| Mary E. Barkworth |
Plants perennial; cespitose from knotty bases or rhizomatous.
Culms 10-350 cm, erect, terete, solitary or in small to large clumps.
Leaves mostly cauline; sheaths open, smooth, sometimes striate;
ligules membranous, ciliate, cilia longer than the membranous bases;
blades flat or involute. Inflorescences terminal, usually exceeding
the upper leaves, 3-70 cm, panicles of 1-75 spikelike branches attached to an
elongate rachis; branches racemosely arranged, alternate, opposite, or
whorled, appressed to strongly divergent, axes 3-sided, spikelets usually sessile
on the 2 lower sides, usually divergent to strongly divergent; disarticulation
beneath the glumes. Spikelets laterally compressed, with 1 floret. Glumes
unequal, strongly keeled; lower glumes shorter than the florets, 1-veined;
upper glumes usually longer than the florets, 1-6-veined; lemmas
shorter than the paleas, 1-3-veined, midveins keeled, lateral veins usually
obscure; paleas thin, papery, 2-veined, obscurely keeled; anthers
3; lodicules sometimes present, truncate, vascularized; styles
2, plumose. Caryopses rarely produced. x = 10. Name from the Greek
spartine, a cord made from Spartium junceum, probably applied
to Spartina because of the tough leaves.
Spartina is a genus of 15-17 species, most of which grow in moist to
wet, saline habitats, both coastal and interior. Reproduction of all the species
is almost entirely vegetative.
There are nine native and two introduced species in the Flora region,
plus three hybrids, one of which is native, the other two being deliberate introductions.
One of the introduced species, S. maritima,
grows in both Europe and at a few locations in Africa; the African populations
may also represent introductions.
On the eastern seaboard of North America, the native species of Spartina
extend as far north as Nova Scotia, but the few species native to the
western seaboard do not extend north of California. Two species, S.
alterniflora and S. densiflora,
have, however, become established as far north as Washington and now
threaten the health of many coastal salt marshes and mud flats (see http://www.spartina.org/
or http://www.ecy.wa.gov/programs/sea/coast/plants/spartina.html). Spartina
alterniflora has also spread across southeastern China, "choking
estuaries, crowding out native grasses, and reducing feed and habitat
for migratory birds"
(Normile 2004).
Mobberley (1956), on whose work this treatment is based, described three groups
within Spartina, but did not give them formal recognition. Most of the
species in the Flora region belong to his third group, which he characterized
as having hard culms, scabrous leaf margins, more or less divergent inflorescence
branches, usually closely imbricate spikelets, and hispid keels on their glumes
and lemmas. Spartina alterniflora
and S. foliosa belong to Mobberley's
second group, the species of which have rather thick, succulent culms, glabrous
blades with smooth margins, and less closely imbricate spikelets than in the
other two groups. Spartina spartinae
is the only species of his first group to grow in the Flora region. Like
other members of the group, it has hard, slender culms, numerous short, closely
imbricate inflorescence branches, spikelets that are hispid or villous, at least
on the keels of the glumes, and no rhizomes.
Reeder (pers. com. 2003) pointed out that some of the
information in Mobberly (1956) has not been supported by subsequent work. In
particular, Reeder and Singh (1967) reported that lodicules are present in at
least three species (Spartina patens,
S. pectinata, and S. spartinae)
and several taxonomists have obtained different chromosome counts. Most of the
chromosome counts reported by Mobberly were multiples of seven, which led to
speculation that the genus belonged in the Pooideae,
but those obtained by other workers have all been multiples of 10. In the descriptions,
Mobberly's counts are listed in [square brackets]; those from other works are
unbracketed. Sources of the other counts are given under the notes
for Spartina and are included in the references listed below.
