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grass family

Habit Plants annual or perennial; usually terrestrial, sometimes aquatic; tufted, mat-forming, cespitose, pluricespitose, or with solitary culms (flowering stems), rhizomes and stolons often well developed. Plants annual or perennial; sometimes cespitose, sometimes rhizomatous.
Culms

annual or perennial, herbaceous or woody, usually erect or ascending, sometimes prostrate or decumbent for much of their length, occasionally climbing, rarely floating;

nodes prominent, sometimes concealed by the leaf sheaths;

internodes hollow or solid, bases meristematic; branching from the basal nodes only or from the basal, middle, and upper nodes;

basal branching extravaginal or intravaginal; branching from the upper nodes intravaginal, extravaginal, or infravaginal.

annual, not woody, usually erect, not branching above the base;

internodes hollow or solid.

Sheaths

usually open, those of the basal leaves sometimes closed;

collars without tufts of hair on the sides;

auricles usually present;

ligules membranous or scarious, sometimes ciliolate, those of the upper and lower cauline leaves usually similar;

pseudopetioles absent;

blades linear to narrowly lanceolate, venation parallel, cross venation not evident, without arm or fusoid cells, surfaces without microhairs, not papillate, cross sections non-Kranz.

Leaves

alternate, 2-ranked, each composed of a sheath and blade encircling the culm or branch;

sheaths usually open, sometimes closed, the margins fused for all or part of their length;

auricles (lobes of tissue extending beyond the margins of the sheaths on either side) sometimes present;

ligules usually present at the sheath-blade junction, particularly on the adaxial surface, abaxial ligules common in the Bambusoideae, membranous, sometimes ciliate, adaxial ligules usually present, of membranous to hyaline tissue, a line of hairs, or a ciliate membrane;

blades usually linear to lanceolate, occasionally ovate to triangular, bases sometimes pseudopetiolate (having a petiole-like constriction), venation usually parallel, sometimes with evident cross veins, occasionally divergent.

Inflorescences

(synflorescences) usually compound, composed of simple or complex aggregations of primary inflorescences, aggregations paniculate, spicate, or racemose or of spikelike branches, often with an evident rachis (central axis), primary inflorescences spikelets, pseudospikelets, or spikelet equivalents;

inflorescence branches usually without obvious bracts.

usually spikes or spikelike racemes, with 1-5 sessile or subsessile spikelets per node, occasionally panicles, sometimes with morphologically distinct sterile and bisexual spikelets within an inflorescence;

pedicels absent or to 4 mm;

disarticulation usually above the glumes and beneath the florets, sometimes in the rachises, sometimes at the inflorescence bases.

Spikelets

with (0-1)2(3-6) glumes (empty bracts) subtending 1-60 florets, glumes and florets distichously attached to a rachilla (central axis);

pseudospikelets with bud-subtending bracts below the glumes.

usually laterally compressed, sometimes terete, with 1-16 bisexual florets, the distal (or only) floret sometimes sterile;

rachillas sometimes prolonged beyond the base of the distal floret.

Glumes

usually with an odd number of veins, sometimes awned.

unequal to equal, shorter than to longer than the adjacent florets, subulate, lanceolate, rectangular, ovate, or obovate, 1-5-veined, absent or vestigial in some species;

florets laterally compressed to terete;

calluses glabrous or hairy;

lemmas lanceolate to rectangular, stiffly membranous to coriaceous, sometimes keeled, 5(7)-veined, veins not converging distally, inconspicuous, apices entire, lobed, or toothed, unawned or awned, awns terminal, unbranched, lemma-awn junction not evident;

paleas usually subequal to the lemmas, sometimes considerably shorter or slightly longer than the lemmas;

lodicules 2, without venation, usually ciliate;

anthers 3;

ovaries with hairy apices;

styles 2, bases free.

Caryopses

ovoid to fusiform, longitudinally grooved, not beaked, pericarp thin;

hila linear;

embryos about 1/3 as long as the caryopses.

