| Mary E. Barkworth |
Plants annual or perennial. Culms 1-500
cm, not woody, usually not branched above the base. Sheaths usually
open, often with coarse hairs at the top; auricles rarely present; ligules of
hairs or membranous, if membranous, often ciliate, cilia sometimes
longer than the membranous base; blades often with stiff, coarse
marginal hairs adjacent to the ligules, glabrous or variously pubescent
elsewhere. Inflorescences terminal, sometimes also axillary,
simple panicles, panicles of 1-many spikelike branches, spikelike racemes,
spikes, or, in 1 genus, a solitary spikelet, in dioecious taxa the
staminate and pistillate inflorescences sometimes morphologically distinct; disarticulation usually
beneath the fertile florets or the glumes but, particularly if the
panicle branches are short, sometimes at the base of the branches. Spikelets usually
laterally compressed, with 1-60 florets, sterile or reduced florets,
if present, usually distal to the bisexual florets. Glumes from
shorter than the adjacent florets to exceeding the distal florets; lemmas 1-3-veined
or 7-13-veined, rarely 5-veined, if with 7-13 veins, the veins often
in 3 groups; lodicules 2, or absent. x = 7, 8, 9, 10,
12.
Most members of the Cynodonteae in the Flora region can be recognized by their possession of two or more of the following characteristics: 1-3- or 7-13-veined lemmas, laterally compressed spikelets, spikelike inflorescence branches, and the presence of coarse hairs near the junction of the sheath and blade. All employ the NAD-ME or PCK C4 photosynthetic pathways, have Kranz blade anatomy, and tend to grow in hot, dry areas. Having said this, it must be acknowledged that each of these characteristics can be found in other tribes and, within the Cynodonteae, there are genera that lack one or more of them. Nevertheless, it is usually fairly easy to recognize members of the tribe.
The tribe Cynodonteae, as interpreted here, includes genera that
are normally placed in two tribes, the Cynodonteae sensu
stricto and the Eragrostideae Stapf (Clayton and Renvoize 1986;
Peterson et al. 2001; Grass Phylogeny Working Group 2001, but see Campbell
1985). Genera 17.01 to 17.34 correspond to the Eragrostideae of
Clayton and Renvoize, and genera 17.35 to 17.53 to the Cynodonteae. The
two are treated as one here because recent morphological, anatomical, and
molecular studies (Van den Borre 1994; Van den Borre and Watson 1997; Hilu
and Alice 2001) indicate that the distinction between the two is artificial,
but there is no agreement on an alternative treatment. Part of the problem
is that several of the genera (e.g., Eragrostis, Chloris, Muhlenbergia,
and Sporobolus) are polythetic (Van
den Borre and Watson 1997; Hilu and Alice 2001). The uncertainty concerning
relationships within the tribe is reflected in the classification of Peterson
et al. (2001), which lists 23 of the 53 genera treated here as being of
uncertain position within the subfamily Chloridoideae.
1 |
Inflorescences
clearly exceeded by the upper leaves, often completely or almost
completely enclosed in the upper leaf sheaths; culms 1-30(75)
cm tall (2) |
Inflorescences usually equaling
or exceeding the upper leaves; culms 1-500 cm tall (13) |
|
Lemmas 3-lobed,
the lobes ciliate; spikelets with 4 florets ..... 17.17 Blepharidachne |
|
Lemmas not 3-lobed or the lobes
not ciliate; spikelets with 1-60 florets (3) |
|
Spikelets (and
often the plants) unisexual (4) |
|
Spikelets bisexual, usually at
least the lowest floret in each spikelet bisexual, in Dasyochloa the
third floret in each spikelet bisexual or pistillate, if pistillate,
the lowest 2 florets staminate (8) |
|
Leaves strongly
distichous; lemmas 9-11-veined; plants unisexual, growing in
saline and alkaline soils Distichlis |
|
Leaves not strongly distichous;
lemmas 1-5-veined; plants unisexual or, if bisexual, with separate
pistillate and staminate inflorescences, growing in a variety
of soils (5) |
|
Spikelets 5-26
mm long; pistillate and staminate inflorescences similar, simple
panicles; glumes and lemmas unawned, mucronate, or 1-awned .....
