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Manual of Grasses for North America

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Grasses are the world’s most important plants. They are the dominant species over large parts of the earth’s land surface, a fact that is reflected in the many different words that exist for grasslands, words such as prairie, veldt, palouse, and pampas to mention just a few. As a group, grasses are of major ecological importance, as soil binders and providers of shelter and food for wild animals, both large and small. Some grasses, such as wheat, rice, corn, barley, rye, tef, and sugar cane are major sources of calories for humans and their livestock; others, primarily bamboos, are used for construction, tools, paper, and fabric. More recently, the seed catalogs that tantalize gardeners each winter have borne witness to an increasing appreciation of the aesthetic value of grasses.

The Manual of Grasses for North America is designed as a successor to the classic volume by Hitchcock and Chase. It reflects current taxonomic thought and includes keys, illustrations, and distribution maps for the nearly 900 native and 400 introduced species that have been found in North America north of Mexico. In addition, it presents keys and illustrations for several species that are known only in cultivation or are of major agricultural significance, either as progenitors of bread wheat and corn or as a major threat to North American agriculture because of their ability to hybridize with crop species. The Manual of Grasses for North America is a major reference work for grasses that will retain its value for many years.

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1. PHAROIDEAE L.G. Clark & Judz.

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The Pharoideae has one tribe, the Phareae, three genera, and twelve species. It is pantropical. In the Americas, it is represented by one genus, Pharus, that extends from Florida to Uruguay and Argentina. The Pharoideae is a basal lineage of the Poaceae, and the first subfamily in which an adaxial ligule and true spikelets are found.

Pl per; rhz, smt csp or stln; monoecious. Clm ann, 10–300 cm, erect to decumbent; intnd usu solid. Lig scarious, smt ciliolate; psdpet present, twisted, placing the abx surface of the bld upmost; bld linear to oblong, not folding or drooping at night, lat veins diverging obliquely from the midveins, cross venation evident. Infl pan, usu espatheate; ult br with 1–2 pist spklt and 1 tml, stmt spklt; dis beneath the pist flt and in the pan br. Spklt unisx, heteromorphic, usu in stmt-pist pairs on brlets, with 1 flt; rchl not prolonged beyond the flt. Stam spklt pedlt, smaller than the pist spklt, lanceolate to ovate, caducous; glm unequal; lo glm absent or much shorter than the up glm; up glm somewhat shorter than the flt; lod minute or absent; anth 6. Pist spklt sessile or shortly pedlt, terete, smt inflated; glm unequal to subequal, shorter than the flt, scarious, entire, smt persistent; lm chartaceous, becoming coriaceous, veins 5 or more, mrg involute or utriculate, partly or wholly covered with uncinate hairs, not terminating in a brchd awn; pal 2-veined; lod absent; stl 1, 3-brchd. Car oblong to linear; hila as long as the car.

 

2. BAMBUSOIDEAE Luerss.

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The Bambusoideae includes two tribes, the woody Bambuseae and the herbaceous Olyreae. Their range includes tropical and temperate regions of Asia, Australia, and the Americas, primarily Central and South America. Three species of Bambuseae are native to the Manual region; there are no native species of Olyreae.

Members of the Bambusoideae grow in temperate and tropical forests, high montane grasslands, along riverbanks, and sometimes in savannahs. They are mainly forest understory or margin plants with a limited ability to reproduce, disperse, or survive outside their forest environment. Many have relatively small geographic ranges, and there is a high degree of endemism. The conservation status of most bamboos is not known; all are intrinsically vulnerable because of their breeding behavior and reliance upon a benign forest habitat. Only the C3 photosynthetic pathway is found in the subfamily.

1. Culms woody, usually taller than 1 m, developing complex vegetative branching from the upper nodes; abaxial ligules present on the foliage leaves, rarely present on the culm leaves . . . . . . . . . . . . . . . . . . . . . . . . . 2. Bambuseae

 

3. EHRHARTOIDEAE Link

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The Ehrhartoideae encompasses three tribes, one of which, the Oryzeae, is native to the Manual region; the Ehrharteae is represented by introduced species. The third tribe, Phyllorachideae C.E. Hubb., is native to Africa and Madagascar. There are approximately 120 species in the Ehrhartoideae. They grow in forests, open hillsides, and aquatic habitats.

Molecular data provide strong support for the close relationship of the Oryzeae and Ehrharteae. Morphologically, they are characterized by spikelets that have a distal unisexual or bisexual floret with up to two proximal sterile florets and, frequently, six stamens in the staminate or bisexual florets.

1. Spikelets with 2 sterile florets below the functional floret, both well-developed, at least the upper sterile floret as long as or longer than the functional floret; glumes from ½ as long as the spikelets to exceeding the florets; culms not aerenchymatous; plants of dry to damp habitats . . . . . . . . . . . . . . . . . . . . . . . 4. Ehrharteae

 

4. POÖIDEAE Benth.

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The subfamily Poöideae includes approximately 3300 species, making it the largest subfamily in the Poaceae. It reaches its greatest diversity in cool temperate and boreal regions, extending across the tropics only in high mountains.

1. Inflorescences 1-sided spikes, the spikelets radial to and partially embedded in the rachises; spikelets with 1 floret each . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. Nardeae

1. Inflorescences panicles, racemes, or 2-sided spikes with spikelets radial or tangential to the rachises, sometimes embedded in the axes, never both radial and embedded; spikelets with 1–30 florets.

2. Cauline leaf sheaths closed for at least ¾ their length; lemmas longer than (4.5)6.5 mm or awned or with prominent, parallel veins.

