicolson, D.H. 1986. Species epithets and gender information. Taxon 35:323-328; Roy, S., J.-P. Simon, and F.-J. Lapointe. 2000. Determination of the origin of the cold-adapted populations of barnyard grass (Echinochloa crus-galli) in eastern North America: A total-evidence approach using RAPD DNA and DNA sequences. Canad. J. Bot. 78:1505-1513; Vickery, J.W. 1975. Echinochloa Beauv. Pp. 189-211 in J.W. Vickery. Flora of New South Wales, No. 19, Gramineae, Part 2 (ed. M.D. Tindale). New South Wales Department of Agriculture, Contributions from the New South Wales National Herbarium Flora Series. National Herbarium of New South Wales, Sydney, New South Wales, Australia. 181 pp.; Wunderlin, R.P. 1988. Guide to the Vascular Plants of Florida. University Press of Florida, Gainesville, Florida, U.S.A. 806 pp.; Yabuno, T. 1962. Cytotaxonomic studies on the two cultivated species and the wild relatives in the genus Echinochloa. Cytologia 27:296-305; Yabuno, T. 1966. Biosystematic study of the genus Echinochloa. J. Jap. Bot. 19:277-323; Yabuno, T. 1984. A biosystematic study on Echinochloa oryzoides (Ard.) Fritsch. Cytologia 49:673-678; Yabuno, T. and H. Yamaguchi. 1996. Hie no hakubutsugaku [A natural history of Echinochloa]. Dow Chemicals and Dow Elanco, Tokyo, Japan. 196 pp. [In Japanese]; Yamaguchi, H. A. Utano, K. Yasuda, A. Yano, and A. Soejima. 2005. A molecular phylogeny of wild and cultivated Ehinochloa in East Asia inferred from non-coding region sequences of trnT-L-F. Weed Biology and Management 5:210-218..Yasuda, K., A. Yano, Y. Nakayama, and H. Yamaguchi. 2001. Identification of Echinochloa oryzicola and E. crus-galli using PCR-RFLP technique. [Abstract]. J. Weed Sci. Technol. 46, Suppl.:204-205. [Text in Japanese, title in Japanese and English].NOTE: In this treatment, measurements of the spikelets do not include the awns. The color of the caryopses is based on fully ripe caryopses. Chromosome numbers cited in this treatment are only those which are confidently believed to refer to the species described here.
1 |
Ligules
of stiff hairs present on the lower leaves; lower florets staminate; plants
perennial (2) |
Ligules almost always absent from all leaves,
the ligule region sometimes pubescent; lower florets sterile or staminate;
plants usually annual, sometimes short-lived perennials (3) |
|
Plants without scaly
rhizomes, sometimes rooting at the lower nodes; lower lemmas usually awned,
sometimes merely apiculate; known outside of experimental plantings ..... 1.
E. polystachya |
|
Plants with short, scaly rhizomes; lower
lemmas unawned, sometimes long-cuspidate; in the Flora region,
known only from experimental plantings ..... 2. E.
pyramidalis |
|
Lower lemmas
usually unawned; spikelets, particularly those near the base of the panicles,
not disarticulating at maturity; upper lemmas wider and longer than the
upper glumes at maturity and, hence, exposed at maturity (4) |
|
Lower lemmas often awned; spikelets disarticulating
at maturity; upper lemmas not or scarcely exceeding the upper glumes in
length and width at maturity (5) |
|
Spikelets
always green and pale at maturity, their apices usually obtuse, varying
to acute; rachis nodes not or only sparsely hispid with papillose-based
hairs; caryopses whitish ..... 9. E.
frumentacea |
|
Spikelets purplish to blackish-brown
at maturity, their apices obtuse to shortly acute; rachis nodes densely
hispid with papillose-based hairs; caryopses brownish ..... 11. E.