Florets

bisexual, staminate, or pistillate, usually composed of a lemma (lower bract) and palea (upper bract), lodicules, and reproductive organs, often laterally or dorsally compressed, sometimes round in cross section;

lemmas usually with an odd number of veins, often awned, bases frequently thick and hard, forming a callus, backs rounded or keeled over the midvein, awns usually 1(-3), arising basally to terminally;

paleas usually with 2 major veins, with 0 to many additional veins between the major veins, sometimes also in the margins, often keeled over the major veins;

lodicules (0)2-3, inconspicuous, usually without veins, bases swelling at anthesis;

stamens usually 3, sometimes 1(2) or 6+, filaments capillary, anthers versatile, usually all alike within a floret, sometimes 1 or 2 evidently longer than the others;

ovaries 1-loculed, with (1)2-3(4) styles or style branches, stigmatic region usually plumose.

Fruits

caryopses, pericarp usually dry and adhering to the seed, sometimes fleshy or dry and separating from the seed at maturity or when moistened;

embryos ⅕ as long as to almost equaling the caryopses, highly differentiated with a scutellum (absorptive organ), a shoot with leaf primordium covered by the coleoptile (shoot sheath), and a root covered by the coleorhiza (root sheath);

hila punctate to linear.

x

= 5,6, 7, 9, 10, 11, 12.

= 7.

Poaceae

Poaceae tribe Triticeae

Distribution
[BONAP county map]
Discussion

The Poaceae or grass family includes approximately 700 genera and 11,000 species (Chen et al. 2006). The two grass volumes in this series treat 10 subfamilies, 25 tribes, 236 genera, and 1373 species. Of these, all the subfamilies, 22 tribes, 136 genera, and 892 species are native to the Flora region; 2 tribes, 78 genera, and 290 species have become established in the region. The remaining taxa include ornamental species; species grown for research purposes; species that, if introduced to the region, would pose a threat to important agricultural species; and a few waifs, i.e., species that have been found in the region but have not become established. Most of the waifs are species that were found on ballast dumps near ports around the turn of the last century.

Grasses constitute the fourth largest plant family in terms of number of species. Nevertheless, the family is clearly more significant than any other plant family in terms of geographic, ecological, and economic importance. Grasses grow in almost all terrestrial environments, including dense forests, open deserts, and freshwater streams and lakes. There are no truly marine grasses, but some species grow within reach of the highest tides.

In addition to being widely distributed, grasses are often dominant or co-dominant over large areas. The importance of such areas to humans is reflected in the many words that exist for grasslands, words such as meadow, palouse, pampas, prairie, savanna(h), steppe, and veldt. Not surprisingly, given their abundance and prevalence, grasses are of great ecological importance as soil stabilizers and as providers of shelter and food for many different animals.

The economic importance of grasses to humans is almost impossible to overestimate. The wealth of individuals and countries is dependent on the availability of such sources of grain as Triticum (wheat), Oryza (rice), Zea (corn or maize), Hordeum (barley), Avena (oats), Secale (rye), Eragrostis (tef), and Zizania (wild rice). Most countries invest heavily in research programs designed to develop better strains of these grasses and the many other grasses that are used for livestock, soil stabilization, and revegetation. Developing improved grasses for recreation areas, such as playing fields, golf courses, and parks, is also a major industry in many parts of the world; increasing recognition of the aesthetic value af grasses is reflected in their prominence in horticultural catalogs.

There are, of course, grasses that are considered undesirable, at least in some parts of the world, but even the most obnoxious grasses may be well-regarded over a portion of their range. For instance, Bromus tectorum (cheatgrass) is a noxious, fire-prone invader of western North American ecosystems; it is also welcomed as a source of early spring feed in some parts of the Flora region. Cynodon dactylon (bermudagrass) is listed as a noxious weed in some jurisdictions; in others it is valued as a lawn grass.