17.23 Eragrostis |
|
Spikelets 2.5-7 mm long; pistillate
and staminate inflorescences strongly dimorphic; staminate inflorescences
with pectinate, spikelike branches; pistillate spikelets with
conspicuously 3-awned glumes or distal florets (7) |
|
Upper glumes of
the pistillate spikelets white, rigid, globose structures terminating
in 3 awnlike teeth; pistillate spikelets with 1 floret, without
rudimentary florets, the floret unawned or shortly 3-awned; staminate
spikelets 4-6 mm long; anthers 2.5-3 mm long, brownish, red,
or orange; widespread species of the central plains ..... 17.48 Buchloë |
|
Upper glumes of the pistillate
spikelets membranous, unawned; pistillate spikelets with one
3-awned floret and a distal 3-awned rudiment; staminate spikelets
3-4 mm long; anthers 2-2.5 mm long, pale; known, within the Flora region,
only from Florida ..... 17.47 Opizia |
|
Lemma margins
with a tuft of hairs at midlength, glabrous elsewhere; blades
with white, thickened margins and sharply pointed ..... 17.18 Munroa |
|
Lemma margins glabrous, or with
hairs but the hairs not forming a tuft at midlength; blades without
white cartilaginous margins, not sharply pointed (9) |
|
Lemmas awned,
the awns 1-11 mm long (10) |
|
Lemmas unawned, sometimes mucronate,
with mucros less than 1 mm long (11) |
|
Plants
stoloniferous; inflorescences 1-2.5 cm long, dense panicles;
lemmas bilobed, the lobes about 1/2 as long as the lemmas; ligules
of hairs ..... 17.15 Dasyochloa |
|
Plants not stoloniferous; inflorescences
1.5-76 cm long, not dense; lemmas entire or minutely bilobed;
ligules membranous, sometimes ciliate ..... 17.19 Leptochloa |
|
Spikelets with
2-20 florets; inflorescences panicles of 2-120 spikelike branches
..... 17.19 Leptochloa |
|
Spikelets with 1(3) florets; inflorescences
simple panicles, often highly contracted, without spikelike branches
(12) |
|
Inflorescences
0.3-7.5 cm long, dense, spikelike or capitate panicles 1-8 times
longer than wide; glumes strongly keeled; plants annual .....
17.31 Crypsis |
|
Inflorescences 1-60 cm long, sometimes
dense and spikelike but, if less than 8 cm long, more than 8
times longer than wide; glumes rounded or weakly keeled; plants
annual or perennial ..... 17.30 Sporobolus |
|
Inflorescences without disarticulating
branches; branches, if present, often more than 4.5 cm long,
variously shaped, including spikelike but not globose, often
with more than 16 spikelets per branch (22) |
|
Upper glumes
with straight or uncinate spinelike projections; spikelets crowded,
the branches condensed into burs ..... 17.52 Tragus |
|
Upper glumes without spinelike
projections; spikelets sometimes crowded, but not forming burlike
clusters (15) |
|
Branches not
fused to the rachises; spikelets more than 3 per branch, usually
all alike, sometimes the proximal spikelet sterile or replaced
with short secondary branches (16) |
|
Branches sometimes fused to the
rachises; spikelets 1-3 per branch, usually some spikelets on
each branch sterile or staminate and 1 pistillate or bisexual
(18) |
|
Both glumes much
longer than the florets, usually exceeding the distal florets;
plants annual ..... 17.22 Dinebra |
|
Lower glumes shorter than or subequal
to the lower florets, 1 or both glumes usually exceeded by the
distal florets; plants usually perennial (17) |
|
Spikelets with
1-2(3) florets; branches disarticulating promptly, before the