1 |
Leaf
blades with smooth or slightly scabrous margins (2) |
Leaf blades with strongly scabrous margins (6) |
|
Panicles branches 2-8
cm long, usually closely appressed and often twisted, the lower branches
evidently less closely imbricate than the upper branches; glumes usually
curved; plants of California and Baja California, Mexico ..... 3. S.
foliosa |
|
Panicle branches 2-24 cm long, usually
loosely appressed or divergent, usually not twisted, lower and upper branches
more or less equally imbricate; glumes straight; plants of varied distribution,
including California and Baja California, Mexico (3) |
|
Glumes usually mostly
glabrous on the sides, sometimes with appressed hairs; panicles with 3-25
branches ..... 2. S. alterniflora |
|
Glumes usually with appressed hairs on
the sides, the margins sometimes glabrous; panicles with 1-12 branches (4) |
|
Ligules 2-3 mm long;
anthers 5-13 mm long, well-filled, dehiscent at maturity ..... 6.
S. anglica |
|
Ligules 0.2-1.8 mm long; anthers 3-10
mm long, sometimes poorly filled and indehiscent at maturity (5) |
|
Ligules 0.2-0.6
mm long; leaf blades 6-12 cm long; anthers 3-6.5 mm long, well-filled,
dehiscent at maturity ..... 4. S. maritima |
|
Ligules 1-1.8 mm long; leaf blades 6-30
cm long; anthers 5-10 mm long, poorly filled and indehiscent at maturity ..... 5. S.
×townsendii |
|
Panicles
smooth in outline, with (6)15-75 tightly appressed panicle branches; branches
0.5-4(7) cm long; plants without rhizomes ..... 1. S.
spartinae |
|
Panicles not smooth in outline, with
2-67 tightly appressed to strongly divergent branches; branches 1-15 cm;
plants with more than 15 panicle branches always strongly rhizomatous,
those with less than 16 branches with or without rhizomes (7) |
|
Plants without rhizomes
or the rhizomes short; culms usually clumped; panicle branches 2-16 (8) |
|
Plants with well-developed rhizomes;
culms usually solitary, sometimes a few together; panicle branches 3-67 (10) |
|
Upper
glumes 1-veined ..... 9. S.
densiflora |
|
Upper glumes 3-4-veined (9) |
|
Spikelets 6-9 mm
long; culms to 200 cm tall; plants of the southeastern United States .....
7. S. bakeri |
|
Spikelets 10-17 mm long; culms to 120
cm tall; plants of the northeastern United States ..... 12. S.
×caespitosa |
|
Rhizomes
whitish; upper glumes 1-veined or with all lateral veins on the same side
of the keels; panicles with 2-15 branches, the branches 1-9 cm long (11) |
|
Rhizomes light brown to brownish-purple;
upper glumes 1-veined or with lateral veins on either side of the keels;
panicles with 3-67 branches, the branches 1.5-15 cm long (13) |
|
Spikelets 6-11 mm long,
ovate to lanceolate; inland plants of western North America, rarely found
east of Lake Winnipeg and the Mississippi Valley ..... 10. S.
gracilis |
|
Spikelets 7-17 mm long, linear-lanceolate
to ovate-lanceolate; usually coastal, also known from a few inland sites
in northeastern North America (12) |
|
Spikelets 7-12 mm long;
blade of the second leaf below the panicles 0.5-4(7) mm wide; plants of
disturbed and undisturbed coastal habitats from the Gulf of St. Lawrence
to the Gulf of Mexico and, as an introduction, on the west coast of North
America ..... 11. S. patens |
|
Spikelets 10-17 mm long; blade of the second
leaf below the panicles 2-7 mm wide; plants of disturbed habitats and
artificial wetlands from Maine to Maryland ..... 12. S.
×caespitosa |
|
Blade of the second
leaf below the panicles 2-5(7) mm wide, usually involute even when fresh;
panicles with 3-9 branches, the branches 3-9 cm long ..... 12. S.
