| Stephen J. Darbyshire |
Plants perennial; cespitose, sometimes
shortly rhizomatous. Culms 7-130 cm, erect. Sheaths open to the
base, with tufts of hairs at the auricle position, sometimes with a line of
hairs around the collar; auricles absent; ligules of hairs; blades
rolled in the bud, flat or involute when dry. Inflorescences terminal;
panicles, racemes, or a solitary spikelet, to 12 cm; rachises, branches,
and pedicels scabrous or hirsute. Spikelets terete or laterally
compressed, with 3-12 florets, terminal floret reduced; disarticulation
beneath the florets, also at the cauline nodes in some species. Glumes
subequal or the lower glumes a little longer than the upper glumes, usually
exceeding the florets (excluding the awns and lemma teeth), lanceolate, chartaceous,
1-7-veined, keels glabrous or sparsely scabrous; rachillas glabrous;
calluses densely strigose on the sides; lemma bodies obscurely
(5)7-11-veined, backs glabrous or pilose, margins usually densely pilose proximally,
apices with 2 acute to aristate lobes, mucronate or awned between the lobes;
awns, when present, geniculate and twisted below the geniculation; paleas
about as long as the lemma bodies, 2-veined, veins scabrous, apices obtuse,
sometimes bifid; lodicules 2, glabrous or with a few hairs; anthers
3, their size depending on whether the florets are chasmogamous or cleistogamous;
ovaries glabrous. Caryopses 1.5-5.5 mm, ovate to obovate, dorsally
flattened, brown; hila linear, 1/3-3/4 as long as the caryopses. Cleistogenes
usually present in the lower sheaths, with 1(-10) florets, not disarticulating;
rachilla internodes about as long as or longer than the adjacent florets;
lemmas coriaceous, glabrous or scabrous near the apex, entire, unawned;
paleas sometimes slightly longer than the lemmas; anthers 3, minute;
ovaries glabrous; caryopses more linear than in the aerial florets.
x = 12. Named for the French botanist Étienne Danthoine, who worked
in the early nineteenth century.
Danthonia is interpreted here as a genus of about 20 species that are native
in Europe, North Africa, and the Americas. Of the eight species found in the Flora
region, seven are native and one is an introduction that is now established.
NOTE: In the key and descriptions, lemma lengths do not include the apical teeth. Callus characteristics are best seen on the middle to upper florets in the spikelet.
1 |
Lemmas mucronate, not awned ..... 1.
D. decumbens |
Lemmas not mucronate, with a twisted, geniculate awn (2) |
|
Calluses of the middle florets from shorter
to slightly longer than wide, convex abaxially; lemma bodies 2.5-6 mm long,
the backs usually pilose, occasionally glabrous or sparsely pilose (3) |
|
Awns 10-17 mm; hairs of the lemma margins
evidently increasing in length distally, longest hairs 2.5-4 mm long ..... 2.
D. sericea |
|
Awns 5-10 mm; hairs of the lemma margins not evidently
increasing in length distally, longest hairs 0.5-2 mm long (4) |
|
Lemma lobes 2-4 mm long, usually 2/3 or
more as long as the lemma bodies, aristate; lower inflorescence branches
usually flexible, divergent after anthesis; pedicels on the lowest inflorescence
branch as long as or longer than the spikelets; leaves not curling at maturity ..... 3.
D. compressa |
|
Lemma lobes 0.5-2 mm long, less than 2/3 as long as the
lemma bodies, acute to aristate; inflorescence branches stiff, appressed
to strongly ascending after anthesis; pedicels on the lowest inflorescence
branch from shorter than to equaling the spikelets; blades usually becoming
curled at maturity ..... 4. D. spicata |
|
Lower inflorescence branches (pedicels
if the inflorescence racemose) stiff, erect; pedicels from shorter than
to as long as the spikelets (6) |
|
Spikelets 1(-3), if 2-3, the inflorescence
a raceme; lemma bodies 5.5-11 mm; mature culms disarticulating at the nodes ..... 8.
