Poaceae tribe Paniceae
Echinochloa
barnyard-grass, cockspur grass
3-800 cm, annual, usually not woody.
10-460 cm, prostrate, decumbent or erect, distal portions sometimes floating, sometimes rooting at the lower nodes;
nodes usually glabrous;
internodes hollow or solid.
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.
basal and/or cauline;
sheaths usually open;
ligules of hairs or membranous, membranous ligules often ciliate, cilia sometimes longer than the membranous base;
blades occasionally pseudopetiolate, seldom disarticulating at maturity.
terminal, sometimes also axillary, occasionally subterranean panicles;
branches sometimes spikelike and secund, sometimes less than 1 cm;
disarticulation usually below the glumes, sometimes at the base of the panicle branches, occasionally below the florets.
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).
usually dorsally compressed, varying to terete or laterally compressed, with 2(3) florets, lower florets staminate, sterile, or reduced, upper florets usually bisexual;
calluses not developed.
plano-convex, with 2(3) florets;
lower florets sterile or staminate;
upper florets bisexual, dorsally compressed.
usually membranous;
lower glumes usually less than 1/2 as long as the spikelets, sometimes absent;
upper glumes usually subequal to the upper florets, occasionally absent;
lower lemmas similar to the upper glumes in length and texture;
upper lemmas indurate, coriaceous, or cartilaginous, with a germination flap at the base, margins usually widely separated and involute at maturity, sometimes flat and hyaline;
upper paleas similar to the upper lemmas in length and texture;
lodicules short;
anthers usually 3;
stigmas usually red.
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.
usually dorsally compressed or terete;
embryos 1/2 or more the length of the caryopses.
ellipsoid, broadly ovoid or spheroid;
embryos usually 0.7-0.9 times as long as the caryopses.
= 9, 10.
= 9.
Poaceae tribe Paniceae
Echinochloa
The tribe Paniceae, which includes about 100 genera and 2000 species, is primarily tropical in distribution. Within the Flora region, it is represented by 27 genera and 262 species, with its greatest representation being in the eastern portion of the contiguous United States (Barkworth and Capels 2000).
The tribe is so morphologically distinct that it was first recognized, in essentially its current sense, by Robert Brown in 1814. Its primary distinguishing features are the unusual spikelet structure combined with the indurate to coriaceous upper florets. Recent molecular studies (Barber et al. 2002; Guissani et al. 2001) show it as comprising two distinct lineages, one of which contains species with a base number of x = 9 and the other, species with x = 10.
Photosynthesis in the Paniceae may follow the C3 pathway or any of three different C4 pathways. Most genera are uniform in this regard, but there are some noteworthy exceptions. Guissani et al. (2001) concluded that the C3 pathway is probably ancestral within the tribe and that two of the three C4 pathways, NAD-ME and PCK, originated only once within the tribe, whereas the NADP-ME pathway originated independently in several different lineages. Most genera are uniform with respect to their photosyntheticd pathway, but there are some noteworthy exceptions.
The germination flap is a small area of soft tissue at the base of the upper lemma through which the primary root of the seedling grows.
(Discussion copyrighted by Flora of North America; reprinted with permission.)
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 probably reflected in the mapping of both E. 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 Echinochloa frumentacea and E. esculenta are domesticated derivatives of E. crus-galli and E. colona, respectively (Yabuno 1962) and that E. oryzoides is very closely related to E. crus-galli (Yabuno 1984). 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; Gonzalez-Andres et al. 1996; Asins 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 that voucher specimens are preserved.
(Discussion copyrighted by Flora of North America; reprinted with permission.)