×caespitosa |
|
Blade of the second leaf below the panicles
5-14 mm wide, flat when fresh; panicles with 5-67 branches, the branches
1.5-15 cm long (14) |
|
Lower glumes 3/4
as long as to equaling the adjacent lemmas; upper glumes awned, the awns
3-8 mm, with glabrous, rarely hispid, lateral veins ..... 13. S.
pectinata |
|
Lower glumes less than 1/2 as long to
2/3 as long as the adjacent lemmas; upper glumes unawned or with awns
up to 2 mm long, usually with hispid lateral veins ..... 8. S.
cynosuroides |
1. Spartina spartinae
(Trin.) Merr.
Gulf Cordgrass
Plants cespitose, not rhizomatous. Culms 40-200 cm, in large clumps,
hard, usually glabrous, nodes frequently exposed. Sheaths mostly glabrous,
throat glabrous, sometimes scabrous; ligules 1-2 mm; blades 1.5-4.5
mm wide, involute when fresh, abaxial surfaces glabrous, adaxial surfaces and
margins scabrous. Panicles 6-70 cm, smoothly cylindrical in outline, with
(6)15-75 branches, internodes shorter than the branches; branches 0.5-4(7)
cm, lower branches often longer than those above, all branches tightly appressed,
closely imbricate, with 10-60 spikelets. Spikelets 5-8(10) mm. Glumes
glabrous or hispidulous, keels hispid; lower glumes 2-8 mm, acuminate;
upper glumes 4-8(10) mm, acuminate to obtuse, keels hispid, lateral veins
1-2, if 2, these on either side of the keel; lemmas 5-6 mm, glabrous or
hispidulous, keels hispid over the distal 2/3, apices usually acuminate or apiculate,
rarely obtuse; anthers 3-5 mm, dark red to purple. 2n = 40 [28,
42].
Spartina spartinae grows from the Gulf coast through Mexico to Costa Rica
in North America and, in South America, in Paraguay and northern Argentina. In
the United States, it grows in sandy beaches, roadsides, ditches, wet meadows,
and arid pastures near the coast, the most inland collection being 60 miles from
the coast. In other parts of its range it sometimes grows well inland in saline
soils where Pinus palustris (longleaf pine) is dominant or co-dominant.
2. Spartina alterniflora Loisel.
Smooth Cordgrass, Spartine Alterniflore
Plants rhizomatous; rhizomes elongate, flaccid,
white, scales inflated, not or only slightly imbricate. Culms to 250
cm tall, (0.3)5-15(20) mm thick, erect, solitary or in small clumps, succulent,
glabrous, having an unpleasant, sulphurous odor when fresh. Sheaths mostly
glabrous, throat glabrous or minutely pilose, lower sheaths often wrinkled;
ligules 1-2 mm; blades to 60 cm long, 3-25 mm wide, lower blades
shorter than those above, usually flat basally, becoming involute distally,
abaxial surfaces glabrous, adaxial surfaces glabrous or sparsely pilose, margins
usually smooth, sometimes slightly scabrous, apices attenuate. Panicles
10-40 cm, with 3-25 branches, often partially enclosed in the uppermost sheath;
branches 5-15 cm, loosely appressed, not twisted, more or less equally
subremote to moderately imbricate throughout the panicle, axes often prolonged
beyond the distal spikelets, with 10-30 spikelets. Spikelets 8-14 mm,
straight, usually divergent, more or less equally imbricate on all the branches.
Glumes straight, sides usually glabrous, sometimes pilose near the base
or appressed pubescent, hairs to 0.3 mm; lower glumes 4-10 mm, acute;
upper glumes 8-14 mm, keels glabrous, lateral veins not present, apices
acuminate to obtuse, occasionally apiculate; lemmas glabrous or sparsely
pilose, apices usually acuminate; paleas slightly exceeding the lemmas,
thin, papery, apices obtuse or rounded; anthers 3-6 mm. 2n = 62 [56, 70].