esculenta |
|
Plants essentially
obligate weeds of rice, growing in the fields; culms erect, densely tufted;
spikelets 3.7-7 mm long; plants resembling rice in their vegetative growth
(6) |
|
Plants not obligate weeds of rice, found
in summer crops and wet places, and often in rice fields; culms sprawling,
decumbent or erect; spikelets 2-5 mm long; plants occasionally resembling
rice vegetatively but, if so, the spikelets less than 3 mm long (7) |
|
Panicles
horizontal or drooping at maturity; spikelets broadly ovate to ovate;
lower lemmas usually awned; caryopses 1.9-3 mm long, the embryos 70-85%
as long as the caryopses ..... 12. E.
oryzoides |
|
Panicles erect to slightly drooping;
spikelets ovate-elliptical; lower lemmas awned or not; caryopses 1.7-2.6
mm long, the embryos 89-98% as long as the caryopses ..... 13.
E. oryzicola |
|
Lower florets
staminate; anthers of the upper florets 1.2-1.7 mm long ..... 3.
E. paludigena |
|
Lower florets sterile; anthers of the
upper florets 0.5-1.2 mm long (8) |
|
Panicle
branches 0.7-2(4) cm long, without secondary branches; spikelets 2-3 mm
long, unawned ..... 8. E. colona |
|
Panicle branches 1-14 cm long, usually
rebranched, the secondary branches often short and inconspicuous; spikelets
2.5-5 mm long, awned or unawned (9) |
|
Upper lemmas
broadly ovate to elliptical, if elliptical, each with a line of minute
(need 25× magnification) hairs across the base of the early-withering
tips (10) |
|
Upper lemmas narrowly ovate to elliptical,
never with a line of minute hairs across the base of the early-withering,
membranous tips (11) |
|
Upper lemmas with rounded
or broadly acute coriaceous apices that pass abruptly into a membranous
tip, a line of minute hairs present at the base of the tip ..... 10.
E. crus-galli |
|
Upper lemmas with acute
or acuminate coriaceous apices that extend into the membranous tip, without
hairs at the base of the tip ..... 4. E.
muricata |
|
Spikelets
2.5-3.4 mm long; lower lemmas unawned or with awns 3-10(15) mm long, curved ..... 6.
E. crus-pavonis |
|
Spikelets 3-5 mm long; lower lemmas usually
with awns 8-25 mm long, typically straight (12) |
|
Blades
10-35(60) mm wide; sheaths usually hispid and the margins ciliate with
prominent papillose-based hairs, sometimes the sheaths only papillose;
lower lemmas awned, the awns 8-25(60) mm long; common in the eastern portion
of the Flora region ..... 5. E.
walteri |
|
Blades 5-10 mm wide; sheaths glabrous
or with papillose-based hairs; lower lemmas unawned or awned, the awns
8-16(50) mm long; in the Flora region, known only from southern
Arizona ..... 7. E. oplismenoides |
1. Echinochloa polystachya (Kunth) Hitchc.
Creeping River Grass
Plants perennial; not rhizomatous. Culms 1-2 m tall, to 1 cm thick,
erect or decumbent and rooting at the lower nodes, upper portion sometimes floating
distally; nodes glabrous or antrorsely villous. Sheaths mostly
glabrous, minutely puberulent, or hispid, hairs papillose-based, throat hispid;
ligules present on the lower leaves, 1-5 mm, of stiff hairs; blades
15-70 cm long, 5-13 mm wide, glabrous. Panicles 13-45 cm, erect, rachis
nodes hispid, hairs 3-6.5 mm, papillose-based, internodes scabrous; primary
branches 4-10 cm, subverticillate, ascending, nodes hispid, hairs 2.5-4
mm, papillose-based, internodes scabrous; secondary branches short, spikelets
subsessile, in clusters. Spikelets 4-7 mm, hispid, hairs appressed, disarticulating
at maturity. Lower glumes at least 1/2 as long as the spikelets; lower
florets staminate; lower lemmas apiculate or awned, awns to 18 mm;
lower paleas subequal to the lower lemmas, often purple; anthers of
lower florets 1.5-3.6 mm, orange; upper lemmas 2.5-5 mm, elliptic
or narrowly ovate, apices obtuse, with a membranous, soon-withering tip; anthers
of upper florets shorter than those of the lower florets. Caryopses
to 3 mm. 2n = 54.