spikelets disarticulate ..... 17.46 Bouteloua |
|
Spikelets with 2-8 florets; branches
disarticulating tardily, initially the spikelets disarticulating
above the glumes ..... 17.25 Pogonarthria |
|
Axes of the branches
extending beyond the base of the distal florets ..... 17.46 Bouteloua |
|
Axes of the branches terminating
at the base of the distal spikelet (19) |
|
Spikelets in
pairs; glumes awned, the lower glumes (1)2(3)-awned, the upper
glumes 1-awned; panicle branches often fused to the rachises
..... 17.34 Lycurus |
|
Spikelets in triplets; glumes
unawned, 1-awned, or 3-awned; panicle branches sometimes appressed,
but not fused, to the rachises (20) |
|
Branches straight
at the base; central spikelets sessile ..... 17.51 Hilaria |
|
Branches sharply curved at the
base; central spikelets sessile or pedicellate (21) |
|
Central spikelets
with 1 bisexual floret; lateral spikelets with 1 floret, varying
to rudimentary ..... 17.50 Aegopogon |
|
Central spikelets with 3-4 florets,
the lowest floret pistillate, bisexual, or staminate, the distal
florets staminate or sterile; lateral spikelets usually with
2 florets ..... 17.49 Cathestecum |
|
Inflorescences
spikes or racemes (23) |
|
Inflorescences simple panicles
(sometimes highly condensed) or panicles of 1-120 spikelike branches
(32) |
|
Spikelets with
1 bisexual or staminate floret and no additional florets (24) |
|
Spikelets with more than 1 floret
but sometimes only 1 floret bisexual, the additional florets
sterile or staminate (26) |
|
Rachises falcate
or curved; both glumes exceeding the florets ..... 17.43 Microchloa |
|
Rachises straight; lower glumes
exceeded by the florets, sometimes absent (25) |
|
Spikelets solitary
at each node; disarticulation below the glumes or the spikelets
not disarticulating ..... 17.53 Zoysia |
|
Spikelets paired, terminal on
branches that are fused to the rachises; disarticulation at the
base of the fused pedicel pairs ..... 17.34 Lycurus |
|
All spikelets
unisexual, the functional florets either staminate or pistillate;
plants either unisexual or with both pistillate and staminate
spikelets (27) |
|
Some or all spikelets bisexual,
the florets bisexual or unisexual, but both staminate and pistillate
florets present within an individual spikelet (29) |
|
Lemmas 9-11-veined;
plants of saline habitats ..... 17.04 Distichlis |
|
Lemmas 3-veined; plants of various
habitats (28) |
|
Lemmas of the
pistillate florets with awns 3.4-6.8 mm long; branches of staminate
inflorescences pectinate; staminate spikelets with 1 floret .....
17.47 Opizia |
|
Lemmas of the pistillate florets
with awns 30-150 mm long; branches of staminate inflorescences
not pectinate; staminate spikelets with 5-20 florets ..... 17.13 Scleropogon |
|
Lemmas of the
pistillate or bisexual florets with awns 30-150 mm long; bisexual
florets rarely found ..... 17.13 Scleropogon |
|
Lemmas of the bisexual florets
unawned or with awns less than 10 mm long; pistillate florets
not present (30) |
|
Inflorescences
5-15 cm long, apparently a pectinate spike, actually a solitary,
pectinate, spikelike branch ..... 17.42 Ctenium |
|
Inflorescences 1.5-10 cm long,
spikes, spikelike racemes, or panicles, linear or densely cylindrical
to ovoid (31) |
|
Inflorescences
linear, (1.5)4-10 cm long; rachises not concealed by the spikelets
..... 17.20 Tripogon |
|
Inflorescences cylindrical to
ovoid, 1.5-5 cm long; rachises concealed by the spikelets .....