Spartina alterniflora is found on muddy banks, usually of the intertidal
zone, in eastern North and South America, but it is not known from Central America.
In addition, it has become established on the west coast of North America, England,
southeastern France, and China. It hybridizes with
S. maritima in Europe, with S.
pectinata in Massachusetts, and with S.
foliosa in California.
The rhizomes and scales of S. alterniflora have large
air spaces, presumably an adaptation to the anaerobic soils of its usual habitat.
Decaploid plants tend to be larger than octoploids, but they cannot be reliably
distinguished without a chromosome count.
Spartina alterniflora is considered a serious threat
to coastal ecosystems in Washington and California. It out-competes many of
the native species in these habitats and frequently invades mud flats and channels,
converting them to marshlands. Pure S. alterniflora grows within the
lower elevational marsh zones in its native range but, in San Francisco Bay,
its hybrids with S. foliosa grow both
below and above the range of that species.
3. Spartina foliosa Trin.
California Cordgrass
Plants occasionally streaked or tinged with purple, rhizomatous; rhizomes
elongate, flaccid, whitish, scales inflated, not closely imbricate. Culms
to 150 cm tall, to 10 mm thick, erect, terete, solitary or in small
clumps, succulent, glabrous, often with adventitious roots from the lower
nodes, having an unpleasant, sulphurous odor when fresh. Sheaths mostly
glabrous, throats sparsely pilose, lower sheaths sometimes somewhat wrinkled; ligules 1-2
mm; blades 8-12 mm wide, flat to loosely involute, glabrous, margins
usually smooth, sometimes slightly scabrous, apices acuminate. Panicles 12-25
cm, with 3-25 branches, smoothly cylindrical, often partially enclosed
in the uppermost sheath; rachises twisted, glabrous; branches 2-8
cm, usually closely appressed and twisted, lower branches noticeably
longer and less closely imbricate than the upper branches, all branches
with axes rarely extending past the distal spikelets, with 8-30 spikelets. Spikelets 8-25
mm, usually appressed, often appearing twisted, those on the lower branches
usually less closely imbricate than those on the upper branches. Glumes usually
curved, sides and keels glabrous, scabrous, or hispid, apices acuminate
to obtuse or rounded; lower
glumes 6-12 mm; upper glumes 8-25 mm, 1-veined; lemmas glabrous
or sparsely appressed pubescent on the sides, keels glabrous, apices obtuse,
rounded or lobed; paleas slightly exceeding the lemmas, thin, papery,
glabrous, apices usually rounded, rarely acuminate; anthers 3-6
mm. 2n = 60 [56].
Spartina foliosa grows in the intertidal zone from northern California
to Baja California, Mexico. Populations in San Francisco Bay are threatened
by various introduced species of Spartina. Of particular concern
is S.
alterniflora, which forms hybrids with S.
foliosa that have a broader ecological amplitude than either parent.
In California, S. foliosa is often confused with S.
densiflora, which is also established in some regions, but S.
foliosa
differs from that species in being rhizomatous and having softer culms
and wider leaf blades.
4. Spartina maritima (Curtis) Fernald
Small Cordgrass
Plants rhizomatous; rhizomes with scales not inflated, not imbricate.
Culms to 80 cm, relatively soft, solitary or in small clumps. Sheaths
mostly glabrous, throat sometimes sparingly pilose, lower sheaths often wrinkled;
ligules 0.2-0.6 mm; blades 6-12 cm long, 5-8 mm wide, loosely involute,
disarticulating from the sheaths, abaxial surfaces glabrous, adaxial surfaces
glabrous or sparsely pilose, margins smooth. Panicles 4-14 cm, with (1)2-3(7)
branches; branches 2-11 cm, alternate, loosely appressed, not twisted,
lower and upper branches more or less equally imbricate, with 5-30 spikelets.