Echinochloa polystachya grows in coastal marshes, often in standing water,
from Texas to Louisiana, and south through Mexico and the Caribbean islands
to Argentina. Echinochloa polystachya var. polystachya
has glabrous culms and leaf sheaths; Echinochloa polystachya var.
spectabilis (Nees
ex Trin.) Mart. Crov. has swollen, pubescent cauline nodes and pubescent
leaf sheaths.
December 2003: Prof. Daniel
B. Ward wrote to Barkworth stating that the record of Echinochloa polystachya
from Florida is based on a cultivated specimen. The map on the Web site and
the above comment have been amended to reflect his correction.
2. Echinochloa pyramidalis (Lam.) Hitchc. & Chase
Antelope Grass
Plants perennial; with short, scaly rhizomes. Culms 1-4.6 m tall,
to 2 cm thick, geniculate or long-prostrate and rooting at the lower nodes, often
floating distally; lower and upper nodes glabrous. Sheaths
mostly glabrous, but usually ciliate at the throat; ligules present on
the lower leaves, 1-5 mm, of stiff hairs, reduced or absent on the upper leaves;
blades 8-75 cm long, 5-30 mm wide. Panicles 15-40 cm, nodes and
internodes scabrous; primary branches 2-7.5 cm, solitary to fascicled,
erect or ascending, simple or compound, nodes and internodes glabrous or hispid,
hairs to 4 mm, papillose-based. Spikelets 2.5-4 mm long, 1-1.8 mm wide,
disarticulating at maturity, finely pubescent or glabrous, greenish to purple
at maturity. Lower florets staminate; lower lemmas unawned, acute
to acuminate or long cuspidate; anthers of lower florets 1-1.5 mm; upper
lemmas apiculate to long cuspidate. Caryopses about 2 mm. 2n
= 54, 72.
Echinochloa pyramidalis is native to Africa, where it is used both as a
cereal and a pasture grass. It has been grown experimentally in Gainesville, Florida,
but it is not established in North America.
3. Echinochloa paludigena Wiegand
Florida Barnyard Grass
Plants annual. Culms to 150 cm, erect. Sheaths glabrous;
ligules absent; blades 15-60 cm long, 8-20 mm wide, scabrous adaxially.
Panicles 8-40 cm, erect to slightly drooping, rachis nodes hispid, hairs
papillose-based; primary branches 2-19 cm, erect to spreading, often widely
spaced, longer branches with secondary branching. Spikelets 3.3-4.5 mm
long, 2.4-2.6 mm wide, disarticulating at maturity, greenish or purplish, scabrous
and hispid, hairs to 1 mm, often papillose-based. Upper glumes about as
long as the spikelets; lower florets staminate; lower lemmas usually
awned, awns 1-15 mm, purplish; lower paleas well-developed; upper lemmas
broadly ovate, narrowing abruptly to the acute or acuminate apices; anthers
of upper florets 1.2-1.7 mm. Caryopses 1.5-1.8 mm. 2n = unknown.
Echinochloa paludigena is native to swamps, riverbanks, and other wet habitats.
Reports from Texas and Louisiana appear to be based on misidentifications; Wunderlin
(1988) considers E. paludigena as a Florida endemic.
4. Echinochloa muricata (P. Beauv.) Fernald
American Barnyard Grass
Plants annual. Culms 80-160 cm, erect or spreading,
sometimes rooting at the lowest nodes, often developing short axillary flowering
shoots at most upper nodes when mature; lower nodes glabrous or puberulent;
upper nodes glabrous. Sheaths glabrous; ligules absent;
blades 1-27 cm long, 0.8-30 mm wide. Panicles of primary culms
7-35 cm, rachises and branches glabrous or hispid, hairs to 3 mm, papillose-based;
primary branches 2-8 cm, usually spreading and rather distant, often
with secondary branches. Spikelets 2.5-5 mm, disarticulating at maturity,
usually purple or streaked with purple, usually hispid, hairs papillose-based.
