Poaceae tribe Stipeae
Piptochaetium
speargrass
annual or perennial, not woody, branches 1 to many at the upper nodes.
4-150 cm, usually erect, sometimes decumbent, glabrous, not branched above the base;
basal branching intravaginal;
prophylls shorter than the sheaths, mostly glabrous, keels usually with hairs, apices bifid, teeth 1-3 mm;
cleistogenes not developed.
open to the base, margins glabrous;
ligules membranous, decurrent, truncate to acute, sometimes highest at the sides, sometimes ciliate;
blades convolute to flat, translucent between the veins, often sinuous distally.
basally concentrated to evenly distributed;
sheaths open, margins not fused, sometimes ciliate distally, basal sheaths sometimes concealing axillary panicles (cleistogenes), sometimes wider than the blade;
collars sometimes with tufts of hair at the sides extending to the top of the sheaths;
auricles absent;
ligules scarious, often ciliate, cilia usually shorter than the base, ligules of the lower and upper cauline leaves sometimes differing in size and vestiture;
pseudopetioles absent;
blades linear to narrowly lanceolate, venation parallel, cross venation not evident, cross sections non-Kranz, without arm or fusoid cells;
epidermes of adaxial surfaces sometimes with unicellular microhairs, cells not papillate.
usually terminal panicles, occasionally reduced to racemes in depauperate plants, sometimes 2-3 panicles developing from the highest cauline node.
terminal panicles, open or contracted, spikelets usually confined to the distal 1/2 of each branch.
usually with 1 floret, sometimes with 2-6 florets, laterally compressed to terete;
rachillas not prolonged beyond the base of the floret in spikelets with 1 floret, prolonged beyond the base of the distal floret in spikelets with 2-6 florets, prolongation hairy, hairs 2-3 mm;
disarticulation above the glumes and beneath the florets.
4-22 mm, with 1 floret;
rachillas not prolonged beyond the base of the floret;
disarticulation above the glumes, beneath the floret.
usually exceeding the floret(s), always longer than 1/4 the length of the adjacent floret, 1-10-veined, narrowly lanceolate to ovate, hyaline or membranous, flexible;
florets usually terete, sometimes laterally or dorsally compressed;
calluses usually well-developed, rounded or blunt to sharply pointed, often antrorsely strigose;
lemmas lanceolate, rectangular, or ovate, membranous to coriaceous or indurate, 3-5-veined, veins inconspicuous, apices entire, bilobed, or bifid, awned, lemma-awn junction usually conspicuous, awns 0.3-30 cm, not branched, usually terminal and centric or eccentric, sometimes subterminal, caducous to persistent, not or once- to twice-geniculate, if geniculate, proximal segment(s) twisted, distal segment straight, flexuous, or curled, not or scarcely twisted;
lodicules 2 or 3;
anthers 1 or 3, sometimes differing in length within a floret;
ovaries glabrous throughout or pubescent distally;
styles 2(3-4)-branched.
subequal, longer than the floret, lanceolate, 3-7(8)-veined;
florets globose to fusiform, terete to laterally compressed;
calluses well developed, sharp or blunt, glabrous or antrorsely strigose, hairs yellow to golden brown;
lemmas coriaceous to indurate, glabrous or pubescent, striate, particularly near the base, smooth, papillose, or tuberculate, often smooth on the lower portion and papillate to tuberculate distally, margins involute, fitting into the grooved palea, apices fused into a crown, awned, lemmas often narrowed below the crown, crowns usually ciliate;
awns caducous to persistent, usually twice-geniculate, first 2 segments usually twisted and hispid, terminal segment straight and scabridulous;
paleas longer than the lemmas, similar in texture, glabrous, sulcate between the veins, apices prow-tipped;
lodicules 2 or 3, membranous, glabrous, blunt or acute;
anthers 3;
ovaries glabrous;
styles 2.
ovoid to fusiform, not beaked, pericarp thin;
hila linear;
embryos less than 1/3 the length of the caryopses.
terete to globose or lens-shaped, x = 11.