D. unispicata |
|
Spikelets (4)5-10; lower inflorescence branches usually
with 2-3 spikelets; lemma bodies 3-6 mm; mature culms not disarticulating
at the nodes ..... 5. D. intermedia |
|
Uppermost cauline blades usually strongly
divergent or reflexed; inflorescences usually racemose; pedicels usually
much longer than the spikelets and usually strongly divergent; lemmas glabrous
or sparsely hairy over the back; mature culms disarticulating at the nodes
..... 7. D. californica |
|
Uppermost cauline blades usually erect to ascending; inflorescences
usually paniculate; pedicels shorter than to as long as the spikelets; lemmas
pilose over the back, at least basally; mature culms not disarticulating
at the nodes ..... 6. D. parryi |
1. Danthonia decumbens (L.) DC.
Mountain Heath-Grass, Danthonie Décombante
Culms 8-60 cm, usually erect, sometimes decumbent, not disarticulating.
Sheaths glabrous or pilose; blades 5-15 cm long, 0.5-4 mm wide,
usually flat, glabrous or sparsely pilose. Inflorescences with up to 15
spikelets; branches erect; lower branches with 1-3 spikelets. Spikelets
6-15 mm; florets usually cleistogamous, rarely chasmogamous. Calluses
of middle florets from as long as to a little longer than wide, convex abaxially;
lemma bodies 5-6 mm, margins glabrous or pubescent for most of their length,
scabrous apically, apices with acute teeth, teeth often scabrous, sometimes scabridulous,
mucronate, not awned, from between the teeth; palea veins swollen at the
base, forming pulvini; anthers of the cleistogamous florets 0.2-0.4 mm,
those of the chasmogamous florets about 2 mm. Caryopses 2.1-2.5 mm long,
1.1-1.8 mm wide. 2n = 24, 36, 124.
Danthonia decumbens grows throughout most of Europe, the Caucasus, and
northern Turkey, and is now established on the west and east coasts of North America.
It grows in heath lands, sandy or rocky meadows, clearings, and sometimes along
roadsides. The species is sometimes placed in the monotypic genus Sieglingia,
as Sieglingia decumbens (L.) Bernh.
2. Danthonia sericea Nutt.
Downy Oatgrass
Culms 50-120 cm, not disarticulating. Sheaths usually villous, occasionally
glabrous; blades 10-30 cm long, 2-4 mm wide, pilose or glabrous, usually
at least the adaxial surface pilose, uppermost cauline blades erect to ascending.
Inflorescences with (5)10-25(30) spikelets; lower branches erect
to ascending, with 2-6 spikelets; pedicels on the lowest branch from shorter
than to subequal to the spikelets. Spikelets 10-20 mm. Calluses of middle
florets from as long as to a little longer than wide, convex abaxially; lemma
bodies 4-6 mm, usually pilose over the back (sometimes glabrous), margins
densely pilose to beyond midlength, hairs evidently increasing in length distally,
longest hairs 2.5-4 mm, longer than apical teeth 2-4.5(5.5) mm, aristate; awns
10-17 mm; anthers to 2.6 mm. Caryopses 1.7-2.4 mm long, 0.8-1.2
mm wide. 2n = 36.
Danthonia sericea is restricted to the eastern United States. It grows
mostly on sand barrens and in open woods on dry soils. A less common form, with
glabrous foliage and lemma backs, is found in bogs, seepage areas, and low moist
areas adjacent to lakes and rivers and has been called D. sericea var.
epilis (Scribn.) Gleason or D. epilis Scribn. [For comments from
Quinn, click here].
Similar patterns of infraspecific variation are also seen in the leaf and lemma
vestiture of D. spicata and D.
californica, but the genetic basis of this variation is probably not
taxonomically significant.
3. Danthonia compressa Austin
Flattened Oatgrass, Danthonie Comprimée
Culms 40-80 cm, disarticulating at the nodes when mature.
Sheaths glabrous, rarely sparsely pilose, usually reddish above the nodes;
blades to 30 cm long, 2-4 mm wide, flexible but not curled at maturity,
glabrous, sometimes scabrous, uppermost cauline blades erect to ascending. Inflorescences
with (4)6-17 spikelets; branches usually flexible, usually divergent,
sometimes strongly so, after anthesis; lower branches with 2-3 spikelets;
pedicels on the lowest branch as long as or longer than the spikelets.