Although grasses are widespread and often dominant in open areas, all evidence points to an origin of the family in forests, most likely in the Southern Hemisphere, at least 55-70 mya (Grass Phylogeny Working Group 2001). Recent evidence from phytoliths (isolated silica bodies commonly produced inside the epidermal cells of grasses and some other plants) embedded in fossil coprolites strongly suggests that grasses evolved earlier in the Cretaceous than previously thought (Prasad et al. 2005). Living representatives of the three earliest lineages of the grass family, together comprising about 30 species, are perennial, broad-leaved plants of relatively small stature, native to tropical or subtropical forests in South America, Africa, southeast Asia, some Pacific Islands, and northern Australia. The major diversification of the family probably occurred in the mid-Cenozoic, and was associated with climatic changes that produced more open habitats. All major lineages of the grass family were present by the middle of the Miocene (Jacobs et al. 1999), and C4 photosynthesis in grasses had evolved by then, as well.

Molecular and morphological data unequivocally support a single origin for the Poaceae (Grass Phylogeny Working Group 2001). The caryopsis, a single-seeded, usually dry and indehiscent fruit with the pericarp usually strongly adherent to the seed, and the laterally positioned, highly differentiated embryo are unique to grasses. Beyond the three early-diverging lineages (Anomochlooideae Potztal, Pharoideae, and Puelioideae L.G. Clark et al.), the great diversity of grasses can be divided into two major lineages: the BEP clade (Bambusoideae, Ehrhartoideae, and Pooideae); and the PACMCAD clade (Panicoideae, Arundinoideae, Chloridoideae, Micrairoideae Pilger, Centothecoideae, Aristidoideae, and Danthonioideae, i.e., the PACCAD clade of volume 25 plus the Micrairoideae, support for recognition of which was obtained after publication of that volume). Relationships among the BEP grass lineages remain uncertain, and some evidence points to the Pooideae as being more closely related to the PACMCAD clade than to the Bambusoideae or Ehrhartoideae. The PACMCAD clade includes all known C4 or warm-season grasses.

The closest relatives of the Poaceae lie within a group of six families, all native primarily to the Southern Hemisphere: Joinvilleaceae Toml. & A.C. Sm., Ecdeiocoleaceae D.F. Cutler &c& Airy Shaw, Restionaceae R. Br., Centrolepidaceae Endl., Anarthriaceae D.E Cutler &c& Airy Shaw, and Flagellariaceae Dumort. (Poales Small, sensu stricto). Joinvilleaceae, Ecdeiocoleaceae, and Poaceae constitute a three family clade, with Ecdeiocoleaceae probably being closer than Joinvilleaceae to the Poaceae (Bremer 2000; Bremer 2002; Michelangeli et al. 2003). Rudall et al. (2005), based on a study of reproductive structures in the Ecdeiocoleaceae, suggest that the grass caryopsis may represent "one end of a transformation series embodied by the reduced gynoecial structure and indehiscent fruit of other Poales such as Flagellaria and Ecdeiocolea" (p. 1441).

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

The Triticeae are primarily north-temperate in distribution. The tribe includes 400-500 species, among which are several important cereal, forage, and range species. Its generic treatment is contentious. Linnaeus (1753) recognized five genera among the species now included in the tribe; Hordeum and Secale are the only two that still have his circumscription. Hordeum is also the only genus to include both annual and perennial species. The lack of agreement concerning the generic treatment of the tribe reflects the prevalence of natural hybridization, introgression, polyploidy, and reticulate relationships among its species. These factors preclude the circumscription of monophyletic groups, and mitigate against the delineation of morphologically coherent groups. Tzvelev (1975) argued that these same factors contribute to the tribe's success by maintaining a "generalist" genome.

The major disagreement in the treatment of the annual genera concerns Triticum and Aegilops. Some (e.g., Kimber and Feldman 1987) advocate treating them as a single genus in recognition of their close genetic similarity; others argue for maintaining them as separate genera (e.g., van Slageren 1994). Love (1984) divided them among 14 genera. They are accepted here in their traditional senses, despite the strong argument for their combination, largely in deference to the wealth of literature, reports, and genetic resources that have been accumulated under these two names. Spontaneous hybridization and introgression between the two are common, and most species of Triticum are derived from hybrids between the two genera. Nevertheless, they differ in their ecology and, to some extent, in their morphology.