1. Plants developing both subterranean and aerial inflorescences, only the subterranean spikelets setting seed | Amphicarpum |
1. Plants developing only aboveground inflorescences, the spikelets setting seed [Amphicarpum is also keyed out here to accommodate situations in which looking for subterranean inflorescences is not permitted or specimens have no underground parts]. | → 2 |
2. Inflorescences spikelike panicles, with the branches partially embedded in the flattened rachises; plants perennial, stoloniferous | Stenotaphrum |
2. Inflorescences panicles, sometimes spikelike, but the branches not embedded in the rachises or the rachises not flattened; plants annual or perennial, sometimes stoloniferous. | → 3 |
3. Most spikelets or groups of 2-11 spikelets subtended by 1-many, distinct to more or less connate, stiff bristles or bracts. | → 4 |
4. Spikelets in groups of 2-11, subtended by 4 flat, narrowly elliptic, coriaceous bracts; terete bristles not present | Anthephora |
4. Spikelets solitary or in groups, subtended by 1-many stiff, terete bristles, sometimes appearing as an extension of the branch; flat, connate bristles sometimes present distal to the terete bristles. | → 5 |
5. Bristles falling with the spikelets at maturity; disarticulation at the base of the reduced panicle branches (fascicles). | → 6 |
6. Bristles plumose or antrorsely scabrous, free or fused no more than 1/2 their length | Pennisetum |
6. Bristles glabrous, smooth, retrorsely scabrous, or strigose, usually at least some bristles fused for more than 1/2 their length | Cenchrus |
5. Bristles persistent; disarticulation below the spikelets. | → 7 |
7. Upper glumes indurate at maturity; lower lemmas somewhat indurate at the base; pedicels subtended by a single bristle | Setariopsis |
7. Upper glumes membranous to herbaceous at maturity; lower lemmas neither constricted nor indurate at the base; pedicels subtended by 1-many bristles. | → 8 |
8. Spikelets subtended by 1-many bristles; paleas of the lower florets usually hyaline to membranous at maturity, rarely absent or reduced; paleal veins not keeled | Setaria |
8. Spikelets subtended by 1 bristle; paleas of the lower florets coriaceous to indurate at maturity, the keels thickened | Ixophorus |
3. All or most spikelets not subtended by stiff bristles, sometimes the terminal spikelet on each branch subtended by a single bristle, and occasionally other spikelets with a single subtending bristle. | → 9 |
9. Terminal spikelet on each branch subtended by a single bristle; other spikelets occasionally with a single stiff subtending bristle | Setaria |
9. None of the spikelets subtended by a stiff bristle. | → 10 |
10. Inflorescences of spikelike branches 1-3.7 cm long, the branch axes extending as a 2.5-4 mm bristle beyond the base of the distal spikelets | Paspalidium |
10. Inflorescences various but, if of spikelike branches, these terminating in a well-developed or rudimentary spikelet. | → 11 |
11. Lower glumes or lower lemmas awned, sometimes shortly so (the awn reduced to a point in Echinochloa colona). | → 12 |
12. Upper florets laterally compressed; spikelets also laterally compressed | Melinis |
12. Upper florets dorsally compressed; spikelets usually dorsally compressed or terete, sometimes laterally compressed. | → 13 |
13. Blades linear to linear-lanceolate, usually more than 10 times longer than wide, with prominent midribs; at least the upper leaves, often all leaves, without ligules; ligules usually absent, particularly from the upper leaves, of hairs when present | Echinochloa |
13. Blades triangular to lanceolate, less than 10 times longer than wide, the midribs not particularly prominent, at least distally; ligules present, of hairs or membranous. | → 14 |
14. Lower glumes awned, the awns exceeding the florets; upper glumes not ciliate-margined; culms trailing on the ground, frequently rooting and branching at the nodes | Oplismenus |
14. Lower glumes unawned or shortly awned, the awns exceeded by the florets; upper glumes ciliate-margined; culms erect or decumbent below, sometimes rooting and branching at the lower nodes | Alloteropsis |
11. Lower glumes and lower lemmas unawned. | → 15 |
15. Upper florets laterally compressed | Melinis |
15. Upper florets dorsally compressed or terete. | → 16 |
16. Upper lemmas and paleas cartilaginous and flexible at maturity; lemma margins flat, hyaline; lower glumes absent or to 1/4 the length of the spikelets. | → 17 |
17. Aerial inflorescences with elongate rachises and glabrous spikelets; spikelets of the aerial panicles rarely setting seed; subterranean spikelets developed, seed-forming | Amphicarpum |
17. Aerial inflorescences of digitate or subdigitate clusters of spikelike branches with glabrous or pubescent spikelets or with elongate rachises and conspicuously pubescent spikelets; aerial spikelets seed-forming; subterranean spikelets not developed. | → 18 |
18. Spikelets ellipsoid to obovoid; inflorescences simple panicles with erect to ascending branches on elongate rachises; branches ascending, not conspicuously spikelike | Anthenantia |
18. Spikelets lanceoloid to ellipsoid; inflorescences usually panicles with digitate or subdigitate clusters of spikelike branches, sometimes simple panicles with strongly divergent branches | Digitaria |
16. Upper lemmas and paleas chartaceous to indurate and rigid at maturity; lemma margins not hyaline, frequently involute; lower glumes varying from absent to subequal to the spikelets or extending beyond the distal floret. | → 19 |
19. Spikelets subtended by a cuplike callus | Eriochloa |
19. Spikelets not subtended by a cuplike callus. | → 20 |
20. At least the upper leaves, often all leaves, without ligules; ligules, when present, of hairs | Echinochloa |
20. All leaves with ligules, ligules membranous or of hairs. | → 21 |
21. Paleas of the lower florets inflated and indurate at maturity; lower and upper florets standing apart from each other when mature | Steinchisma |
21. Paleas of the lower florets neither inflated nor indurate at maturity; lower and upper florets closely appressed to each other when mature, [revert to left side] | → 22 |