Upper glumes about as long as the spikelets; lower florets sterile;
lower lemmas unawned or awned, awns to 16 mm; lower paleas well-developed;
upper lemmas broadly obovoid or orbicular, narrowing to an acute or acuminate
coriaceous portion that extends into the membranous tip, boundary between the
coriaceous and membranous portions not marked by minute hairs; anthers
0.4-1.1 mm. Caryopses 1.2-2.5 mm, broadly obovoid or spheroid, yellowish;
embryos 1.4-2 mm, 80-91% as long as the caryopses. 2n = 36.
Echinochloa muricata is native to North America, growing from
southern Canada to northern Mexico in moist, often disturbed sites (but
not rice fields). It resembles E. crus-galli in
gross morphology and ecology, but differs consistently by the characters
used in the key. The two varieties tend to be distinct, but there is
some overlap in both morphology and geography.
1 |
Spikelets 2.5-3.8 mm long; lower lemmas unawned
or awned, the awns to 10 mm long ..... var. microstachya |
| Spikelets 3.5-5 mm long; lower lemmas usually awned, the awns 6-16 mm long ..... var. muricata |
Echinochloa muricata var. microstachya Wiegand
Échinochloa de l'Ouest
Spikelets 2.5-3.8 mm. Lower glumes 0.9-1.6 mm;
upper glumes 2.8-3.8 mm; lower lemmas unawned or awned, awns to
10 mm; anthers 0.4-0.7 mm.
Echinochloa muricata var. microstachya is the common variety in
the western part of North America, extending east to the Missouri River and
the Texas panhandle.
Echinochloa muricata (P. Beauv.) Fernald var. muricata
Échinochloa Piquant
Spikelets 3.5-5 mm. Lower glumes 1-2.6 mm; upper
glumes3-5 mm; lower lemmas usually awned, awns 6-16 mm, occasionally
unawned; anthers 0.5-1.1 mm.
Echinochloa muricata var. muricata is the common variety in eastern
North America.
5. Echinochloa walteri (Pursh) A. Heller
Coast Barnyard Grass
Plants annual. Culms (30)50-200+ cm tall, to 2.5
cm thick; nodes pilose or villous, upper nodes usually with sparser and
shorter pubescence, occasionally glabrous. Lower sheaths usually hispid,
hairs papillose-based, sometimes just papillose; upper sheaths hispid
or glabrous; ligules absent; blades to 55 cm long, 10-35(60) mm
wide, scabrous. Panicles 8.5-35 cm, erect to slightly drooping, nodes
hispid, hairs 3.5-5 mm, papillose-based, sometimes sparsely so, internodes usually
glabrous, sometimes hispid, hairs papillose-based; primary branches 1-10
cm, loosely erect, not concealed by the spikelets, nodes usually hispid, hairs
papillose-based, sometimes glabrous, internodes scabrous, sometimes also sparsely
hispid, hairs papillose-based; secondary branches present on the longer
primary branches. Spikelets 3-5 mm, disarticulating at maturity, scabrous
to variously muricate and hairy, hairs usually not papillose-based, margins
sometimes with a few papillose-based hairs. Lower glumes usually more
than 1/2 as long as the spikelets, abruptly narrowing to a fine, 0.5 mm point;
lower florets sterile; lower lemmas usually awned, awns 8-25(60)
mm; lower paleas subequal to the lower lemmas; upper lemmas 3-5
mm long, about 1.5 mm wide, not or scarcely exceeding the upper glumes, narrowly
ovate to elliptical, coriaceous portion subacute, tips acuminate, membranous,
without a line of hairs at the base of the tip; anthers 0.6-1(1.2) mm.
Caryopses 1.2-1.8 mm, brownish; embryos 52-77% as long as the
caryopses. 2n = 36.
