| Grass Phylogeny Working Group Lynn G. Clark |
Plants annual or perennial;
synoecious, monoecious, or dioecious; primarily herbaceous, habit varied. Culms annual,
usually solid, sometimes somewhat woody, sometimes decumbent, often
branched above the base. Leaves distichous; sheaths usually
open; auricles usually absent; abaxial ligules usually
absent, occasionally present as a line of hairs; adaxial ligules membranous,
sometimes also ciliate, or of hairs, sometimes absent; bladessometimes
pseudopetiolate; mesophyll radiate or non-radiate; adaxial
palisade layer absent; fusoid cells usually absent; arm
cells usually absent; kranz anatomy absent or present; midribsusually
simple, rarely complex; adaxial bulliform cells present; stomata with
triangular or dome-shaped subsidiary cells; bicellular microhairs usually
present, with a long, narrow distal cell; papillae absent or
present. Inflorescences ebracteate (Paniceae) or bracteate
(most Andropogoneae) panicles, racemes, spikes, or complex arrangements
of rames (in the Andropogoneae), usually bisexual, sometimes
unisexual; disarticulation usually below the glumes, frequently
in the secondary and higher order axes of the inflorescences. Spikelets bisexual
or unisexual, frequently paired or in triplets, the members of each
unit usually with pedicels of different lengths or 1 spikelet sessile. Glumes usually
2, equal or unequal, shorter or longer than the adjacent florets, sometimes
exceeding the distal florets; florets 2, usually dorsally compressed,
sometimes terete or laterally compressed; lower florets sterile
or staminate, frequently reduced to a lemma; upper florets usually
bisexual; lemmas hyaline to coriaceous, lacking uncinate hairs,
often terminally awned; awns single; paleas of bisexual floretswell-developed,
reduced, or absent; lodicules usually 2, sometimes absent, cuneate,
free, fleshy, usually glabrous; anthers 1-3; ovaries usually
glabrous; haustorial synergids absent; style branches2,
free and close or fused at the base. Caryopses: hila usually
punctate; endosperm hard, without lipid; starch grains simple; embryos large
in relation to the caryopses, usually waisted; epiblasts usually
absent; scutellar cleft present; mesocotyl internode elongated; embryonic
leaf margins usually overlapping, rarely just meeting. x =
5, (7), 9, 10, (12), (14).
The subfamily Panicoideae is most abundant in tropical and subtropical
regions, particularly mesic portions of such regions, but several species
grow in temperate regions of the world. Within the Flora region,
the Panicoideae are represented by 59 genera and 364 species. They
are most abundant in the eastern United States (Barkworth and Capels 2000).
Photosynthesis may be either C3 or C4.
All three pathways are found in the subfamily, but the PCK and NAD-ME variants
appear to have evolved only once, while the NADP-ME pathways seems to have
evolved several different times (Giussani et al. 2001).
The Panicoideae were first recognized as a distinct unit by Brown
(1814), earlier than any of the other subfamilial taxa of the Poaceae.
Its early recognition is undoubtedly attributable to its distinctive spikelets.
Recognition of the tribe Gynerieae is
recent (Sánchez-Ken and Clark 2001) and its placement in the Panicoideae,
rather than the Centothecoideae,
should be regarded as tentative.
Spikelets with two florets are found in many other subfamilies, but rarely
do they follow the pattern of the lower floret being sterile or staminate
and the upper floret bisexual. Development of unisexual florets within
the Panicoideae appears to be consistent across the subfamily (LeRoux
and Kellogg 1999), but differs from that in the Ehrhartoideae (Zaitcheck
et al. 2000).
The Paniceae and Andropogoneae have
their conventional interpretation in this Flora, so far as the North
American taxa are concerned. Molecular studies, however, while strongly
supporting the monophyly of the Andropogoneae, show the Paniceae to
be paraphyletic, with two distinct clades. In one of these clades, most
taxa have a chromosome base number of x = 9, but some have x =
10, and the taxa are pan-tropical in origin. The taxa in the other clade,
with one exception, have a chromosome base number of x = 10 and
are American in origin. This latter clade is sister to the Andropogoneae,
which also have a chromosome base number of x =
10 (Gómez-Martínez and Culham 2000; Giussanni et al. 2001;
Barber et al. 2002).
1 |
Blades
of leaves on the lower 1/2 of the culms disarticulating from
the sheaths; plants 2-15 m tall, unisexual, without axillary
inflorescences; blades with midribs 5-15 mm wide ..... 24. Gynerieae |
Blades of most or all cauline
leaves remaining attached to the sheaths; plants 0.05-6 m tall,
usually bisexual, sometimes with unisexual inflorescences, often
with axillary inflorescences; blades with midribs 0.2-5 mm wide
(2) |
|
Glumes usually
conspicuously unequal; lower glumes usually greatly exceeded
by the upper florets; upper glumes from subequal to longer than
the distal florets; lemmas of the upper florets usually coriaceous
to indurate; disarticulation usually beneath the glumes, not
in the axes of theinflorescence branches ..... 25. Paniceae |
|
Glumes usually subequal, usually
exceeding and concealing the florets; lemmas of the upper florets
hyaline to membranous; disarticulation frequently in the axes
of the inflorescence branches ..... 26. Andropogoneae |