22. Inflorescences of 1-sided, spikelike primary branches. | → 23 |
23. Spikelets with the lower lemmas and lower glumes (if present) adjacent to the branch axes. | → 24 |
24. Lower glumes absent | Axonopus |
24. Lower glumes present on all or most spikelets. | → 25 |
25. Upper lemmas rugose and verrucose; panicle branches in 2 or more ranks, sometimes verticillate | Urochloa |
25. Upper lemmas smooth; panicle branches in 1 rank | Moorochloa |
23. Spikelets with the upper lemmas and upper glumes (if present) adjacent or appressed to the branch axes. | → 26 |
26. Both glumes absent from all or almost all spikelets, the terminal spikelet on a branch sometimes with upper glumes | Reimarochloa |
26. Upper or both glumes present on all spikelets. | → 27 |
27. Upper lemmas smooth to slightly rugose; lower glumes usually absent | Paspalum |
27. Upper lemmas rugose and verrucose; lower glumes present, from 1/5 as long as the spikelets to equaling them | Urochloa |
22. Inflorescences usually panicles with well-developed secondary branches, sometimes spikelike panicles or panicles with spikelike, but not 1-sided, branches. | → 28 |
28. Inflorescences dense, the spikelets concealing at least the distal 1/2 of the rachises. | → 29 |
29. Upper glumes slightly to strongly saccate, 5-13-veined; panicle branches often fused to the rachises; blades 1.5-22 mm wide; culm internodes hollow | Sacciolepis |
29. Upper glumes not saccate, 3-7-veined; panicle branches not fused to the rachises; blades 12-28 mm wide; culm internodes filled with aerenchyma | Hymenachne |
28. Inflorescences more or less open panicles, the spikelets not concealing the rachises. | → 30 |
30. Lower glumes with saccate bases; glumes and lemmas with woolly pubescent apices; culms weakly lignified, rooting at the nodes | Lasiacis |
30. Lower glumes not saccate basally; glumes and lemmas glabrous or with short, straight hairs, apices sometimes with a tuft of hairs but never woolly pubescent; culms usually not lignified, if lignified, not rooting at the nodes. | → 31 |
31. Lemmas of the upper florets rugose and verrucose; panicle branches usually spikelike and 1-sided, alternate or subopposite, less frequently verticillate | Urochloa |
31. Lemmas of the upper florets usually smooth, if rugose the panicle branches neither verticillate nor 1-sided and spikelike. | → 32 |
32. Plants developing aerial and subterranean panicles; aerial spikelets lanceoloid, often without lower glumes; upper lemmas with flat margins | Amphicarpum |
32. Plants developing aerial, but not subterranean, panicles; spikelets ovoid to ellipsoid or lanceoloid; lower glumes present; upper lemmas with involute margins. | → 33 |
33. Blades of the basal leaves clearly distinct from the cauline leaves; basal leaves ovate to lanceolate, cauline leaves with longer and narrower blades; basal leaves forming a distinct winter rosette | Dichanthelium |
33. Blades of the basal and cauline leaves similar, usually linear to lanceolate, varying from filiform to ovate; basal leaves not forming a distinct winter rosette. | → 34 |
34. Panicles terminating the culms usually appearing in late spring; branches usually developing from the lower and middle cauline nodes in summer, the branches rebranching 1 or more times by fall; upper florets not disarticulating at maturity, plump | Dichanthelium |
34. Panicles terminating the culms usually appearing after midsummer; branches usually not developing branches from the lower and middle cauline nodes, when present, rarely rebranched; upper florets disarticulating or not very plump at maturity | Panicum |
1. Ligules of stiff hairs present on the lower leaves; lower florets staminate; plants perennial. | → 2 |
2. Plants without scaly rhizomes, sometimes rooting at the lower nodes; lower lemmas usually awned, sometimes merely apiculate; known outside of experimental plantings | E. polystachya |
2. Plants with short, scaly rhizomes; lower lemmas unawned, sometimes long-cuspidate; in the Flora region, known only from experimental plantings | E. pyramidalis |
1. 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 |
3. 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 |
4. 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 | E. frumentacea |
4. Spikelets purplish to blackish-brown at maturity, their apices obuse to shortly acute; rachis nodes densely hispid with papillose-based hairs; caryopses brownish | E. esculenta |
3. Lower lemmas often awned; spikelets disarticulating at maturity; upper lemmas not or scarcely exceeding the upper glumes in length and width at maturity. | → 5 |
5. 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 |
6. 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 | E. oryzoides |
6. 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 | E. oryzicola |
5. 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 |
7. Lower florets staminate; anthers of the upper florets 1.2-1.7 mm long | E. paludigena |
7. Lower florets sterile; anthers of the upper florets 0.5-1.2 mm long. | → 8 |
8. Panicle branches 0.7-2(4) cm long, without secondary branches; spikelets 2-3 mm long, unawned | E. colona |
8. 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 |
9. Upper lemmas broadly ovate to elliptical, if elliptical, each with a line of minute (need 25x magnification) hairs across the base of the early-withering tips. | → 10 |
10. 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 | E. crus-galli |
10. Upper lemmas with acute or acuminate coriaceous apices that extend into the membranous tip, without hairs at the base of the tip | E. muricata |
9. Upper lemmas narrowly ovate to elliptical, never with a line of minute hairs across the base of the early-withering, membranous tips. | → 11 |
11. Spikelets 2.5-3.4 mm long; lower lemmas unawned or with awns 3-10(15) mm long, curved | E. crus-pavonis |
11. Spikelets 3-5 mm long; lower lemmas usually with awns 8-25 mm long, typically straight. | → 12 |
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 | E. walteri |
12. 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 | E. oplismenoides |
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