Echinochloa walteri grows in wet places, often in shallow water and brackish
marshes. It is a native species, found in both disturbed and undisturbed sites
although not in rice fields. Occasional specimens of E. walteri with
glabrous lower sheaths and short awns can be distinguished from E.
crus-pavonis by their less dense panicles.
6. Echinochloa crus-pavonis (Kunth) Schult.
Gulf Barnyard Grass
Plants annual or short-lived perennials. Culms
30-150 cm; nodes glabrous. Sheaths glabrous, often purplish; ligules
absent; blades 12-60 cm long, 10-25 mm wide, glabrous. Panicles
10-30 cm, erect or drooping, nodes sparsely hispid, hairs papillose-based, internodes
glabrous; primary branches to 14 cm, nodes sometimes sparsely hispid,
hairs papillose-based, internodes usually glabrous; secondary branches
to 3 cm. Spikelets 2.5-3.4 mm long, 1.2-1.4 mm wide, disarticulating
at maturity. Upper glumes subequal to the spikelets; lower florets
sterile; lower lemmas unawned or awned, awns 3-10(15) mm, curved; lower
paleas absent, vestigial, or well-developed; upper lemmas narrowly
elliptic, not or scarcely exceeding the upper glumes, acute or obtuse, with
a well-differentiated, early-withering tip, glabrous or pubescent at the base
of the tip, hairs not forming a line across the base; anthers 0.5-0.7
mm. Caryopses 1.2-1.5 mm long, 1-1.3 mm wide; embryos 50-70% as
long as the caryopses. 2n = 36.
Echinochloa crus-pavonis is a native species found in scattered locations
from British Columbia to Arizona, east to Florida, and south into South America.
It favors marshes and wet places at lower elevations, often being found in the
water.
1 |
Lower paleas more than 1/2 as long as the lemmas;
panicles usually drooping ..... var. crus-pavonis |
Lower paleas absent or much less
than 1/2 as long as the lemmas; panicles usually stiffly erect
..... var. macera |
Echinochloa crus-pavonis (Kunth) Schult. var. crus-pavonis
Panicles usually drooping. Lower paleas 1/2 or more as long as
the lower lemmas.
Echinochloa crus-pavonis var. crus-pavonis is generally the
more southern of the two varieties, extending through Mexico and the Caribbean
to Bolivia and Argentina. It appears, presumably as an adventive species, as
far north as Humboldt County, California.
Echinochloa crus-pavonis var. macera (Wiegand)
Gould
Panicles usually erect, stiff. Lower paleas absent or much less
than 1/2 as long as the lower lemmas.
Echinochloa crus-pavonis var. macera extends south only as
far as northern Mexico.
7. Echinochloa oplismenoides (E. Fourn.) Hitchc.
Chihuahuan Barnyard Grass
Plants annual. Culms to 100 cm, erect, succulent, glabrous, branching
from the lower nodes. Sheaths glabrous or hispid with papillose-based hairs;
ligules absent or the ligule region pubescent; blades 10-35 cm long,
5-10 mm wide. Panicles 15-30 cm, narrow; primary branches appressed
to ascending, with papillose-based hairs at the base of the spikelets. Spikelets
4-5 mm, disarticulating at maturity. Glumes with hairs over the veins,
glabrous, scabrous, or hispid between the veins; upper glumes about equal
to the spikelets, muticous or awned, awns to 1 mm; lower florets sterile;
lower lemmas unawned or awned, awns 8-16(50) mm; lower paleas absent
or hyaline and subequal to the lemmas; upper lemmas 4-4.5 mm long, 1.7-1.9
mm wide, elliptic; anthers 0.5-0.7 mm, purple. Caryopses2.7-2.9
mm long, 1.7-1.8 mm wide, elliptic in outline, mucronate; embryos about
75% as long as the caryopses; hila obovate. 2n = unknown.