Spikelets (7)10-16 mm. Calluses of middle florets about as long
as wide, convex abaxially; lemma bodies 2.5-5 mm, pilose over the back,
sometimes sparsely so, margins pilose to beyond midlength, distal hairs 0.5-2
mm, apical teeth 2-4 mm, aristate, (1/2)2/3 or more as long as the lemma bodies;
awns 6-10 mm; anthers to 2.2 mm. Caryopses 1.7-2.6 mm long,
0.7-1.1 mm wide. 2n = 36.
Danthonia compressa grows in open and semi-shaded areas, including meadows,
open woods, and woodland openings. Although not a true pioneer species, it may
sometimes occur as a weed in perennial crops. It is restricted to eastern North
America.
4. Danthonia spicata (L.) P. Beauv. ex Roem. & Schult.
Poverty Oatgrass, Danthonie à Épi
Culms (7)10-70(100) cm, disarticulating at the nodes
when mature. Sheaths pilose or glabrous; blades 6-15(20) cm long,
0.8-3(4) mm wide, usually becoming curled at maturity, glabrous or pilose, uppermost
cauline blades erect to ascending. Inflorescences with 5-10(18) spikelets;
branches stiff, appressed to strongly ascending after anthesis; lower
branches with 1-3 spikelets; pedicels on the lowest branch from shorter
than to equaling the spikelets. Spikelets 7-15 mm. Calluses of middle
florets about as long as wide, convex abaxially; lemma bodies 2.5-5
mm, usually pilose (sometimes glabrous) over the back, margins pilose to about
midlength, longest hairs 0.5-2 mm, apical teeth 0.5-2 mm, acute to aristate,
less than 2/3 as long as the lemma bodies; awns 5-8 mm; anthers
to 2.5 mm. Caryopses 1.5-2(2.3) mm long, 0.7-1 mm wide. 2n = 31,
36.
Danthonia spicata grows in dry rocky, sandy, or mineral soils, generally
in open sunny places. Its range includes most of boreal and temperate North
America and extends south into northeastern Mexico.
Phenotypically, Danthonia spicata is quite variable,
expressing different growth forms under different conditions (Dore and McNeill
1980; Darbyshire and Cayouette 1989). Slow clonal growth, extensive cleistogamy,
and limited dispersal contribute to the establishment of morphologically uniform
populations, some of which have been given scientific names. For instance, D.
spicata var. pinetorum Piper is sometimes applied to depauperate
plants and D allenii Austin misapplied to more robust or second growth
plants (Dore and McNeill 1980).
Plants of shady or moist habitats often lack the distinctive
curled or twisted blades usually found on plants growing in open habits. Such
plants, which tend to have smaller spikelets and pilose foliage, have been called
D. spicata var. longipila Scribn. & Merr. The terminal inflorescence
is usually primarily cleistogamous, but plants with chasmogamous inflorescences
are found throughout the range of the species. Chasmogamous plants differ in
having divergent inflorescence branches at anthesis, larger anthers, and well-developed
lodicules.
5. Danthonia intermedia Vasey
Timber Oatgrass, Danthonie Intermédiaire
Culms 10-50(70) cm, not disarticulating at maturity. Sheaths usually
glabrous; blades 5-10 cm long, 1-3.5 mm wide, glabrous or slightly pilose.
Inflorescences with (4)5-10 spikelets; branches stiff, appressed
or strongly ascending; lower branches with (1)2-3(5) spikelets; pedicels
on the lowest branch shorter than the spikelets. Spikelets 11-15(19)
mm. Calluses of middle florets longer than wide, concave abaxially; lemma
bodies 3-6 mm, glabrous over the back, densely pilose along the margins,
teeth 1.5-2.5 mm, acute to acuminate or aristate; awns 6.5-8 mm; anthers
usually tiny, sometimes to 4 mm. Caryopses (2)2.3-3 mm long, 0.7-1.1
mm wide. Cleistogenes rarely produced. 2n = 36, 98.
Danthonia intermedia grows in boreal and alpine meadows, open woods,
and on rocky slopes and northern plains. Its range extends from Kamchatka, Russia,
to North America, south along the cordillera, and east, through boreal and alpine
regions, to Quebec and Newfoundland and Labrador. Its primarily cleistogamous
reproduction has probably facilitated its establishment and spread through more
boreal and alpine habitats than other members of the genus.
Tzvelev (1976) treats the American plants as Danthonia intermedia Vasey
subsp. intermedia and the Russian plants, which have 2n = 18,
as Danthonia intermedia subsp. riabuschinskii (Kom.) Tzvelev.