17.01 Fingerhuthia |
|
Inflorescences
simple panicles, sometimes highly contracted, even spikelike
in appearance; spikelike branches not evident (33) |
|
Inflorescences panicles of spikelike
branches, the branches digitately or racemosely arranged on the
rachises (57) |
|
Spikelets usually
with only 1 floret, occasionally with 2-3 florets (34) |
|
Spikelets with more than 1 floret
(39) |
|
Ligules membranous,
hyaline, or coriaceous, sometimes ciliate; lemmas 3-veined (occasionally
appearing 5-veined), usually awned, sometimes unawned or mucronate
(35) |
|
Ligules of hairs; lemmas 1(3)-veined,
unawned, sometimes mucronate (37) |
|
Lemmas and paleas
densely sericeous over the veins and margins, glabrous between
the veins ..... 17.16 Blepharoneuron |
|
Lemmas and paleas glabrous or
variously hairy but not densely sericeous over the veins and
margins (36) |
|
Lemmas usually
awned or mucronate; spikelets usually with 1 floret ..... 17.33 Muhlenbergia |
|
Lemmas unawned or mucronate; spikelets
frequently with 2-3 florets ..... 17.23 Eragrostis |
|
|
Panicles 0.3-4(7.5)
cm long, 3-15 mm wide, spikelike or capitate, 1-8 times longer
than wide; plants annual ..... 17.31 Crypsis |
|
Panicles 1-80 cm long, 2-600 mm
wide, dense to open, if less than 8 cm long, often 10 or more
times longer than wide; plants annual or perennial (38) |
|
Calluses usually
glabrous or almost so; paleas glabrous; fruits falling free of
the lemma and palea ..... 17.30 Sporobolus |
|
Calluses evidently hairy, the
hairs 1/4-7/8 as long as the lemmas; paleas hairy; fruits falling
with the lemma and palea ..... 17.32 Calamovilfa |
|
Lemmas with 1-3 veins (occasionally
with scabrous lines that may be mistaken for additional veins)
(47) |
|
Spikelets unisexual;
plants almost always unisexual, occasionally bisexual (41) |
|
All spikelets with at least 1
bisexual floret (43) |
|
Lemmas 9-11-veined;
glumes 2-7 veined; plants rhizomatous and/or stoloniferous, found
in saline or alkaline soils ..... 17.04 Distichlis |
|
Lemmas 1-6-veined; lower glumes
of the staminate spikelets 1-veined, those of the pistillate
spikelets 1-5-veined; plants stoloniferous or rooting at the
lower nodes, not rhizomatous, not found in saline or alkaline
soils (42) |
|
Plants perennial,
stoloniferous; paleas of the pistillate florets completely surrounding
the ovaries, the intercostal region coriaceous ..... 17.06 Allolepis |
|
Plants annual, rooting at the
lower nodes; paleas of the pistillate florets not completely
surrounding the ovaries, the intercostal region membranous or
hyaline ..... 17.23 Eragrostis |
|
Glumes longer
than the adjacent lemmas (44) |
|
Glumes shorter than the adjacent
lemmas (45) |
|
Glumes exceeded
by the distal florets; spikelets with 3-7 bisexual florets plus
reduced distal florets; lemmas of the bisexual florets 5-7-veined
throughout ..... 17.03 Swallenia |
|
Glumes exceeding the distal florets;
spikelets with 2-4 florets, only the lowest floret bisexual;
lemmas of the bisexual florets 3-veined basally, 5-7-veined distally
..... 17.01 Fingerhuthia |
|
Spikelets ovate-elliptical
to ovate-triangular, 15-50 mm long, 6-16 mm wide; lower florets
sterile, without paleas ..... 17.02 Uniola |
|
Spikelets usually elliptical to
lanceolate, 1-26 mm long, 0.6-9 mm wide; lower florets in each
spikelet bisexual, with paleas (46) |
|
Calluses glabrous
or sparsely pubescent; lemmas (1)3(5)-veined; spikelets 1-26
mm long, 0.6-9 mm wide ..... 17.23 Eragrostis |
|
Calluses densely pubescent; lemmas
5-, 7-, or 9-veined; spikelets 10-16 mm long, 2.5-5 mm wide .....