Spikelets 10-15 mm. Glumes straight, mostly appressed pubescent,
only the margins glabrous; lower glumes 7-10 mm, narrow, acuminate, obtuse,
or rounded; upper glumes 10-15 mm, acuminate (rarely obtuse); lemmas
mostly appressed pubescent, margins and basal portion of the keels glabrous, apices
acuminate; anthers 3-6.5 mm, well-filled, dehiscent at maturity. 2n
= 60 [56].
Spartina maritima is a European species that has been reported as growing
in Mississippi (Kartesz and Meacham 1999); the record has not been verified for
this treatment. It also grows in Africa, possibly as an introduction.
5. Spartina ×townsendii H. Groves & J. Groves
Townsend's Cordgrass
Plants rhizomatous; rhizomes whitish, scales not inflated, not closely
imbricate. Culms to 150 cm, relatively hard, solitary or in small clumps.
Sheaths mostly glabrous, throats pilose, lower sheaths often wrinkled;
ligules 1-1.8 mm; blades 6-30 cm long, 4-12 mm wide, diverging 20-45°
from the culms, flat proximally, involute distally, both surfaces glabrous, margins
smooth. Panicles 15-25 cm, with 2-10 branches; branches 4-24 cm,
loosely appressed, with 10-30 spikelets. Spikelets 16-22 mm. Glumes
mostly appressed pubescent, margins glabrous or sparingly hispidulous; lower
glumes 8-14 mm, linear, acuminate to obtuse; upper glumes 16-22 mm,
acuminate to obtuse; lemmas mostly pubescent, keels glabrous near the base,
margins glabrous throughout, apices obtuse to rounded or obscurely lobed; anthers
5-10 mm, poorly filled, indehiscent at maturity. 2n = 62,60+2.
Spartina ×townsendii is a sterile hybrid between the European S.
maritima and the American S. alterniflora.
It seems to have formed spontaneously at several locations in Europe, often taking
over the areas formerly occupied by its progenitors. At some locations it has
given rise to the fertile amphiploid S. anglica,
from which it differs morphologically in its narrower, less divergent upper blades,
shorter ligules, shorter, less hairy spikelets, and poorly filled, indehiscent
anthers. Spartina ×townsendii has been used throughout the world for tideland
reclamation because it is easy to establish, but it displaces native species.
6. Spartina anglica C.E. Hubb.
English Cordgrass
Plants rhizomatous; rhizomes elongate, flaccid, thick, whitish,
imbricate. Culms 30-130 cm, forming large clumps. Sheaths glabrous,
rounded dorsally; ligules 2-3 mm; blades 10-46 cm long, 6-15 mm
wide, persistent or deciduous, flat or involute, adaxial surfaces ridged, not
scabrous, margins smooth or slightly scabrous, sharply pointed, blades of upper
leaves strongly divergent. Panicles 12-40 cm, with 2-12, more or less equally
spaced branches; branches 16-25 cm, erect or somewhat divergent, axes pubescent,
extending up to 5 cm beyond the spikelets; disarticulation at the base
of the glumes, spikelets falling intact at maturity. Spikelets 14-21 mm
long, 2-3 mm wide, narrowly oblong, appressed, closely imbricate. Glumes
straight, sides appressed pubescent, keels ciliate or hispid, acute; lower
glumes 10-14 mm, 2/3-4/5 as long as the upper glumes, 1-veined; upper glumes
exceeding the floret, 3-6-veined; lemmas shorter than the upper glumes,
shortly appressed pubescent, 1-3-veined, acute; paleas a little longer
than the lemmas; anthers 5-13 mm, well-filled, dehiscent at maturity. 2n
= 122-124.
Spartina anglica is a naturally formed amphidiploid, derived from S.
×townsendii, that was first recognized as a separate species
in 1968. It has been introduced (like S. ×townsendii) for
reclamation of tidal mudflats. It differs from Spartina ×townsendii in its
wider and more widely divergent upper blades, longer ligules, longer, more hairy
spikelets, and longer, well-filled anthers.
