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Habit Plants annual or perennial; monoecious, inflorescences unisexual or bisexual with the pistillate spikelets basal and the staminate spikelets distal. Plants annual or perennial; synoecious, monoecious, or dioecious; primarily herbaceous, habit varied.
Culms

(0.2)0.5-6 m tall, 1-5 cm thick, solitary or several to many together, monopodial, often branching (branches frequently highly reduced and hidden within the subtending leaf sheath), usually succulent when young, becoming woody with age;

lower nodes with prop roots;

internodes pith-filled.

annual, usually solid, sometimes somewhat woody, sometimes decumbent, often branched above the base.

Leaves

not aromatic, cauline, distichous;

sheaths open;

auricles sometimes present;

ligules membranous, shortly ciliate;

blades 2-12 cm wide, flat.

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;

blades sometimes pseudopetiolate;

mesophyll radiate or non-radiate;

adaxial palisade layer absent;

fusoid cells usually absent;

arm cells usually absent;

kranz anatomy absent or present;

midribs usually 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(-4), 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 florets well-developed, reduced, or absent;

lodicules usually 2, sometimes absent, cuneate, free, fleshy, usually glabrous;

anthers 1-3;

ovaries usually glabrous;

haustorial synergids absent;

style branches 2, free and close or fused at the base.

Caryopses

subspherical to dorsally compressed;

hila round;

embryos about 2/3 as long as the 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).

Pistillate

or partially pistillate inflorescences terminal on axillary branches;

staminate inflorescences (tassels) paniculate, of 1-many branches or rames, sometimes with secondary and tertiary branching.;

pistillate spikelets solitary, sessile, with 1 floret;

pedicels and pedicellate spikelets suppressed;

lower glumes exceeding the floret, indurate on the central, exposed portion, hyaline on the margins, concealing the caryopses at maturity.;

pistillate spikelets in subsessile pairs, each spikelet with 1 functional floret;

glumes shorter than the spikelets, indurate basally, hyaline distally;

lower florets suppressed.

Wild

TAXA: Pistillate inflorescences solitary, distichous rames (ears), these often tightly clustered in false panicles, each usually wholly or partially enclosed by a thin prophyll and an equally thin bladeless leaf sheath;

rames composed of 5-15 spikelets in 2 ranks;

disarticulation in the rame axes, dispersal units (fruitcases) consisting of an indurate, shiny rame segment and its embedded spikelet.;

wild TAXA: Staminate panicles terminal on the culms and primary branches, sometimes also on the secondary branches and pistillate inflorescences;

rames distichous, similar in thickness and structure, axes disarticulating below the sessile spikelets after pollination, abscission layers evident.

Domesticated

TAXON: Pistillate inflorescences solitary, polystichous spikes (ears) terminating reduced branches, each spike surrounded by several to many, often bladeless leaf sheaths and a prophyll (husks), with 60-1000+ spikelets in 8-24 rows, neither spikes nor spikelets disarticulating at maturity.; domesticated TAXON: Staminate panicles terminal on the culms, central axes always much thicker than the lateral branches and irregularly polystichous, lateral branches distichous to more or less polystichous, not disarticulating, without abscission layers below the sessile spikelets.

All

TAXA: lemmas and paleas hyaline, unawned;

lodicules absent;

ovaries glabrous;

styles (silks) 2, appearing solitary by being fused except at the very tip, filamentous, sides stigmatic.; all TAXA: Staminate spikelets in sessile-pedicellate pairs, each with 2 staminate florets;

glumes membranous to chartaceous, stiff to flexible, sometimes with a pair of winged keels, 5-14(28)-veined, acute;

lemmas and paleas hyaline;

lodicules 2;

anthers 3.

x

= 10.

