| Mary E. Barkworth |
Plants usually perennial. Culms
7-600 cm, annual, not woody, often reddish or purple, particularly at the nodes,
often branched above the base. Sheaths open; ligules usually scarious
to membranous, ciliate or not; blades mostly well-developed, leaves subtending
an inflorescence or an inflorescence unit
often with reduced blades. Photosynthetic pathway NADP-ME; bundle
sheaths single. Inflorescences terminal, frequently on both the culms
and their branches, sometimes also axillary, usually of 1-many spikelike branches,
these in digitate clusters of 1-13+ on a peduncle or attached, directly or indirectly,
to elongate rachises, often partially to almost completely enclosed by the subtending
leaf sheath at maturity, in some taxa axillary inflorescences composed of multiple-stalked
pedunculate clusters of inflorescence branches subtended by a modified leaf;
disarticulation usually in the branch axes beneath the sessile florets,
the dispersal unit being a sessile floret, the internode to the next sessile
floret, the pedicel, and the pedicellate spikelet (branches with disarticulating
axes are termed rames in the following accounts),
sometimes beneath the glumes, the branch axes remaining intact. Spikelets
in unequally pedicellate pairs, sessile-pedicellate pairs, or triplets, or apparently
solitary and sessile, pedicellate spikelets and sometimes the pedicels reduced
or absent, triplets usually with 1 sessile and 2 pedicellate spikelets, terminal
spikelet units on the branches often with 2 pedicellate spikelets even if the
others have only 1 (all spikelet units with 2 sessile and 1 pedicellate spikelets
in Polytrias). Spikelet pairs or triplets
homogamous (spikelets in the unit sexually
alike) or heterogamous (spikelets in the
unit sexually dissimilar); spikelets of unequally pedicellate pairs usually
homogamous and homomorphic; spikelets in
sessile-pedicellate pairs or triplets usually heterogamous and heteromorphic;
sessile spikelets usually bisexual; pedicellate spikelets usually
smaller than the sessile spikelets, often staminate or sterile, sometimes absent.
Spikelets usually with 2 florets (1 in Polytrias).
Glumes exceeding and usually concealing the florets (excluding the awns),
rounded or dorsally compressed, usually tougher than the lemmas; lower florets
in bisexual or pistillate spikelets sterile or staminate, often reduced to a
hyaline scale; upper florets bisexual or pistillate, lemmas often hyaline,
sometimes with an awn that exceeds the glumes; lodicules cuneate; anthers
usually 3. Pedicels free or fused to the rachis internodes. Pedicellate
spikelets variable, sometimes similar to the sessile spikelets, sometimes
differing in sexuality and shape, sometimes missing. x = usually 9 or
10, or possibly 5 with 9 and 10 reflecting ancient polyploidy.
The tribe Andropogoneae includes about 87 genera and 1060 species, of
which 31 genera and 102 species have been found in the Flora region;
some of these have not become established. The tribe is common in tropical and
subtropical regions, particularly in areas with significant summer rains, such
as the central plains of North America. Two of the grasses that used to dominate
the prairies of central North America, Andropogon
gerardii and Schizachyrium scoparium (Big and Little Bluestem, respectively), are member of the Andropogoneae.
The reddish-purplish coloration that characterizes the culms and leaves of many
Andropogoneae gives a striking aspect to grasslands (and lawns) dominated
by its members.
Members of the Andropogoneae differ from those of Paniceae in the reduced lemmas and paleas of their florets and, usually, in their
paired, unequally pedicellate spikelets, disarticulating inflorescence branches
(rames), and the manner in which these branches are aggregated into inflorescences.
Unequally pedicellate spikelet pairs are found in many other tribes, but they
are more common, and the pedicels more strikingly unequal in length, in the
Andropogoneae. Recent molecular work supports recognition of the tribe
with one modification of its traditional limits, the incorporation of Arundinella and Tristachya (Kellogg 2000). There
is less agreement on the tribes internal structure and its relationship to the
Paniceae (Clayton and Renvoize 1986; Kellogg 2000; Spangler 2000; Guissani
et al. 2001).