Treatment of the perennial species is more contentious. Restriction of Agropyron to what are known in English as the crested wheatgrasses is universally accepted; most taxonomists also accept the placement of alkaline-tolerant species that are strongly rhizomatous or have short, subulate glumes in Leymus. Pseudoroegneria, Pascopyrum, and Thinopyrum are less accepted. They are widely accepted by those working in genetic resources, but less so by those involved in floristics who prefer to include them in Elymus; all were traditionally included in Agropyron. Another area of disagreement is the treatment of Elytrigia Desv., Roegneria K. Koch, and Hystrix Moench. Species sometimes placed in Elytrigia are here included in Elymus, Thinopyrum, or Pseudoroegneria; species of Roegneria in Elymus; and species of Hystrix in Elymus or Leymus. Wide acceptance of a single treatment is hampered by the existence of differing taxonomic traditions, and by the lack of a coordinated international examination of morphological variation among the tribe's species.

The treatment followed here is strongly influenced by the treatments of Love (1984) and Dewey (1984), particularly with respect to the perennial genera. Both advocated using genomic constitution as the basis for generic delimitation in the tribe. The genomic constitution of individual species is determined by observing meiotic chromosome pairing in hybrids. The base chromosome number in the tribe is seven. If a hybrid between two tetraploids forms 7 quadrivalents and 14 bivalents at meiosis, its parents are considered to have one similar set of chromosomes or haplome, and one dissimilar haplome. The three haplomes can then be assigned codes. For example, one parent might be said to have the E and F haplomes, or an EF genomic constitution, and the other the E and L haplomes, or an EL genomic constitution. The prevalence of polyploids and the ease of forming hybrids in the Triticeae has enabled cytogeneticists to build up a rather complete picture of the genomic constitution of its members. This led to the discovery that there is a strong, but not perfect, correlation between morphology and genomic constitution. The haplome codes used in this volume are those endorsed by the International Triticeae Consortium (http://herbarium.usu.edu/Triticeae/genmsymb.htm). Molecular tools reveal a pattern that is, in general, consistent with the cytogenetic data, but they often reveal an underlying complexity that cannot be discerned using only classical cytogenetic techniques.

