| Mary E. Barkworth |
Plants annual or perennial; habit various. Culms 3-800
cm, annual, usually not woody. Leaves 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. Inflorescences 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. Spikelets 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. Glumes 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. Caryopses usually dorsally compressed
or terete; embryos 1/2 or more the length of the caryopses. x =
9, 10.
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; Giussani 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. Giussani 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 photosynthetic 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.
1 |
.Plants
developing both subterranean and aerial inflorescences, only
the subterranean spikelets setting seed ..... 25.04 Amphicarpum |
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 and specimens
have no underground parts] (2) |
|
Inflorescences
spikelike panicles, with the branches partially embedded in the
flattened rachises; plants perennial, stoloniferous ..... 25.22 Stenotaphrum |
|
Inflorescences panicles, sometimes
spikelike, but the branches not embedded in the rachises or the
rachises not flattened; plants annual or perennial, sometimes
stoloniferous (3) |
|
Most spikelets
or groups of 2-11 spikelets subtended by 1-many, distinct to
more or less connate, stiff bristles or bracts (4) |
|
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) |
|
Spikelets in groups
of 2-11, subtended by 4 flat, narrowly elliptic, coriaceous bracts;
terete bristles not present ..... 25.17 Anthephora |
|
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) |
|
Bristles falling
with the spikelets at maturity; disarticulation at the base of
the reduced panicle branches (fascicles) (6) |
|
Bristles persistent; disarticulation
below the spikelets (7) |
|
Bristles plumose
or antrorsely scabrous, free or fused no more than 1/2 their
length ..... 25.15 Pennisetum |
|
Bristles glabrous, smooth, retrorsely
scabrous, or strigose, usually at least some bristles fused for
more than 1/2 their length ..... 25.16 Cenchrus |
|
Upper glumes indurate
at maturity; lower lemmas somewhat indurate at the base; pedicels
subtended by a single bristle ..... 25.19 Setariopsis |
|
Upper glumes membranous to herbaceous
at maturity; lower lemmas neither constricted nor indurate at
the base; pedicels subtended by 1-many bristles (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 ..... 25.20 Setaria |
|
Spikelets subtended by 1 bristle;
paleas of the lower florets coriaceous to indurate at maturity,
the keels thickened ..... 25.18 Ixophorus |
|
Terminal spikelet
on each branch subtended by a single bristle; other spikelets
occasionally with a single stiff subtending bristle ..... 25.20 Setaria |
|
None of the spikelets subtended
by a stiff bristle (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
..... 25.21 Paspalidium |
|
Inflorescences various but, if
of spikelike branches, these terminating in a well-developed
or rudimentary spikelet (11) |
|
Lower glumes
or lower lemmas awned, sometimes shortly so (the awn reduced
to a point in Echinochloa
colona) (12) |
|
Lower glumes and lower lemmas
unawned (15) |
|
Upper florets
laterally compressed; spikelets also laterally compressed .....
25.12 Melinis |
|
Upper florets dorsally compressed;
spikelets usually dorsally compressed or terete, sometimes laterally
compressed (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 ..... 25.07 Echinochloa |
|
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) |
|
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 ..... 25.06 Oplismenus |
|
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 ..... 25.03 Alloteropsis |
|
Upper florets
laterally compressed ..... 25.12 Melinis |
|
Upper florets dorsally compressed
or terete (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) |
|
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) |
|
Aerial inflorescences
with elongate rachises and glabrous spikelets; spikelets of the
aerial panicles rarely setting seed; subterranean spikelets developed,
seed-forming ..... 25.04 Amphicarpum |
|
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) |
|
Spikelets ellipsoid
to obovoid; inflorescences simple panicles with erect to ascending
branches on elongate rachises; branches ascending, not conspicuously
spikelike ..... 25.02 Anthenantia |
|
Spikelets lanceoloid to ellipsoid;
inflorescences usually panicles with digitate or subdigitate
clusters of spikelike branches, sometimes simple panicles with
strongly divergent branches ..... 25.01 Digitaria |
|
Spikelets subtended
by a cuplike callus ..... 25.14 Eriochloa |
|
Spikelets not subtended by a cuplike
callus (20) |
|
At least the
upper leaves, often all leaves, without ligules; ligules, when
present, of hairs ..... 25.07 Echinochloa |
|
All leaves with ligules, ligules
membranous or of hairs (21) |
|
Paleas of the
lower florets inflated and indurate at maturity; lower and upper
florets standing apart from each other when mature ..... 25.24 Steinchisma |
|
Paleas of the lower florets neither
inflated nor indurate at maturity; lower and upper florets closely
appressed to each other when mature (22) |
|
Inflorescences
of 1-sided, spikelike primary branches (23) |
|
Inflorescences usually panicles
with well-developed secondary branches, sometimes spikelike panicles
or panicles with spikelike, but not 1-sided, branches (28) |
|
Spikelets with
the lower lemmas and lower glumes (if present) adjacent to the
branch axes (24) |
|
Spikelets with the upper lemmas
and upper glumes (if present) adjacent or appressed to the branch
axes (26) |
|
Lower glumes
absent ..... 25.25 Axonopus |
|
Lower glumes present on all or
most spikelets (25) |
|
Upper lemmas
rugose and verrucose; panicle branches in 2 or more ranks, sometimes
verticillate ..... 25.13 Urochloa |
|
Upper lemmas smooth; panicle branches
in 1 rank ..... 25.11 Moorochloa |
|
Both glumes absent
from all or almost all spikelets, the terminal spikelet on a
branch sometimes with upper glumes ..... 25.27 Reimarochloa |
|
Upper or both glumes present on
all spikelets (27) |
|
Upper lemmas
smooth to slightly rugose; lower glumes usually absent .....
25.26 Paspalum |
|
Upper lemmas rugose and verrucose;
lower glumes present, from 1/5 as long as the spikelets to equaling
them ..... 25.13 Urochloa |
|
Inflorescences
dense, the spikelets concealing at least the distal 1/2 of the
rachises (29) |
|
Inflorescences more or less open
panicles, the spikeletsnot concealing the rachises (30) |
|
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 ..... 25.08 Sacciolepis |
|
Upper glumes not saccate, 3-7-veined;
panicle branches not fused to the rachises; blades 12-28 mm wide;
culm internodes filled with aerenchyma ..... 25.23 Hymenachne |
|
Lower glumes
with saccate bases; glumes and lemmas with woolly pubescent apices;
culms weakly lignified, rooting at the nodes ..... 25.05 Lasiacis |
|
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) |
|
Lemmas of the
upper florets rugose and verrucose; panicle branches usually
spikelike and 1-sided, alternate or subopposite, less frequently
verticillate ..... 25.13 Urochloa |
|
Lemmas of the upper florets usually
smooth, if rugose the panicle branches neither verticillate nor
1-sided and spikelike (32) |
|
Plants developing
aerial and subterranean panicles; aerial spikelets lanceoloid,
often without lower glumes; upper lemmas with flat margins .....
25.04 Amphicarpum |
|
Plants developing aerial, but
not subterranean, panicles; spikelets ovoid to ellipsoid or lanceoloid;
lower glumes present; upper lemmas with involute margins (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 .....
25.09 Dichanthelium |
|
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) |
|
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 ..... 25.09 Dichanthelium |
|
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 ..... 25.10 Panicum |