Inflorescence Structures
Describing inflorescence structures in the Andropogoneae is not simple.
There is a basic pattern, but its many modifications have resulted in great
structural diversity. The following paragraphs provide an overview of this diversity
and explain the words and phrases used in describing it.
Spikelets
Members of the Andropogoneae, like those of the Paniceae, generally have two florets per spikelet, the lower floret usually being
reduced in size and sterile or staminate, and the upper floret bisexual. Despite this similarity, spikelets of the two tribes are easy to distinguish.
In the Paniceae, the lowest glume is usually much shorter than the floret,
and the upper florets usually have lemmas that are thicker and tougher than
the glumes and lower lemmas. In the Andropogoneae, the glumes usually
exceed and enclose both florets, and are thicker and tougher than the lemmas.
The florets of the Andropogoneae tribe contrast strongly with the glumes,
having hyaline or thinly membranous lemma bodies and hyaline paleas, or, in
many cases, no palea. They are almost always completely concealed by the glumes,
except that the upper floret often has an awn that projects beyond the glumes.
In some Andropogoneae, the glumes are merely thickly membranous, but
most genera have coriaceous or indurate glumes. The lower glumes are sometimes
tougher and larger than the upper glumes, and may even conceal the upper glumes
as, for example, in Heteropogon. In such
genera, the lower glumes may be mistaken for lemmas. In dioecious species, or
monoecious species with strongly differentiated staminate and pistillate spikelets,
the staminate spikelets usually have softer glumes than the pistillate spikelets.
Spikelet Units
The basic element of the inflorescence structure in the Andropogoneae
is the spikelet unit. These units usually consist of pairs of spikelets,
one sessile and one pedicellate (e.g., Saccharum
bengalense), but they may consist of a pair of unequally pedicellate
spikelets (e.g., Miscanthus sacchariflorus) or of three spikelets (e.g., Chysopogon
fulvus). If there are three spikelets in the unit, one is usually
sessile and the other two pedicellate, but a few genera, such as Polytrias, have two sessile spikelets and one pedicellate spikelet.
Unequally pedicellate spikelet pairs or triplets are found in other tribes,
but in the Andropogoneae they usually differ in size, shape, and sexuality.
Spikelet units with spikelets that differ in their sexuality are described as
heterogamous; those with sexually similar spikelets are said to be homogamous.
Spikelet units with morphologically dissimilar spikelets are heteromorphic (e.g., Andropogon longiberbis);
those with morphologically similar spikelets are homomorphic (e.g., Chrysopogon zizanioides). In most Andropogoneae, the spikelet units are heterogamous and
heteromorphic. The sessile spikelets usually contain a bisexual or pistillate
floret, and often exhibit features such as awns and calluses that are related
to seed dispersal and establishment (Peart 1984); the pedicellate spikelets
are usually staminate, sterile, vestigial, or even absent. In some genera the
situation is reversed, the pedicellate spikelets being bisexual or pistillate,
and the sessile spikelets staminate or sterile. Sterile and staminate spikelets
are sometimes morphologically similar to the pistillate or bisexual spikelets,
but usually lack the features associated with seed dispersal and establishment.
A few genera have no staminate or sterile spikelets, merely
empty pedicels associated with the bisexual sessile spikelets, as in Sorghastrum
or even, as in Arthraxon, with only a stump
where the pedicel and its spikelet would be.
Inflorescence Structure
Further complexity is introduced to the Andropogoneae inflorescence structure
by the manner in which the spikelet units are aggregated and the mode of disarticulation.
Three patterns can be identified. The simplest pattern consists of inflorescences
similar to those common in other tribes, in which neither the rachis nor the
inflorescence branches break up at maturity. Genera with such inflorescences
(e.g., Miscanthus and Imperata) have unequally pedicellate spikelets, and disarticulation is below the
glumes. Such inflorescences are, however, in the minority within the Andropogoneae.