Echinochloa oplismenoides was first found in the United States, in southern
Arizona, in 1993 (Fishbein 1995). It was previously known only from Mexico, with
a range that extends from northwestern Mexico to Guatemala. The southern Arizonan
plants were found near a cattle tank in wet grasslands. Fishbein stated that it
was impossible to tell whether they represented a previously overlooked native
species or an introduction.
8. Echinochloa colona (L.) Link
Awnless Barnyard Grass
Plants annual; erect or decumbent, cespitose or spreading,
rooting from the lower cauline nodes. Culms 10-70 cm; lower nodes
glabrous or hispid, hairs appressed; upper nodes glabrous. Sheaths
glabrous; ligules absent, ligule region frequently brown-purple; blades
8-22 cm long, 3-6(10) mm wide, mostly glabrous, sometimes hispid, hairs papillose-based
on or near the margins. Panicles 2-12 cm, erect, rachises glabrous or
sparsely hispid; primary branches 5-10, 0.7-2(4) cm, erect to ascending,
spikelike, somewhat distant, without secondary branches, axes glabrous or sparsely
hispid, hairs 1.5-2.5 mm, papillose-based. Spikelets 2-3 mm, disarticulating
at maturity, pubescent to hispid, hairs usually not papillose-based, tips acute
to cuspidate. Lower glumes about 1/2 as long as the spikelets; upper
glumes about as long as the spikelets; lower florets usually sterile,
occasionally staminate; lower lemmas unawned, similar to the upper glumes;
lower paleas subequal to the lemmas; upper lemmas 2.6-2.9 mm,
not or scarcely exceeding the upper glumes, elliptic, coriaceous portion rounded
distally, passing abruptly into a sharply differentiated, membranous, soon-withering
tip; anthers 0.7-0.8 mm. Caryopses 1.2-1.6 mm, whitish; embryos
63-83% as long as the caryopses. 2n = 54.
Echinochloa colona is widespread in tropical and subtropical regions.
It is adventive and weedy in North America, growing in low-lying, damp to wet,
disturbed areas, including rice fields. The unbranched, rather widely-spaced
panicle branches make this one of the easier species of Echinochloa to
recognize.
Hitchcock (1913) considered that colonum was a non-declining
contraction, but dictionaries of Linnaeus' time treated it as a declining
adjective. Because Linnaeus was the first to name the species (as Panicum
colonum),
it seems best to follow the practice considered correct in his day; hence E.
colona. (See also Nicolson
1986.)
9. Echinochloa frumentacea Link
Siberian Millet, White Panic
Plants annual. Culms 70-150 cm, erect, glabrous. Sheaths
glabrous; ligules absent; blades 8-35 cm long, 3-20(30) mm
wide, glabrous. Panicles 7-18 cm, erect to slightly drooping at
maturity, rachises not or only sparsely hispid, nodes with papillose-based
hairs; branches
numerous, appressed or ascending, spikelike, not or only sparsely hispid,
hairs papillose-based; primary branches 1.5-4 cm, glabrous or sparsely
hispid, hairs to 3 mm, papillose-based; secondary branches, if
present, usually concealed by the densely packed spikelets; longer
pedicels 0.2-0.5 mm.
Spikelets 3-3.5 mm, often with 1 sterile and 2 bisexual florets,
not disarticulating at maturity (particularly those near the bases of
the panicles), scabrous or short-hispid but without papillose-based hairs,
green and pale at maturity, apices usually obtuse, varying to acute. Upper
glumes narrower and shorter than the upper
lemmas; lower florets sterile; lower lemmas unawned; lower
paleas
subequal to the lower lemmas; upper lemmas 2.5-3 mm, ovate to elliptic,
coriaceous portion terminating abruptly at the base of the membranous
tip; anthers
0.8-1 mm. Caryopses 1.7-2.2 mm long, 1.6-1.8 mm wide, whitish; embryos
66-86% as long as the caryopses. 2n = 36 [J.D.