17.26 Vaseyochloa |
|
Florets unisexual
(48) |
|
At least 1 floret in each spikelet
bisexual (50) |
|
Staminate and
pistillate florets strongly dimorphic; plants unisexual or bisexual,
bisexual plants with unisexual or bisexual spikelets; pistillate
spikelets with 3-5 functional florets and lemma awns (30)50-150
mm long; staminate spikelets with 5-10(20) florets and unawned
or shortly awned (to 3 mm) lemmas ..... 17.13 Scleropogon |
|
Staminate and pistillate florets
similar; plants unisexual; spikelets with 4-60 florets, all or
almost all functional; lemmas 1.5-10.5 mm, unawned, sometimes
mucronate (49) |
|
Plants perennial,
stoloniferous; paleas of the pistillate florets completely surrounding
the ovaries, the intercostal region coriaceous ..... 17.06 Allolepis |
|
Plants annual, rooting at the
lower nodes; paleas of the pistillate florets not completely
surrounding the ovaries, the intercostal region membranous or
hyaline ..... 17.23 Eragrostis |
|
Lemmas, including
the calluses, glabrous or inconspicuously hairy; lemma apices
usually entire, sometimes minutely toothed (51) |
|
Lemma bodies conspicuously hairy over the veins
and/or calluses conspicuously hairy; lemma apices usually with
emarginate, bilobed, or trilobed apices, sometimes entire (52) |
|
Spikelets with
(1)2-60 florets; lemmas unawned, sometimes mucronate; ligules
usually membranous and ciliate or ciliolate, sometimes of hairs
..... 17.23 Eragrostis |
|
Spikelets with 1(2-3) florets; lemmas often awned,
sometimes unawned or mucronate; ligules membranous, sometimes ciliolate,
not ciliate ..... 17.33 Muhlenbergia |
|
Leaf margins
evidently cartilaginous ..... 17.14 Erioneuron |
|
Leaf margins not cartilaginous (53) |
|
Palea keels long
hairy distally, the distal hairs 0.5-2 mm long ..... 17.12 Triplasis |
|
Palea keels glabrous or with hairs less than 0.5
mm long (54) |
|
Lemmas unawned,
the midveins sometimes excurrent up to 0.5 mm (55) |
|
Lemmas awned, the awns 1-7 mm long (56) |
|
Lemmas rounded
to truncate, emarginate to bilobed; all 3 lemma veins often pilose
basally ..... 17.10 Tridens |
|
Lemmas acute, entire or with 3 minute teeth, glabrous
or shortly pubescent on the distal 2/3, the pubescence not confined
to the veins ..... 17.11 Redfieldia |
|
Plants 80-500
cm tall; panicles 35-73 cm long, plumose; lemma margins pilose;
lemma apices bifid, awned from between the teeth; awns about
3 mm long ..... 17.08 Neyraudia |
|
Plants to 2-90 cm tall; panicles 6-30 cm long, not
plumose; lemma margins sparsely pilose; lemma apices 3-4-lobed
or -toothed and 3-awned; central awns 5-7 mm long, lateral awns
6-7 mm long ..... 17.09 Triraphis |
|
Inflorescence
branches 1 or more, if more than 1, arranged in terminal, digitate
clusters, sometimes with additional branches or whorls below
the terminal cluster (58) |
|
Inflorescence branches more than 1, racemosely arranged
on the rachises (71) |
|
Inflorescences
with 1(2) falcate branches (59) |
|
Inflorescences with more than 1 branch or, if only
1, the branch not strongly falcate (60) |
|
Spikelets with
2 well-developed sterile or staminate florets below the bisexual
florets; additional sterile or staminate florets present distal
to the bisexual floret ..... 17.42 Ctenium |
|
Spikelets usually with 1, rarely 2, florets, the
lowest or only floret bisexual ..... 17.43 Microchloa |
|
Plants unisexual
(61) |
|
Plants bisexual, all spikelets with at least 1 bisexual
floret (62) |
|
Staminate spikelets
4-6 mm long, with 2 florets; upper glumes of the pistillate spikelets
indurate, white ..... 17.48 Buchloë |
|
Staminate spikelets 3-4 mm long, with 2 florets;
upper glumes of the pistillate spikelets membranous ..... 17.47 Opizia |
|
Spikelets with
more than 1 bisexual floret (63) |
|
Spikelets usually with only 1 bisexual floret (occasionally
2 in some genera), often with additional staminate, sterile, or
modified florets (67) |
|
Panicle branches
0.4-7 cm long, terminating in a point ..... 17.29 Dactyloctenium |
|
Panicle branches 1-22 cm long, terminating in a
functional or rudimentary spikelet (64) |
|
Disarticulation
eventually below the glumes, initially below the lemmas and caryopses,
the paleas persistent; panicle branches terminating in a rudimentary