A more common situation is for the spikelets to be in sessile-pedicellate
pairs and disarticulation to be in the branch axes, immediately below the attachment
of the sessile spikelets. The resulting dispersal unit consists of the spikelet
pair plus the internode that extends from the sessile spikelet to the next most
distal sessile spikelet. These disarticulating inflorescence branches, termed
rames in this Flora, form the basic unit of the typical Andropogoneae
inflorescence. In other publications, the rames are often called racemes,
a word that is restricted in this Flora to an entire infloresence, not
just an inflorescence branch.
Rames are usually composed of several spikelet units, but sometimes
of only one. The spikelets may be evenly distributed, or the base of the rame
axis may be naked. Individual plants may bear few to many rames, and the rames
themselves may be aggregated in a wide array of primary and secondary arrangements;
they may also be branched.
One or more rames may be borne on a single stalk. If this stalk is attached
to a rachis, the unit formed by the stalk and its rame(s) constitutes an inflorescence
branch. Such a pattern is seen, for example, in Sorghum
halepense and Bothriochloa
bladhii. A more common situation is for one or more rames to be attached
digitately to a common stalk, the peduncle. This peduncle may terminate a culm
(as in Dichanthium annulatum
or Elionurus) or be axillary to a subtending
leaf (as in Andropogon hallii
and Hemarthria altissima).
Each peduncle and its associated rame(s) constitutes an inflorescence unit.
False panicles represent a further level of complexity.
In these, the inflorescence units terminate rays, each of which has a
prophyll, a 2-veined structure, in its axil. Several rays may develop
within the axil of a single leaf sheath, and rays may themselves give rise to
subtending leaves with multiple rays in their axils. The result is a complex,
tiered inflorescence in which only the ultimate units are easily described.
Such inflorescences are found, for example, in Andropogon
glomeratus and Cymbopogon
citratus. Fortunately, identification of the Andropogoneae
does not require analyzing false panicles, merely their ultimate inflorescence
units.
In another inflorescence pattern, the rame axes are thick and
the pedicels are either closely appressed or even fused to the rame axes. In
these genera, the pedicellate spikelets are often highly reduced or absent.
Pistillate rames of Tripsacum and
wild taxa of Zea represent an extreme example
of this pattern. In these genera, the sessile spikelets are completely embedded
in the rame axes, the lower glumes being indurate and completely concealing
the florets. Less extreme examples are seen in Coelorachis
and Hackelochloa.
1 |
Leaves
smelling of lemon oil or citronella, the sheaths without glandular depressions
on the keel; plants perennial, not reaching reproductive maturity in the
Flora region when grown outdoors .....
26.16 Cymbopogon |
Leaves usually not aromatic or, if aromatic
and smelling of citronella, the sheaths with glandular depressions along
the keel and plants annual; plants reaching reproductive maturity in the
Flora region (2) |
|
All spikelets unisexual,
the pistillate and staminate spikelets in separate inflorescences or the
pistillate spikelets below the staminate spikelets in the same inflorescence
(3) |
|
Some spikelets bisexual (usually the
sessile or more shortly pedicellate spikelet of each spikelet pair or
triplet) (5) |
|
Pistillate spikelets
completely concealed within a hard, globose, beadlike structure (a modified
leaf sheath) from which the staminate rames protrude ..... 26.30 Coix |
|
Pistillate spikelets exposed or enclosed
by 1 or more subtending leaf sheaths and a hyaline prophyll; staminate
spikelets either distal on the same branch or in a separate inflorescence
on the same plant (4) |
|
Staminate and pistillate
spikelets in the same inflorescence and on the same branch, the staminate