= 7, 8, 10, 11, 12.
Poaceae tribe Stipeae
Piptochaetium
The tribe Stipeae includes about 15 genera and approximately 500 species. It grows in Africa, Australia, South and North America, and Eurasia. In Australia, South America, and Asia, it is often the dominant grass tribe over substantial areas. It is not present in southern India, and is represented by only one native species in southern Africa. Most species grow in arid or seasonally arid, temperate regions.
Morphological considerations have led to the Stipeae being placed in three different subfamilies (Pooideae, Bambusoideae, and Arundinoideae) in the past, and even to recognition as a subfamily. Molecular data support its treatment as an early diverging lineage within the Pooideae (Soreng and Davis 1998; Grass Phylogeny Working Group 2001) that is more closely related to the Meliceae than the core pooid tribes.
Decker (1964) suggested including Ampelodesmos in the Stipeae on the basis of the cross sectional anatomy of its leaf blades. His suggestion is supported, not always strongly, by molecular studies (Soreng and Davis 1998; Grass Phylogeny Working Group 2001; Jacobs et al. 2006). The usual alternative is to treat Ampelodesmos as the only genus of a closely related, monospecific tribe, the Ampelodesmeae (Conert) Tutin, because it is so distinct from other members of the Stipeae, being, for example, the only member of the tribe with more than 1 floret in its spikelets and rachillas that are prolonged beyond the base of the terminal floret in a spikelet.
The lowest chromosome number known in the Stipeae is 2n =18 (Prokudin et al. 1977), suggesting that all members of the tribe are ancient polyploids. The wide range of base numbers listed is based on numbers for the various genera. The primary basic chromosome number for the tribe is probably 5 or 6, with higher numbers reflecting ancient euploidy.
(Discussion copyrighted by Flora of North America; reprinted with permission.)
Piptochaetium is primarily South American, being particularly abundant in Argentina. It has 27 species. Four species are native in the Flora region; two South American species are established at a single location in Marin County, California.
All stipoid species with elongate florets were included in Stipa by Hitchcock (1935, 1951) and other North American taxonomists. Parodi (1944) argued that Piptochaetium should be expanded to include those species of Stipa with elongated florets that shared with Piptochaetium its distinctive lemma and palea morphology, hair color, and leaf anatomy. His interpretation is now universally accepted.
The basal chromosome number of Piptochaetium is probably 11. This interpretation implies that counts of In = 42 (Gould 1965 and Reeder 1968 for P. fimbriatum; Reeder 1977 for P. pringlei) represent an aneuploid reduction from 2n = 44. It is also possible that the base number is 7, as is common in the Pooideae.
(Discussion copyrighted by Flora of North America; reprinted with permission.)
1. Spikelets with 2-6 florets | Ampelodesmos |
1. Spikelets with 1 floret. | → 2 |
2. Paleas sulcate, longer than the lemmas; lemma margins involute, fitting into the paleal groove; lemma apices not lobed | Piptochaetium |
2. Paleas flat, from shorter than to longer than the lemmas; lemma margins convolute or not overlapping; lemma apices often lobed or bifid. | → 3 |
3. Prophylls exceeding the leaf sheaths; plants cultivated as ornamentals. | → 4 |
4. Panicles contracted; lemma awns once-geniculate | Macrochloa |
4. Panicles open; lemma awns twice-geniculate | Celtica |
3. Prophylls concealed by the leaf sheaths; plants native, introduced, sometimes cultivated as ornamentals. | → 5 |
5. Flag leaf blades up to 12 mm long; basal leaves overwintering | Oryzopsis |
5. Flag leaf blades more than 10 mm long; basal leaves not overwintering. | → 6 |
6. Plants with multiple stiff branches from the upper nodes; pedicels sometimes plumose; species cultivated as ornamentals in the Flora region | Austrostipa |