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

Key

Key to Tribes

1. Leaf blades with divergent veins; spikelets unisexual and dimorphic, the pistillate lemmas with uncinate hairs (Pharoideae; FNA 24:11)
Phareae
1. Leaf blades with parallel veins; spikelets bisexual, unisexual, or modified into plantlets, the pistillate lemmas never with uncinate hairs.
→ 2
2. Culms perennial, woody or herbaceous, often developing complex branching systems from the upper nodes; leaves on the upper portion of the culms, or distal on the branches, usually pseudopetiolate (Bambusoideae).
→ 3
3. Culms woody, to 30 m tall; leaves strongly dimorphic, those of the main culms (culm leaves) with expanded sheaths and often with reduced, non-photosynthetic blades, those of the branches (foliage leaves) with abaxial ligules; blades of the distal leaves not folding at night or under stress; florets bisexual; plants native or introduced, often cultivated (FNA 24:15)
Bambuseae
3. Culms herbaceous, to 3.5 m tall or climbing; leaves not strongly dimorphic; blades of the distal leaves often folding at night or under stress; florets unisexual; plants known only in cultivation in the Flora region (FNA 24:29)
Olyreae
2. Culms usually annual, sometimes facultatively perennial, rarely woody, sometimes branching from the upper nodes but the branching system not complex; leaves usually not pseudopetiolate.
→ 4
4. Spikelets almost always with 2 florets, the lower florets in the spikelets always sterile or staminate, frequently reduced to lemmas, occasionally missing, the upper florets bisexual, staminate, or sterile, unawned or awned from the lemma apices or, if the lemmas bilobed, from the sinuses; glumes membranous and the upper lemma stiffer than the lower lemma, or both florets reduced and concealed by the stiff to coriaceous glumes; rachilla not prolonged beyond the second floret (Panicoideae, in part).
→ 5
5. Glumes flexible, membranous, the lower glumes usually shorter than the upper glumes, sometimes missing, the upper glumes usually subequal to or exceeded by the upper floret; lower lemmas membranous; upper lemmas usually coriaceous to indurate, sometimes membranous; upper paleas similar in texture; spikelets usually single or in pairs, occasionally in triplets and all pedicellate, often shortly so) (FNA 25:353)
Paniceae
5. Glumes stiff, coriaceous to indurate, often subequal, at least 1 and usually both exceeding the upper floret (excluding the awn); both lemmas hyaline; paleas hyaline or absent; most spikelets in pairs or triplets, at least 1 spikelet in each group usually sessile; pedicels shorter or only a little longer than the sessile spikelets (FNA 25:602)
Andropogoneae
4. Spikelets either with other than 2 florets or, if with 2, the lower floret bisexual or the upper floret awned from the back or base of the lemma, or the spikelets bulbiferous; glumes usually membranous; lemmas scarious to indurate; rachilla sometimes prolonged beyond the distal floret.
→ 6
6. Spikelets with 1 floret; lemmas terminating in a 3-branched awn (the lateral branches sometimes greatly reduced); callus well developed; ligules usually of hairs, sometimes ciliate membranes, the cilia longer than the membranous base (Aristidoideae; FNA 25:314)
Aristideae
6. Spikelets with more than 1 floret or, if only 1, the lemma not terminating in a 3-branched awn; callus development various; ligules various.
→ 7
7. Spikelets with 1 sexual floret or the spikelets bulbiferous; glumes absent or less than ¼ as long as the adjacent floret; lower glumes, if present, without veins, upper glumes, if present, veinless or 1-veined.
→ 8
8. Upper glumes present, 1-veined; lower glumes absent or much shorter than the upper glumes and lacking veins (Pooideae, in part; FNA 24:57)
Brachyelytreae
8. Both glumes absent or lacking veins.
→ 9
9. Inflorescences 1-sided spikes; triangular in cross section, [Pooideae, in part; FNA 24:62)
Nardeae
9. Inflorescences panicles; spikelets laterally compressed or terete.
→ 10
10. Culms aerenchymatous, 20-500 cm long; plants of wet places, often emergent, sometimes floating; lemmas of the bisexual or pistillate florets 3-14-veined; paleas 3-10-veined (Ehrhartoideae, in part; FNA 24:36)