spikelet ..... 17.28 Acrachne |
|
Disarticulation above the glumes, usually also below
the florets; panicle branches terminating in a functional spikelet
(65) |
|
Lemmas 3-awned,
the central awns 8-12 mm long ..... 17.38 Trichloris |
|
Lemmas not 3-awned (66) |
|
Lemmas usually with hairs over the veins, at least basally, the
apices often toothed, sometimes mucronate or awned ..... 17.19 Leptochloa |
|
Lemmas glabrous, the apices entire, neither mucronate
nor awned ..... 17.27 Eleusine |
|
Spikelets usually
without sterile or modified florets; lemmas unawned ..... 17.44 Cynodon |
|
Spikelets with 1 or more sterile florets distal
to the bisexual floret; lemmas of the bisexual florets often awned
(68) |
|
Lowest lemmas
in the spikelets 3-awned, the central awns 8-12 mm long, the
lateral awns 0.5-12 mm long ..... 17.38 Trichloris |
|
Lowest lemmas in the spikelets usually unawned or
with a single awn, if 3-awned, the lateral awns less than 0.5 mm
long (69) |
|
Spikelets dorsally
compressed ..... 17.37 Enteropogon |
|
Spikelets laterally compressed or terete (70) |
|
Upper glumes
truncate or bilobed; lowest lemmas unawned or with an awn to
1.2 mm long ..... 17.36 Eustachys |
|
Upper glumes acute to acuminate; lowest lemmas usually
awned, the awns to 37 mm long ..... 17.35 Chloris |
|
All spikelets
unisexual (72) |
|
All spikelets with at least 1 bisexual floret (73) |
|
Staminate spikelets
4-6 mm long, the anthers 2.5-3 mm long; pistillate spikelets
with 1 unawned or shortly awned floret; widespread species of
the central plains ..... 17.48 Buchloë |
|
Staminate spikelets 3-4 mm long, the anthers 2-2.5
mm long; pistillate lemmas with awns 3.4-6.8 mm long; in the Flora region,
known only as an occasional escape from lawns and experimental
plots in Florida ..... 17.47 Opizia |
|
Inflorescence
branches woody, terminating in hard, sharp points ..... 17.24 Cladoraphis |
|
Inflorescence branches not particularly stiff or
rigid, terminating in spikelets or points (74) |
|
Spikelets with
more than 1 bisexual floret, sometimes also with reduced florets
(75) |
|
Spikelets with 1 bisexual floret, sometimes with
sterile, rudimentary, or modified florets distal to the bisexual
floret (79) |
|
Lemmas 7-11-veined,
mucronate, the mucros 0.1-0.3 mm long; not established in the Flora region
..... 17.05 Aeluropus |
|
Lemmas 3-veined, unawned, mucronate, or awned, the
awns often much more than 1 mm long; established in the Flora region
(76) |
|
Lower glumes
exceeding the lowest lemmas, sometimes exceeding the distal lemmas
..... 17.21 Trichoneura |
|
Lower glumes not or only slightly exceeding the
lowest lemmas (77) |
|
Inflorescences
with 50 or more closely spaced, arcuate, tardily deciduous branches
..... 17.25 Pogonarthria |
|
Inflorescences with 2-120 straight, non-disarticulating
branches (78) |
|
Spikelets with
(2)3-12(20) bisexual florets ..... 17.19 Leptochloa |
|
Spikelets with 2-4 florets, but only the lowest
1(2) florets bisexual ..... 17.41 Gymnopogon |
|
Functional spikelets
with sterile, rudimentary, or modified florets distal to the
bisexual floret (80) |
|
Functional spikelets with only 1 floret, lacking
sterile, rudimentary, or modified florets (82) |
|
Spikelets widely
spaced to slightly imbricate, appressed to the branch axes .....
17.41 Gymnopogon |
|
Spikelets densely imbricate, varying from appressed
to strongly divergent (81) |
|
Inflorescence
branches usually solitary at each node (sometimes only 1 per
panicle); spikelets laterally compressed or terete ..... 17.46 Bouteloua |
|
Inflorescence branches usually more than 1 at the
lower nodes; spikelets dorsally compressed ..... 17.37 Enteropogon |
|
Spikelets distant
to slightly imbricate, appressed to the branches; branches strongly
divergent (83) |
|
Spikelets clearly imbricate, appressed to strongly
divergent; branches appressed to strongly divergent (84) |
|
Blades with thick,
white margins and a well-developed midrib ..... 17.40 Schedonnardus |
|
Blades lacking both thick, white margins and well-developed
midribs ..... 17.41 Gymnopogon |
|
Spikelets laterally
compressed, appressed to divergent ..... 17.45 Spartina |
|
Spikelets dorsally compressed, appressed ..... 17.39 Willkommia |