Zea

Poaceae subfam. panicoideae

Distribution
from FNA
AL; AR; AZ; CA; CO; CT; FL; GA; IA; ID; IL; IN; KS; KY; LA; MA; MD; ME; MI; MN; MO; MS; MT; NC; NE; NH; NJ; NM; NV; NY; OH; OR; PA; RI; SC; TN; TX; UT; VA; VT; WA; WI; WV; WY; PR; ON; QC; Virgin Islands
[BONAP county map]
Discussion

Zea is an American genus of five species, four of which are native to montane Mexico and Central America. The fifth species, Z. nicaraguensis H.H. litis & B.F. Benz, is said to have been ubiquitous at one time in coastal Pacific Nicaragua, but is now known from only four or five small populations near sea level in seasonally flooded savannahs and riverine forests inland from the Bay of Fonseca, Nicaragua.

The often weedy, wild taxa, known as 'teosinte', are used in plant breeding as well as in developmental and evolutionary studies. The genus has also been the focus of physiological and genetic research, mostly involving the domesticated taxon, Zea mays subsp. mays. Examples of such work include Barbara McClintock's Nobel prize-winning discovery that genes can "jump" from one chromosome to another, and recent work on the evolution of tassel morphology (e.g., Westerbergh and Doebley 2002).

Zea mays subsp. mays, the most widespread taxon in the genus, was first domesticated about 7,000 years ago and soon became widely planted in the Americas. It is now grown in all warmer parts of the world and is the world's third most important crop plant. No other American grass has such agricultural importance.

In the Flora region, Zea mays subsp. mays is widely grown commercially; Z. luxurians is sometimes grown for forage; while Z. diploperennis and Z. perennis, and the other subspecies of Z. mays, are almost completely confined to research plantings.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

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 (Sanchez-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 (Gomez-Martinez and Culham 2000; Giussanni et al. 2001; Barber et al. 2002).

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Key
1. Pistillate inflorescences terete, with 2+ rows of paired spikelets, each spikelet with a functional floret, hence the spikelets in 4+ rank; staminate spikelets of wild taxa only slightly imbricate; lower glumes of the staminate spikelets flexible and translucent, loosely enclosing the upper glumes before anthesis, rounded on the back, the lateral veins not more prominent than those between, never forming winged keels; plants annual (sect. Zea)
Z. mays
1. Pistillate inflorescences somewhat flattened, with 2 rows of solitary spikelets, hence the spikelets appearing 2-ranked; staminate inflorescences with densely imbricate spikelets; lower glumes of the staminate spikelets stiff, not translucent, strongly enclosing the upper glumes before anthesis, with more or less flat backs, the lateral veins evidently more prominent than those between, keeled and winged distally; plants annual or perennial (sect. Luxuriantes).
→ 2
2. Plants annual; lower glumes of the staminate spikelets narrowly winged; pistillate inflorescence units 1-many per node, usually enclosed by their subtending leaf sheaths, occasionally with 1-2 rames on naked peduncles that exceed the subtending leaf sheaths
Z. luxurians
2. Plants perennial, rhizomatous; lower glumes of the staminate spikelets strongly winged; pistillate inflorescence units 1-4(5) per node, almost always exceeding the subtending leaf sheaths.
→ 3
3. Rhizomes to 15 cm long, with internodes 0.2-0.6 cm long, often forming scaly, tuberous short shoots; culms to 3.5 m tall and 3 cm thick; leaf blades to 40 cm long and 4-5.5 cm wide
Z. diploperennis
3. Rhizomes to 40 cm long or more, with internodes 1-6 cm long, lacking tuberlike shoots; culms to 2.5 m tall and 1.5-2 cm thick; leaf blades often to 65(80) cm long and 2-4.5 cm wide
Z. perennis
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
Gynerieae
1. 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
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 the inflorescence branches
Paniceae
2. 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
Andropogoneae
Source FNA vol. 25, p. 696. Author: Hugh H. litis;. FNA vol. 25, p. 351. Author: Grass Phylogeny Working Group;.
Parent taxa Poaceae > subfam. Panicoideae > tribe Andropogoneae Poaceae
Subordinate taxa
Z. diploperennis, Z. luxurians, Z. mays, Z. perennis
Name authority L. Link
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