7. Spartina bakeri Merr.
Sand Cordgrass
Plants cespitose, bases knotty, not rhizomatous. Culms to 200 cm,
in large, dense clumps, indurate, often branching from the lower nodes. Sheaths
smooth to striate, glabrous; ligules 0.5-2 mm; blades 10-50 cm long,
3-7 mm wide, usually involute, rarely flat, abaxial surfaces glabrous, adaxial
surfaces and margins scabrous, apices acuminate. Panicles 8-25 cm, usually
shallowly sinuous or lobed in outline, with 3-16 branches; branches 2-6
cm, usually appressed, moderately imbricate, axes glabrous, sometimes somewhat
scabrous on the angles, with 10-30 spikelets. Spikelets 6-9 mm. Glumes
with hispid keels and hispidulous margins, apices acuminate; lower glumes
3-6 mm, to 2/3 as long as the upper glumes; upper glumes 6-9 mm, hispidulous,
3-4-veined, lateral veins 2-3, prominent, on 1 side of the keel; lemmas mostly
glabrous, keels hispid, margins glabrous or hispid, apices acute to obtuse, sometimes
obscurely lobed; anthers about 5 mm, well-filled, dehiscent at maturity.
2n = 40 [42].
Spartina bakeri grows on sandy maritime beaches and other salt water sites
in the southeastern coastal states and on the shores of inland, freshwater lakes
in Florida. Its inflorescence is similar to that of S.
patens, but the branches of S. patens usually diverge from the
rachises at maturity, whereas those of S. bakeri remain appressed. Spartina
bakeri is distinct from most other species of Spartina in North America
in forming dense clumps and in being able to grow in freshwater habitats.
8. Spartina cynosuroides (L.) Roth
Big Cordgrass
Plants strongly rhizomatous; rhizomes elongate, purplish-brown or
tan, scales closely imbricate. Culms 100-350 cm tall, 1-2 cm thick, hard,
solitary or few together. Sheaths smooth to striate, mostly glabrous, throats
often densely pilose, lower sheaths often wrinkled; ligules 1-3 mm; blades
6-20 mm wide, flat or involute, glabrous on both surfaces, margins strongly scabrous,
apices acuminate, second blade below the panicles 5-15 mm wide, usually flat.
Panicles 15-40 cm, not smooth in outline, with 5-67 branches; branches
6-15 cm, usually spreading, with 10-70 spikelets. Spikelets 9-14 mm. Glumes
with hispid keels and hispidulous margins; lower glumes 3-7 mm, from less
than 1/2 as long as to 2/3 as long as the adjacent lemmas, linear, acute; upper
glumes 9-14 mm, usually more than twice as long as the lower glumes, exceeding
the florets, mostly glabrous or hispidulous, keels scabrous or hispid, trichomes
to 0.3 mm, 2 lateral veins prominent, 1 on each side of the keel, usually hispid,
apices unawned or awned, the awns to 2 mm; lemmas glabrous or hispidulous,
sometimes glabrous proximally and hispidulous distally, apices obtuse to rounded,
sometimes shallowly bilobed; anthers 4-6 mm, well-filled, dehiscent. 2n
= 40 [28, 42].
Spartina cynosuroides grows in brackish estuaries, tidal lagoons and bays,
and in maritime habitats bordering the strand and intertidal zones. It grows primarily
on the eastern and Gulf coasts of the United States, but has also been found in
Michigan, possibly introduced by shipping. Reports from South Dakota are based
on a misidentification.