6. Danthonia parryi Scribn.
Parry's Oatgrass
Culms 30-80(100) cm, not disarticulating at maturity. Sheaths
glabrous or sparsely pubescent; blades 15-25 cm long, to 4 mm wide, glabrous
or scabrous (rarely pilose), uppermost cauline blades erect or diverging less
than 20° from the culm at maturity. Inflorescences usually paniculate,
sometimes racemose, with (3)4-11 spikelets; branches appressed to ascending,
somewhat flexible; pedicels on the lowest branch from shorter than to
as long as the spikelets. Spikelets 16-24 mm. Calluses of middle florets
longer than wide, concave abaxially; lemma bodies 5.5-10 mm, backs usually
pilose, especially near the base (rarely glabrous), margins pilose, teeth 2.5-8
mm, aristate; awns 12-15 mm; anthers to 6.5 mm. Caryopses
rarely produced, 3.5-5.2 mm long, 0.9-1.8 mm wide. 2n = 36.
Danthonia parryi is endemic to western North America and is often a major
component of grasslands on the eastern foothills of the Rocky Mountains. It
grows in open grassland, open woods, and rocky slopes, at elevations up to 4000
m. It rarely produces caryopses in the terminal inflorescences. This and its
somewhat intermediate morphology have led to speculation that it is derived
from hybridization between D. californica
and D. intermedia.
7. Danthonia californica Bol.
California Oatgrass
Culms (10)30-130 cm, disarticulating at the nodes at maturity. Sheaths
glabrous or pilose, upper sheaths usually glabrous or unevenly pilose; blades
10-30 cm long, (1)2-5(6) mm wide, flat to rolled or involute, glabrous or pilose,
uppermost cauline blades strongly divergent to reflexed at maturity. Inflorescences
usually racemose, with (2)3-6(10) widely-spreading spikelets; branches
flexible, strongly divergent to reflexed at maturity, pulvini usually present
at the base; pedicels on the lowest branch longer than the spikelets,
often crinkled. Spikelets (10)14-26(30) mm. Calluses of middle florets
usually longer than wide, concave abaxially; lemma bodies 5-10 mm, glabrous
or sparsely pilose over the back, margins pubescent (rarely glabrous), apical
teeth (2)4-6(7) mm, aristate; awns (7)8-12 mm; anthers to 4 mm.
Caryopses 2.5-4.2 mm long, 1.3-1.6 mm wide. 2n = 36.
Danthonia californica grows in prairies, meadows, and open woods. It
has a disjunct distribution, one portion of its range being located in western
North America, the other in Chile. An introduced population has been found at
Mansfield, Massachusetts.
Plants with pilose foliage have been called D. californica
var. americana (Scribn.) Hitchc. and plants with sparsely pilose lemma
backs D. californica var. macounii Hitchc., but the variation
does not appear to be taxonomically significant.
8. Danthonia unispicata (Thurb.)
Munro ex Vasey
One-Spike Oatgrass
Culms (10)15-30(42) cm, disarticulating at the nodes at maturity. Sheaths
usually densely pilose, hairs sometimes papillose-based (upper sheaths sometimes
glabrous); blades 3-8(20) cm long, 1-3 mm wide, both surfaces sparsely
to densely pilose, sometimes also scabrous or hirsute (rarely glabrous). Inflorescences
with 1-2(3) spikelets, if more than 1, racemose; pedicels stiff, appressed,
shorter than the spikelets. Spikelets (8)12-26 mm. Calluses of middle
florets longer than wide, concave abaxially; lemma bodies 5.5-11 mm,
glabrous over the back (rarely with a few scattered hairs), margins pilose (rarely
glabrous), apical teeth 1.5-7 mm, acute to aristate; awns 5.5-13 mm; anthers
to 3.5 mm. Caryopses 2.2-4 mm long, about 1 mm wide. 2n = 36.
Danthonia unispicata is restricted to western North America, where
it grows in prairies and meadows, on rocky slopes, and in dry openings
up to timberline in the mountains. It differs from D.
californica in its shorter stature, usually densely pilose foliage,
short, erect pedicels, and the usually erect cauline leaf blades. It is
closely related to D. californica,
and some authors prefer to treat it as Danthonia californica var. unispicata
Thurb.