Munshi, J.K. Pal, V. Bansikar, and N. Pandit. 1994. Eco-cytological studies
of Echinochloa frumentacea (Roxb.) Link of Diara Lands of Bhagalpur,
Bihar. Proc. Indian Sci, Congr. 81:112],
54.
Echinochloa frumentacea originated in India, and possibly also in Africa.
It is grown for grain, fodder, and beer, although not as extensively as in the
past. It is found occasionally in the contiguous United States and southern Canada,
the primary source being birdseed mixes. It used to be confused with E.
esculenta, from which it differs in its whitish caryopses and proportionately
smaller embryos. Hybrids between E. frumentacea and E.
colona are partially fertile; those with E. esculenta are sterile.
10. Echinochloa crus-galli (L.) P. Beauv.
Barnyard Grass, Échinochloa Pied-de-Coq
Plants annual. Culms 30-200 cm, spreading, decumbent
or stiffly erect; nodes usually glabrous or the lower nodes puberulent.
Sheaths glabrous; ligules absent, ligule region sometimes pubescent;
blades to 65 cm long, 5-30 mm wide, usually glabrous, occasionally sparsely
hirsute. Panicles 5-25 cm, with few-many papillose-based hairs at or
below the nodes of the primary axes, hairs sometimes longer than the spikelets;
primary branches 1.5-10 cm, erect to spreading, longer branches with
short, inconspicuous secondary branches, axes scabrous, sometimes also sparsely
hispid, hairs to 5 mm, papillose-based. Spikelets 2.5-4 mm long, 1.1-2.3
mm wide, disarticulating at maturity. Upper glumes about as long as the
spikelets; lower florets sterile; lower lemmas unawned to awned,
sometimes varying within a branch, awns to 50 mm; lower paleas subequal
to the lemmas; upper lemmas broadly ovate to elliptical, coriaceous portion
rounded distally, passing abruptly into an early-withering, acuminate, membranous
tip that is further demarcated from the coriaceous portion by a line of minute
hairs (use 25× magnification); anthers 0.5-1 mm. Caryopses
1.3-2.2 mm long, 1-1.8 mm wide, ovoid or oblong, brownish; embryos 59-86%
as long as the caryopses. 2n = 54.
Echinochloa crus-galli is a Eurasian species that is now widely established
in the Flora region, where it grows in moist, disturbed sites, including
rice fields. Some North American taxonomists have interpreted Echinochloa
crus-galli much more widely; others treat it as here, but recognize several
infraspecific taxa based on such characters as trichome length and abundance,
and awn length. There are several ecological and physiological ecotypes within
the species, but the correlation between most of these and the species morphological
variation has not been established, so no infraspecific taxa are recognized
here.
11. Echinochloa esculenta (A. Braun) H. Scholtz
Japanese Millet
Plants annual. Culms 80-150 cm tall, 4-10 mm thick, glabrous. Sheaths
glabrous; ligules absent, ligule region sometimes pubescent; blades
10-50 cm long, 5-25 mm wide. Panicles 7-30 cm, dense, rachis nodes densely
hispid, hairs papillose-based, internodes scabrous; primary branches 2-5
cm, erect or spreading, simple or branched, often incurved at maturity, nodes
hispid, hairs papillose-based, internodes usually scabrous; longer pedicels
0.5-1 mm. Spikelets 3-4 mm long, 2-2.5 mm wide, not or only tardily disarticulating
at maturity, obtuse to shortly acute, purplish to blackish-brown at maturity.
Upper glumes narrower and shorter than the upper lemmas; lower florets
sterile; lower lemmas usually unawned; lower paleas shorter and
narrower than the lemmas; upper lemmas longer and wider than the upper
glumes, broadly ovate to ovate-orbicular, shortly apiculate, exposed distally
at maturity; anthers 1-1.2 mm. Caryopses 1.2-2.3 mm, brownish; embryos
84-96% as long as the caryopses. 2n = 54.