3. Ovary apices glabrous; styles fused at the base, divergent, naked on the lower portion, plumose distally; lemmas often with a purplish band in the distal ½, usually unawned; distal 1–3 florets often reduced to lemmas, the lower 1–2 lemmas often enclosing the terminal lemmas; lodicules about 0.2–0.5 mm long, truncate, fleshy, without a distal membranous portion . . . . . . . . . . . . . . . . . . . . . 9. Meliceae

 

5. ARUNDINOIDEAE Burmeist.

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The Arundinoideae are interpreted here as including one tribe, the Arundineae.

Pl usu per; csp or not, smt rhz, smt stln. Clm 15–1000 cm, ann, hrb to somewhat wd, intnd usu hollow. Lvs usu mostly cauline, often conspicuously distichous; shth usu open; aur usu absent; abx lig usu absent (of hairs in Hakonechloa); adx lig memb or of hairs, if memb, often ciliate; bld without psdpet, smt deciduous at maturity; mesophyll usu non-radiate (radiate in Arundo); adx palisade layer absent; fusoid cells absent; arm cells usu absent (present in Phragmites); Kranz anatomy absent; midribs simple; adx bulliform cells present; stomatal subsidiary cells low dome-shaped or triangular; bicellular microhairs usu present, usu with long, narrow tml cells; papillae usu absent. Infl usu tml, ebracteate, usu pan, occ spicate or rcm. Spklt lat compressed, with 1–several bisx flt or all flt unisx and the species dioecious; flt 1–several, terete or lat compressed, distal flt often rdcd; dis above the glm. Glm 2, from shorter than the adjacent lm to exceeding the distal flt; lm (3)5–7-veined, lanceolate to elliptic, acute to acuminate, smt awned; awns 1 or 3, if 3 not fused into a single bas column; pal subequal to the lm; lod 2, usu free, occ joined at the base, fleshy, usu glab, not, scarcely, or heavily vascularized; anth (1)2–3; ov glab; sty 2, usu free, bases close together. Car usu punctate (long-linear in Molinia); endosperm hard, without lipid; starch grains compound; haustorial synergids absent; emb usu large compared to the car, waisted or not; epiblasts absent; scutellar cleft present; mesocotyl intnd elongate; emb lf mrg usu meeting (overlapping in Hakonechloa).

 

6. CHLORIDOIDEAE Kunth ex Beilschm.

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The subfamily Chloridoideae is most abundant in dry, tropical and subtropical regions. In the Manual region, it reaches its greatest diversity in the southwestern United States. Almost all its members, and all those in the Manual region, have C4 photosynthesis.

There is considerable disagreement concerning the tribal treatment within the Chloridoideae, the number of tribes recognized varying from two to eight. The treatment presented here is conservative in recognizing the Orcuttieae and Pappophoreae as distinct tribes. It departs from most other treatments in merging all other North American taxa into a single tribe, the Cynodonteae.

1. Leaves with little or no distinction between the sheath and blade; ligules not present; plants annual, viscid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18. Orcuttieae

1. Leaves clearly differentiated into sheath and blade; ligules present; plants annual or perennial, not viscid.

 

7. DANTHONIOIDEAE N.P. Barker & H.P. Linder

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8. ARISTIDOIDEAE Caro

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The subfamily Aristidoideae includes one tribe, the Aristideae.

Pl ann or per; usu csp. Clm ann, erect, solid or hollow, usu unbrchd. Lvs distichous; shth usu open; aur absent; abx lig absent or of hairs; adx lig memb and ciliate or of hairs; bld without psdpet; mesophyll cells radiate or non-radiate; adx palisade layer absent; fusoid cells absent; arm cells absent; Kranz anatomy absent or present, when present, with 1 or 2 parenchyma shth; midribs simple; adx bulliform cells present; stomatal subsidiary cells dome-shaped or triangular; bicellular microhairs present, with long, slender, thin-walled tml cells. Infl tml, not lfy, usu pan, smt spikes or rcm. Spklt bisx, with 1 flt; rchl extension absent; dis above the glm. Glm 2, usu longer than the flt, usu acute or acuminate; flt terete or lat compressed, with well-developed cal; lm 1- or 3-veined, more or less coriaceous, with a germination flap, lm mrg overlapping at maturity and concealing the pal, apc evidently 3-awned; awn bases often forming a column, lat awns occ rdcd or absent; pal less than ½ as long as the lm; lod usu present, 2, free, memb, glab, heavily vascularized; anth 1–3; ov glab; haustorial synergids absent; sty 2, free to the base but close. Car usu fusiform, falling with the lm and pal attached; hila short or long, linear; endosperm hard, without lipid; stch g compound; emb small or large relative to the car; epiblasts absent; scutellar cleft present or absent; mesocotyl intnd elongated; embryonic lf mrg meeting.

 

9. CENTOTHECOIDEAE Soderstr.

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The subfamily Centothecoideae is one of the subfamilies that cannot be characterized by a suite of morphological characteristics, but anatomical, micromorphological, and nucleic acid data all support its recognition. It is most abundant in warm-temperate woodlands and tropical forests.

1. Spikelets 4–50 mm long, with 1–15 florets, the lowest florets sometimes sterile, the upper florets bisexual; disarticulation at the base of the florets or the base of the spikelets; leaves not pseudopetiolate; culms 35–150 cm tall; plants not reedlike . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21. Centotheceae

1. Spikelets 1.2–1.8 mm long, with 2(3–4) florets, the lower florets sterile, the upper florets bisexual; disarticulation at the pedicel bases, subsequently below the spikelets; leaves pseudopetiolate; culms 150–400 cm tall; plants reedlike . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22. Thysanolaeneae

 

10. PANICOIDEAE Link

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