9. Spartina densiflora Brongn.
Densely-Flowered Cordgrass
Plants cespitose, rarely rhizomatous; rhizomes, when present,
short, to 10 mm thick. Culms 27-150 cm, forming large clumps, indurate,
usually with short extravaginal shoots appressed to the culms. Sheaths
glabrous, lower sheaths smooth, indurate and shining, upper sheaths dull and
somewhat striate; ligules 1-2 mm; blades 12-43 cm long, 3-8 mm
wide, involute when fresh, abaxial surfaces glabrous, adaxial surfaces and margins
scabrous, apices acuminate. Panicles 10-30 cm long, 4-8 mm wide, sinuous
in outline, often twisted, with 2-15 branches; branches 1-11 cm long,
longer branches narrower than the shorter branches, all branches tightly appressed,
moderately imbricate, axes not prolonged beyond the distal spikelets, with 10-30
spikelets. Spikelets 8-14 mm. Glumes glabrous or sparsely hispidulous,
keels hispidulous, margins sparsely hispidulous; lower glumes 4-7 mm,
usually obtuse; upper glumes 8-14 mm, 1-veined, usually acuminate; lemmas
minutely hispidulous, keels glabrous proximally, hispidulous distally, apices
acuminate to obtuse; paleas acuminate, keels glabrous basally, hispidulous
distally; anthers 3-5 mm. 2n = 60.
Spartina densiflora is native to South America, where it grows in coastal
marshes and at inland sites. It was introduced to Humboldt Bay, Humboldt County,
California, possibly during the nineteenth century. It is now established there
and in several locations around San Francisco Bay and in Washington, Oregon,
and Texas, as well as the Mediterranean coast of Europe. In California, it has
often been mistaken for S. foliosa, from
which it differs in its indurate culms, narrow, inrolled leaves, and cespitose
growth habit and tendency to grow among Salicornia in the upper intertidal
zone or in open mud.
The chromosome count of 2n = 60 was obtained by Gerish (1979), who reported
it for Spartina foliosa, but Spicher
and Josselyn (1985) demonstrated that the plants he worked with were almost
certainly S. densiflora, a species that hitherto had been misidentified
as the native S. foliosa. In 2008, Ayres et al. (2008) and Fortune et al. (2008) reported that S. densiflora has 2n = 70.
10. Spartina gracilis Trin.
Alkali Cordgrass
Plants strongly rhizomatous; rhizomes elongate, 1.5-5 mm thick,
whitish, scales not inflated, closely imbricate. Culms 40-100 cm tall,
2-3.5 mm thick, usually solitary, erect, terete, indurate, glabrous. Sheaths
smooth or striate, mostly or completely glabrous, throats occasionally ciliate;
ligules 0.5-1 mm; blades 6-30 cm long, 2.5-8 mm wide, flat, becoming
involute, abaxial surfaces glabrous, adaxial surfaces scabrous, margins scabrous.
Panicles 8-25 cm, not smooth in outline, with 3-12 branches; branches
1.5-8 cm, alternate, differing only slightly in length and spacing within a
panicle, usually appressed, rarely spreading, with 10-30 spikelets. Spikelets
6-11 mm, ovate to lanceolate. Glumes with glabrous or sparingly hispidulous
margins, apices acute or mucronate; lower glumes 3-7 mm, sides narrow,
glabrous or sparsely pubescent, keels glabrous or strigose; upper glumes
6-10 mm, usually equaling the florets, keels strigose, hairs 0.2-0.5 mm, lateral
veins 2, inconspicuous, both on the same side of the keel; lemmas glabrous
or sparsely hirsute, keels hirsute, at least distally, hairs 0.3-1 mm, margins
sparsely hairy, apices obtuse to rounded, sometimes obscurely lobed; paleas
sparsely hispid distally, obtuse to slightly rounded; anthers 2.5-5 mm,
well-filled, dehiscent at maturity. 2n = 40 [42].
Spartina gracilis is found on the margins of alkaline lakes and along
stream margins and river bottoms. Its range extends from the southern portion
of the Northwest Territories, Canada, to central Mexico.