Oryzeae
10. Culms not aerenchymatous, 2-300 cm tall; plants of wet or dry habitats but not emergent or floating; lemmas of the bisexual or pistillate florets 1-3-veined; paleas 2-veined.
→ 11
11. Culms 2-19 cm tall; plants of cold or damp habitats, not rhizomatous; sheaths of the flag leaves closed for at least 1/2 their length; caryopses exposed at maturity (Pooideae, in part; FNA 24:378)
Poeae
11. Culms 5-300 cm tall; plants usually of warm or dry habitats, often rhizomatous; sheaths of the flag leaves open to the base; caryopses not exposed at maturity (Chloridoideae, in part; FNA 25:14)
Cynodonteae
7. Spikelets usually with more than 1 sexual floret; usually with 2 glumes, 1 or both glumes often longer than ¼ the length of the adjacent floret and/or with more than 1 vein, always longer in taxa with 1 sexual floret.
→ 12
12. Lemmas unawned, flabellate or with (5)7-15 awnlike teeth (Chloridoideae, in part).
→ 13
13. Plants not viscid, usually perennial; ligules present, composed of hairs (FNA 25:285)
Pappophoreae
13. Plants viscid annuals; ligules absent (FNA 25:290)
Orcuttieae
12. Lemmas awned or unawned, lanceolate, rectangular, or ovate, apices entire, mucronate, bilobed, or bifid, occasionally 4-lobed or 4-5-toothed, sometimes erose.
→ 14
14. Cauline leaf sheaths closed for 1/2 their length or more; glumes usually exceeded by the distal florets, sometimes greatly so (Pooideae, in part).
→ 15
15. Spikelets 5-80 mm long, not bulbiferous; lemmas usually awned, often bilobed or bifid, veins convergent distally; ovary apices hairy (FNA 24:193)
Bromeae
15. Spikelets 0.7-60 mm long, sometimes bulbiferous; lemmas often unawned, not both bilobed/bifid and with convergent veins; ovary apices usually glabrous.
→ 16
16. Lemma veins (4)5-15, usually prominent, parallel distally; spikelets 2.5-60 mm long, not bulbiferous (FNA 24:67)
Meliceae
16. Lemmas veins 1-9, often inconspicuous, usually convergent distally; spikelets 0.7-18(20) mm long, sometimes bulbiferous (FNA 24:378)
Poeae
14. Cauline leaf sheaths open for at least 1/2 their length; glumes exceeding or exceeded by the distal florets.
→ 17
17. Spikelets with 1 floret; lemmas terminally or subterminally awned, the junction of the awn and lemma conspicuous; rachillas not prolonged beyond the base of the floret (Pooideae, in part; FNA 24:109)
Stipeae
17. Spikelets with 1-60 florets; lemmas unawned or awned, awns basal to terminal, if terminal or subterminal, the lemma-awn junction not conspicuous; rachillas often prolonged beyond the base of the distal floret.
→ 18
18. Ligules, at least of the flag leaves, of hairs, a ciliate ridge or membrane bearing cilia longer than the basal ridge or membrane; leaves usually hairy on either side of the ligule; auricles absent.
→ 19
19. Lemmas of the fertile florets with 3-11 inconspicuous veins, never glabrous, if with 3 veins, pilose throughout or with transverse rows of tufts of hair, if with 5-11 veins, the margins pilose proximally, the hairs not papillose-based; lemma apices usually bilobed or bifid and awned or mucronate from the sinus, if acute to acuminate, the lemmas pilose; awns twisted proximally (Danthonioideae; FNA 25:298)
Danthonieae
19. Lemmas of the fertile florets usually with 1-3 conspicuous veins, sometimes with 3 inconspicuous veins or 5-11 veins, often glabrous, if with 3 veins, usually glabrous throughout or hairy over the veins, sometimes the margins with papillose-based hairs; lemma apices acute to obtuse, bilobed, or 4-lobed, often mucronate or awned from the sinuses; awns usually not twisted.
→ 20
20. Lemmas 1-11-veined, veins glabrous or hairy, margins without papillose-based hairs; rachillas and calluses not pilose, sometimes strigose or strigulose; basal internodes of the culms not persistent, not swollen and clavate (Chloridoideae, in part; FNA 25:14)
Cynodonteae
20. Lemmas 3(5)-veined, veins glabrous, margins sometimes with papillose-based hairs; rachillas or calluses pilose or the basal internodes of the culms persistent, often swollen and clavate (Arundinoideae, in part; FNA 25:7)
Arundineae
18. Ligules membranous, if ciliate, the cilia shorter than the membranous base; leaves usually glabrous on either side of the ligule; auricles present or absent.