spikelets distal to the pistillate spikelets ..... 26.28 Tripsacum |
|
Staminate and pistillate inflorescences
usually separate; staminate inflorescences terminal on the culms and branches;
pistillate inflorescences terminal on axillary peduncles, sometimes aggregated
in false panicles ..... 26.29 Zea |
|
Spikelets apparently
solitary and sessile, the pedicellate spikelets absent; pedicels absent
or present (6) |
|
Spikelets in sessile-pedicellate orunequally
pedicellate pairs or triplets, the pedicellate spikelets often smaller
than the sessile spikelets, sometimes rudimentary (8) |
|
Culms decumbent, scrambling;
leaf blades ovate to ovate-lanceolate; pedicels absent or shorter than
3 mm ..... 26.18 Arthraxon |
|
Culms erect; leaf blades lanceolate to
linear-lanceolate; pedicels always present, usually longer than 3 mm (7) |
|
Inflorescences terminal
and axillary, composed of digitate clusters of 1-13 rames on a common
peduncle; peduncles subtended by, and often partially included in, a modified
leaf ..... 26.15 Andropogon |
|
Inflorescences terminal, with elongate
rachises and brancheswith several to many rames; peduncles and branches
not subtended by a modified leaf ..... 26.09 Sorghastrum |
|
Pedicels strongly appressed
or fused to the thick rame axes, or the rames with only 1 spikelet unit,
this a triplet with 2 unequally pedicellate spikelets; bisexual spikelets
usually unawned; inflorescences of rames (9) |
|
Pedicels free; rame or branch internodes
slender, sometimes thickened distally; bisexual spikelets usually awned;
inflorescences of rames with the spikelets in sessile-pedicellate pairs
or of non-disarticulating branches with the spikeletsin unequally pedicellate
pairs (17) |
|
Lower glumes of the sessile
spikelets rugose, pitted, tuberculate, or alveolate or the keels winged
or with spinelike projections at the base (10) |
|
Lower glumes of the sessile spikelets
smooth or scabrous, not sculptured, the keels without spinelike projections
(13) |
|
Keels
of the lower glumes with spinelike projections on the base, sometimes
winged distally, the surface between the keels smooth; spikelets unawned
..... 26.25 Eremochloa |
|
Keels of the lower glumes winged throughout
or not winged, the surface between the keels rough, rugose, pitted, tuberculate,
or alveolate; spikelets unawned or awned (11) |
|
Sessile spikelets awned
..... 26.13 Ischaemum |
|
Sessile spikelets unawned (12) |
|
Plants perennial; sessile
spikelets ovate, the lower glumes smooth, rugose, or pitted ..... 26.24
Coelorachis |
|
Plants annual; sessile spikelets hemispherical,
the lower glumes alveolate ..... 26.27 Hackelochloa |
|
Inflorescences false
panicles; individual rames to 1 cm long, with 1 spikelet unit; spikelet
units composed of 1 sessile and 2 unequally pedicellate and dissimilar
spikelets ..... 26.14 Apluda |
|
Inflorescences usually solitary rames,
sometimes with 2 rames in a digitate cluster; individual rames 2-15 cm
long, with more than 1 spikelet unit; spikelet units composed of sessile
pedicellate pairs, the pedicellate spikelets often rudimentary or absent
(14) |
|
Pedicels appressed,
but not fused, to the rame axes (15) |
|
Pedicels at least partially fused to
the rame axes (16) |
|
Pedicellate spikelets
1-3 mm long ..... 26.24 Coelorachis |
|
Pedicellate spikelets 4-8 mm long .....
26.22 Elionurus |
|
Plants perennial; sheaths
mostly glabrous, sparsely ciliate basally ..... 26.23 Hemarthria |
|
Plants annual; sheaths with stiff, papillose-based
hairs 1-3 mm long ..... 26.26 Rottboellia |
|
All spikelet units homogamous,
frequently also homomorphic (18) |
|
All or most spikelet unitsheterogamous,
usually also heteromorphic, sometimes the proximal units on the rames
or racemes homomorphic and homogamous (24) |
|
Terminal inflorescences
a single rame or a digitate or subdigitate cluster of rames (19) |
|
Terminal inflorescences with elongated
rachises (20) |
|
Terminal inflorescences
a digitate or subdigitate cluster of (1)2-6 rames; rames 3-7 cm long .....