Echinochloa esculenta was derived from E.
crus-galli in Japan, Korea, and China. It is cultivated for fodder, grain,
or birdseed. It has sometimes been included in Echinochloa
frumentacea, from which it differs in its brownish caryopses and longer
pedicels. Hybrids between E. crus-galli and E. esculenta are fully
fertile, but those with E. frumentacea are sterile.
12. Echinochloa oryzoides (Ard.) Fritsch
Early Barnyard Grass
Plants annual. Culms 40-120 cm, erect, densely
tufted; nodes glabrous. Sheaths glabrous; ligules absent;
blades lax or drooping, 7-20 cm long, 4-12 mm wide, mostly glabrous.
Panicles 8-17(25) cm, lax, horizontal to strongly drooping, rachis nodes
hispid, hairs to 4 mm, papillose-based, internodes glabrous; primary branches
to 5 cm, appressed to the rachises, mostly simple, glabrous or sparsely
hispid, hairs to 3.5 mm, papillose-based, particularly at the nodes. Spikelets
3.7-7 mm long, 1.9-2.4 mm wide, disarticulating at maturity, broadly ovate to
ovate. Lower glumes usually 1/4-2/5 as long as the spikelets, occasionally
1/2 as long or longer; upper glumes subequal to the spikelets; lower
florets sterile; lower lemmas similar in size to the spikelet, usually
awned, awns to 5 cm; lower paleas well-developed; upper lemmas
3.5-4.5 mm, similar in length and width to the upper glumes, broadly elliptic
to ovate, with an acute, greenish tip; anthers to 0.8 mm. Caryopses
1.9-3 mm, light brown or tan; embryos 70-85% as long as the caryopses.
2n = 54.
Echinochloa oryzoides is a common weed of rice fields throughout the
world, growing in the flooded portions of the fields. It was included in Echinochloa
oryzicola by Gould et al. (1972), but it differs in its shorter embryo,
lax, strongly drooping panicle, and earlier (June-July) flowering period. This
flowering period is also earlier than that of Oryza.
In addition, E. oryzoides is usually conspicuously awned, having longer
awns than even the awned variants of E. oryzicola, and it is rarely obviously
pubescent on the cauline nodes, leaf sheaths, and collars. The earliest known
collection of Echinochloa oryzoides in the United States was made in
1925 (Barrett and Seaman 1980).
13. Echinochloa oryzicola (Vasinger) Vasinger
Late Barnyard Grass
Plants annual. Culms 40-150 cm, erect or nearly
so, densely tufted; lower nodes usually antrorsely scabrous or villous;
upper nodes glabrous. Lower sheaths densely pubescent; upper
sheaths glabrous or pubescent at the throat, and sometimes on the collar;
ligules absent; blades stiff, ascending, lower blades pubescent,
upper blades usually glabrous. Panicles 8-20 cm, erect to slightly drooping,
rachis nodes hispid, with papillose-based hairs to 5.6 mm, internodes usually
scabrous, sometimes also with a few papillose-based hairs; primary branches
to 4 cm. Spikelets 4-6 mm, ovoid to ellipsoid, disarticulating at maturity.
Lower glumes usually at least 1/2 as long as the spikelets; upper
glumes equaling or exceeding the upper florets; lower florets sterile;
lower lemmas often thickened and somewhat coriaceous, unawned or awned,
awns to 1.5 mm; lower paleas well-developed; upper lemmas broadly
ovate to elliptical, coriaceous portion rounded distally, passing abruptly into
an early-withering, acuminate, membranous tip that is further demarcated from
the coriaceous portion by minute hairs (use 25× magnification); anthers
0.9-1.2 mm. Caryopses 1.7-2.6 mm, brownish; embryos 89-98% as
long as the caryopses. 2n = 36.
Like Echinochloa oryzoides, Echinochloa
oryzicola is an introduced weed of rice fields, where it grows in the flooded
portion, with the rice. The two are quite distinct, with E. oryzicola
flowering after Oryza and having a longer embryo
and an erect panicle. It is also more likely to have evidently pubescent cauline
nodes, leaf sheaths, and collars than E. oryzoides and is never conspicuously
awned.