→ 21
21. Inflorescences panicles or unilateral racemes, not spikelike, without spike-like branches; spikelets solitary, the lowest 0-4 florets in a spikelet sterile or staminate, the distal florets sexual.
→ 22
22. Spikelets with (1)2-25 bisexual florets; all lemmas similar in size and shape; glumes and lemmas membranous (Centothecoideae).
→ 23
23. Culms 35-150 cm tall; spikelets with (2)3-26 florets, including the lowest (0)1-4 sterile or staminate florets; lower glumes (1)2-9-veined (FNA 25:344)
Centotheceae
23. Culms 150-400 cm tall; spikelets with 2-4 florets, including the lowest sterile floret; glumes 0-1-veined (FNA 25:349)
Thysanolaeneae
22. Spikelets with 1 bisexual or unisexual floret; lemmas of the sterile florets usually differing in size and shape from those of the sexual floret; glumes membranous, lemmas of the sexual florets firmer.
→ 24
24. Lemmas of the lower florets coriaceous, at least the upper exceeding the sexual floret (Ehrhartoideae, in part; FNA 24:33)
Ehrharteae
24. Lemmas of the lower florets membranous, often both much shorter than the sexual floret, sometimes subequal to it, sometimes only 1 sterile floret present (Pooideae, in part; FNA 24:378)
Poeae
21. Inflorescences panicles, racemes, or spikes; spikelets sometimes in pairs or triplets, sterile florets, if any, distal to the bisexual or pistillate florets.
→ 25
25. Lemmas with 1-3 or 9-11 conspicuous veins; sheaths open; blade cross sections with Kranz leaf anatomy (Chloridoideae, in part; FNA 25:14)
Cynodonteae
25. Lemmas with (1)3-15 often inconspicuous veins, if with 3 conspicuous veins, the sheaths closed; sheaths open or closed; blade cross sections without Kranz leaf anatomy.
→ 26
26. Inflorescences spikes or spikelike; spikelets 1-5+ per node, at least 1 spikelet sessile or subsessile (Pooideae, in part).
→ 27
27. Upper glumes 5-9-veined; spikelets subsessile and solitary at the nodes; auricles absent (FNA 24:187)
Brachypodieae
27. Upper glumes 1-5-veined; spikelets 1-5+ per node, usually at least 1 sessile at each node, sometimes highly reduced branches present; auricles present or absent.
→ 28
28. Inflorescences with 1-5 spikelets at a node, if 3, usually with 1 sessile and 2 pedicellate spikelets, if 1, the spikelet tangential to or embedded in the rachis, with 2 glumes, the glumes facing each other; ovaries with hairy apices; auricles often present (FNA 24:238)
Triticeae
28. Inflorescences spikelike panicles with highly reduced branches, or spikes with spikelets radial to the rachises and all but the terminal spikelet with only 1 glume, or spikes with spikelets tangential to the rachises and having 2 glumes adjacent to each other; ovaries with glabrous apices; auricles usually absent (FNA 24:378)
Poeae
26. Inflorescences panicles, with no sessile spikelets.
→ 29
29. Caryopses with a thick pericarp forming a distinct apical knob or beak at maturity; lemmas 3(5)-veined (Pooideae, in part; FNA 24:64)
Diarrheneae
29. Caryopses usually with a thin pericarp, never with a distinct apical beak or knob; lemmas 3-9-veined.
→ 30
30. Glumes subulate, stiff; lemmas unawned or with awns to 4 mm long (Pooideae, in part; FNA 24:238)
Triticeae
30. Glumes lanceolate, membranous; lemmas awned or unawned, awn length varied.
→ 31
31. Rachillas hairy, hairs 2-3 mm long; lemmas coriaceous; plants established in California, sometimes cultivated as ornamentals (Pooideae, in part; FNA 24:109)
Stipeae
31. Rachillas glabrous to hairy, hairs shorter than 2 mm; lemmas membranous to coriaceous; plants native, established, or cultivated.
→ 32
32. Leaves to 2 cm wide, usually not conspicuously distichous; culms 0.01-2.75 m tall, usually less than 1 cm thick (Pooideae, in part; FNA 24:378)
Poeae
32. Leaves 2-10 cm wide, often conspicuously distichous; culms 2-10(15) m tall, often more than 1 cm thick.
→ 33
33. Lower cauline blades disarticulating, upper cauline blades forming a flat, fan-shaped arrangement (Panicoideae, in part; FNA 25:352)
Gynerieae
33. Lower cauline blades persistent, upper cauline blades not forming a flat, fan-shaped arrangement (Arundinoideae, in part; FNA 25:7)
Arundineae
1. Spikelets 2-7 at all or most nodes.
→ 2