6. Plants not branching at the upper nodes, or with a few, flexible branches; pedicels never plumose; species native, established introductions, or cultivated as ornamentals. | → 7 |
7. Apices of the leaf blades sharp and stiff; caryopses obovoid, often with 3 smooth ribs at maturity; cleistogenes usually present | Amelichloa |
7. Apices of the leaf blades acute to acuminate, never both sharp and stiff; caryopses fusiform, ovoid or obovoid, without ribs; cleistogenes sometimes present. | → 8 |
8. Lemma margins strongly overlapping their whole length at maturity, lemma bodies usually rough throughout, apices not lobed; paleas 1/4 - 1/2 the length of the lemmas, without veins, glabrous | Nassella |
8. Lemma margins usually not or only slightly overlapping for some or all of their length at maturity, strongly overlapping in some species with smooth lemmas, lemma bodies usually smooth on the lower portion, apices often 1-2-lobed; paleas from 1/3 as long as to equaling or slightly exceeding the lemmas, 2-veined at least on the lower portion, usually with hairs or both lemmas and paleas glabrous. | → 9 |
9. Calluses 1.5-6 mm long, sharply pointed; plants perennial or annual, if perennial, awns 65-500 mm long, if annual, awns 50-100 mm long; panicle branches straight. | → 10 |
10. Lower ligules densely hairy, upper ligules less densely hairy or glabrous; plants perennial | Jarava |
10. Ligules glabrous or inconspicuously pubescent, lower and upper ligules alike in vestiture; plants perennial or annual. | → 11 |
11. Plants perennial; florets 7-25 mm long; awns scabrous or pilose on the first 2 segments, the terminal segment scabrous, or if pilose, the hairs 1-3 mm long | Hesperostipa |
11. Plants annual or perennial, if annual, the florets 4-7 mm long and the awns not plumose, if perennial, the florets 18-27 mm long and the awns plumose on the terminal segment, the hairs 5-6 mm long | Stipa |
9. Calluses 0.1-2 mm long, blunt to sharply pointed; plants perennial; awns 1-70 mm; panicle branches straight or flexuous. | → 12 |
12. Florets usually dorsally compressed at maturity, sometimes terete; paleas as long as or longer than the lemmas and similar in texture and pubescence; lemma margins separate for their whole length at maturity | Piptatherum |
12. Florets terete or laterally compressed at maturity; paleas often shorter than the lemmas, sometimes less pubescent, sometimes as long as the lemmas and similar in texture and pubescence; lemma margins often overlapping for part or all of their length at maturity. | → 13 |
13. Glumes without evident venation, glume apices rounded to acute; plants subalpine to alpine, sometimes growing in bogs | Ptilagrostis |
13. Glumes with 1-3(5) evident veins or the glume apices attenuate; plants growing from near sea level to subalpine or alpine habitats, not growing in bogs. | → 14 |
14. Lemma bodies with evenly distributed hairs of similar length or completely glabrous, sometimes with longer hairs around the base of the awn; basal segment of the awns sometimes with hairs up to 2 mm long | Achnatherum |
14. Lemma bodies with hairs to 1 mm long over most of their length, with strongly divergent hairs 3-8 mm long on the distal 1/4, or the basal segment of the awns with hairs 3-8 mm long | Jarava |
1. Florets 6.5-22 mm long; culms 40-130 cm tall. | → 2 |
2. Lemmas hairy; awns 19-35 mm long | P. pringlei |
2. Lemmas glabrous; awns 40-120 mm long. | → 3 |
3. Florets 7-13 mm long; awns 40-75 mm long | P. avenaceum |
3. Florets 13.5-22 mm long; awns 62-120 mm long | P. avenacioides |
1. Florets 2.3-5.5 mm long; culms 20-95 cm tall. | → 4 |
4. Lemmas golden brown, hairy, the hairs easily rubbed off | P. fimbriatum |
4. Lemmas dark brown, glabrous. | → 5 |
5. Awns 10-16 mm long; blades 0.8-1.5 mm wide; distal margin of the lemma crowns straight | P. setosum |
5. Awns 15-25 mm long; blades 0.2-0.4 mm wide; distal margin of the lemma crowns sometimes slightly to strongly revolute | P. stipoides |
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