26.06 Microstegium |
|
Terminal inflorescences solitary rames;
rames 2-3 cm long ..... 26.05 Polytrias |
|
Spikelets in unequally
pedicellate pairs; disarticulation below the glumes, the branches remaining
intact at maturity (21) |
|
Spikelets in sessile-pedicellate pairs
or triplets; disarticulation in the rames, below the sessile spikelets
(22) |
|
Spikelets usually awned;
inflorescence branches usually 7-35 cm long ..... 26.03 Miscanthus |
|
Spikelets unawned; inflorescence branches
1-7 cm long ..... 26.04 Imperata |
|
Culms to 100 cm tall,
often decumbent and straggling; terminal inflorescences composed of 2-6
subdigitately to racemosely arranged rames ..... 26.06 Microstegium |
|
Culms 40-600 cm tall, erect; terminal
inflorescences panicles, with more than 6 primary branches; branches usually
with 2 or more rames (23) |
|
Panicle branches alternate,
with multiple rames; rames with more than 5 spikelet units ..... 26.02
Saccharum |
|
Panicle branches subverticillate, with
1-3 rames; rames with 2-5 spikelet units ..... 26.01 Spodiopogon |
|
Terminal inflorescences
with elongated rachises (25) |
|
Terminal inflorescences and individual
inflorescence units without elongated rachises, composed of 1-13 rames,
or a raceme in which disarticulation occurs below the pedicellate spikelets
(27) |
|
Rame internodes and
pedicels with a translucent median line ..... 26.12 Bothriochloa |
|
Rame internodes and pedicels without
a translucent median line (26) |
|
Sessile spikelets terete
or laterally compressed, calluses usually sharp, sometimes blunt .....
26.10 Chrysopogon |
|
Sessile spikelets dorsally compressed,
calluses usually blunt, sometimes sharp ..... 26.08 Sorghum |
|
Inflorescences composed
of rames, disarticulation being in the rame axes; rame internodes and
pedicels with a translucent median groove ..... 26.12 Bothriochloa |
|
Inflorescences composed of rames, with
disarticulation in the axes or a spikelike raceme (occasionally of 2 subdigitate
spikelike branches) with disarticulation below the pedicellate spiklets;
inflorescence internodes and pedicels without a translucent median groove
(28) |
|
All spikelet units in
the inflorescence heterogamous (29) |
|
Basal spikelet units on each rame homomorphic
and homogamous, sterile or staminate, unawned (31) |
|
Disarticulation occurring
below the pedicellate spikelets, not in the inflorescence axes; pedicellate
spikelets bisexual and awned, awns (4)6-15 cm long, pilose on the column,
the hairs 1-2 mm long; sessile or subsessile spikelets staminate or sterile,
unawned ..... 26.07 Trachypogon |
|
Disarticulation occurring below the sessile
spikelets, in the inflorescence axes; pedicellate spikelets staminate,
sterile, rudimentary, or absent, unawned or with awns to 6 mm long; sessile
spikelets bisexual or pistillate, awned (30) |
|
Rames usually solitary
on the peduncles, occasionally 2; rame internodes cupulate or fimbriate
distally; lower glumes of the sessile spikelets veined between the keels
..... 26.17 Schizachyrium |
|
Rames usually 2-13 on the peduncles,
occasionally solitary; rame internodes neither fimbriate nor cupulate
distally; lower glumes of the sessile spikelets usually without veins
between the keels ..... 26.15 Andropogon |
|
Awns 5-15 cm long; rames
with 3-10 homogamous spikelet units ..... 26.20 Heteropogon |
|
Awns 1-5(19) cm long; rames with 1-2
homogamous spikelet units (32) |
|
Inflorescences terminal
on the culms, axillary inflorescences not present or few in number .....
26.11 Dichanthium |
|
Inflorescences terminal and axillary,
axillary inflorescences numerous (33) |
|
Homogamous spikelets
distinctive, forming an involucre around the rame bases ..... 26.21 Themeda |
|
Homogamous spikelets not distinctive,
not forming an involucre around the rame bases ..... 26.19 Hyparrhenia |