2. Spikelets 3 at each node, the central spikelets sessile, the lateral spikelets usually pedicellate, sometimes all 3 spikelets sessile in cultivated plants; spikelets with 1 floret, usually only the central spikelet with a functional floret, the florets of the lateral spikelets usually sterile and reduced, in cultivated plants all florets functional or those of the lateral spikelets functional and those of the central spikelet reduced
Hordeum
2. Spikelets usually other than 3 at each node, if 3, all three sessile; spikelets with 1-11 florets, if 1 floret, additional reduced or sterile florets present distal to the functional floret in at least 1 spikelet per node.
→ 3
3. Lemmas strongly keeled, keels conspicuously scabrous distally, scabrules 0.5-0.8 mm long; lemma awns straight
Secale
3. Lemmas rounded proximally, sometimes keeled distally, keels not or inconspicuously scabrous distally; lemma awns straight, flexuous, or variously curved.
→ 4
4. Plants annual, weedy; spikelets with only 1 bisexual floret
Taeniatherum
4. Plants perennial, usually not weedy; spikelets usually with more than 1 bisexual floret.
→ 5
5. Lemma awns (0)1-120 mm long; anthers 0.9-6 mm long; blades with well-spaced, unequally prominent veins on the adaxial surfaces
Elymus
5. Lemmas usually unawned or with awns up to 7 mm long, if awns 16-35 mm long, anthers 6-8 mm; blades usually with closely spaced, equally prominent veins on the adaxial surfaces.
→ 6
6. Disarticulation in the spikelets, beneath the florets; plants sometimes cespitose, often rhizomatous
Leymus
6. Disarticulation tardy, in the rachises; plants cespitose, not rhizomatous
Psathyrostachys
1. Spikelets 1 at all or most nodes.
→ 7
7. Spikelets usually more than 3 times the length of the middle rachis internodes, usually divergent, sometimes ascending; rachis internodes 0.2-5.5 mm long.
→ 8
8. Glumes with 2 prominent keels, keels with tufts of hair
Dasypyrum
8. Glumes initially with 1 keel, sometimes 2-keeled at maturity, keels glabrous or hairy, hairs never in tufts.
→ 9
9. Plants annual; anthers 0.4-1.4 mm long; spikes 0.8-4.5 cm long
Eremopyrum
9. Plants perennial; anthers 3-5 mm long; spikes 1.3-15 cm long
Agropyron
7. Spikelets 72-3 times the length of the middle rachis internodes, appressed or ascending; rachis internodes 3-28 mm long.
→ 10
10. Glumes subulate to narrowly lanceolate, tapering from below midlength, 1(3)-veined at midlength. 11. Glumes lanceolate, tapering to acuminate apices from near midlength or below, keels curving to the side distally; plants always rhizomatous
Pascopyrum
11. Glumes subulate to lanceolate, tapering from below midlength, keels straight or almost so; plants often rhizomatous
Leymus
10. Glumes lanceolate, rectangular, ovate, or obovate, narrowing beyond midlength, often the in distal 1/4 (1)3-5(7)-veined at midlength.
→ 11
12. Plants annual; glumes often with lateral teeth or awns, midveins smooth throughout.
→ 13
13. Glumes rounded over the midveins; plants weedy
Aegilops
13. Glumes keeled over the midveins; plants cultivated, sometimes escaping
Triticum
12. Plants perennial; glumes without lateral teeth or awns, midveins sometimes scabrous.
→ 14
14. Glumes stiff, truncate, obtuse, or acute, unawned; glume keels smooth proximally, usually scabrous distally
Thinopyrum
14. Glumes flexible, acute to acuminate, sometimes awn-tipped; glume keels usually uniformly smooth or scabrous their whole length, sometimes smooth proximally and scabrous distally.
→ 15
15. Spikelets distant, not or scarcely reaching the base of the spikelet above on the same side of the rachis; anthers 4-8 mm long
Pseudoroegneria
15. Spikelets usually more closely spaced, reaching midlength of the spikelet above on the same side of the rachis; anthers 0.7-7 mm long
Elymus
Source FNA vol. 24, p. 3. Author: Lynn G. Clark; Elizabeth A. Kellogg;. FNA vol. 24, p. 238. Author: Mary E. Barkworth;.
Parent taxa Poaceae > subfam. Pooideae
Subordinate taxa
Aegilops, Agropyron, Dasypyrum, Elymus, Eremopyrum, Hordeum, Leymus, Pascopyrum, Psathyrostachys, Pseudoroegneria, Secale, Taeniatherum, Thinopyrum, Triticum
Synonyms family gramineae
Name authority Barnhart Dumort.
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