11. Spartina patens (Aiton) Muhl.
Saltmeadow Cordgrass, Spartine Étalée
Plants strongly rhizomatous; rhizomes elongate,
1-6 mm thick, whitish, scales not imbricate. Culms 15-150 cm tall, 1-6
mm thick, usually solitary, indurate. Sheaths glabrous or mostly glabrous,
throats occasionally short-pilose; ligules about 0.5(1) mm; blades
10-50 cm long, 0.5-4(7) mm wide, involute when fresh, abaxial surfaces glabrous,
adaxial surfaces scabrous, margins strongly scabrous; blade of the second leaf
below the panicles 4-40 cm long, 0.5-3(7) mm wide. Panicles 3-15 cm,
not smooth in outline even if the branches appressed, with 2-15 branches; branches
1-7 cm, alternate, differing only slightly in length and spacing within a panicle,
appressed to strongly divergent, with 10-30 spikelets. Spikelets 7-12
mm, linear lanceolate to ovate lanceolate. Glumes glabrous or sparsely
hispidulous on the sides, keels scabrous to hispidulous, trichomes (1) 1.5-2.5
mm, apices acuminate; lower glumes 3-8 mm, linear; upper glumes
7-12 mm, with 2 lateral veins, these on the same side of the keel, usually hispid,
apices acuminate, acute, or obtuse; lemmas mostly glabrous or sparsely
hispidulous, keels hispid distally, apices obtuse, rounded, or obscurely lobed;
anthers 3-5 mm, well-filled, indehiscent. 2n = 40 [28, 35, 42,
56].
Spartina patens grows in coastal salt and brackish waters. It is native
to the east coast of North and Central America, extending through the Caribbean
Islands to the north coast of South America, but is now established at scattered
locations on the west coast of Canada and the United States. On the east coast,
it is usually one of the dominant components of coastal salt marshes, frequently
extending from the dry, sandy beach above the intertidal zone well up into the
drier portions of the marshes. The older inland collections are from areas associated
with brine deposits or saline soils, but there is some indication that the species
range is increasing inland because of the use of salt to de-ice roads in winter.
The inflorescence of Spartina patens is similar to that
of S. bakeri when young, but its inflorescence
branches usually diverge at maturity, whereas those of S. bakeri remain
appressed.
Spartina patens is probably one of the parents of S. ×caespitosa,
S. pectinata being the other. Unlike
S. ×caespitosa, S. patens grows in both disturbed and undisturbed
habitats.
12. Spartina ×caespitosa
A.A. Eaton
Mixed Cordgrass
Plants rhizomatous or not; rhizomes, when present, thick, usually
purplish-brown, scales closely imbricate. Culms to 120 cm tall, 1-3 mm
thick, indurate, solitary or in small, dense clumps. Sheaths mostly glabrous,
throats glabrous or short-pilose; ligules 0.5-1 mm; blades 8-56
cm long, 2-6(7) mm wide, usually involute, abaxial surfaces glabrous, adaxial
surfaces glabrous or scabrous, margins strongly scabrous, blade of the second
leaf below the panicles 8-56 cm long, 2-5(7) mm wide. Panicles 9-20 cm,
not smoothly cylindrical, with 3-9 branches; branches 3-9 cm, appressed
or spreading, with 20-50 spikelets. Spikelets 10-17 mm, lanceolate to ovate-lanceolate.
Glumes glabrous or sparsely hispidulous, keels glabrous, hispid in whole
or in part, or ciliate; lower glumes 4-9 mm, acuminate or awned; upper
glumes 10-17 mm, exceeding the florets, keels hispid, lateral veins prominent,
1 on each side of the keel or 2-3 on 1 side of the keel, apices acuminate or awned;
lemmas glabrous or sparsely hispidulous, apices obtuse, rounded, obscurely
lobed, or apiculate; anthers 3-6 mm, poorly filled, indehiscent. 2n
= [42].
Spartina ×caespitosa
is found in disturbed areas of the drier portions of salt and brackish marshes,
at some distance above the intertidal zone. It occurs sporadically along the coast
from Maine to Maryland, a region where its putative parents, S.
pectinata and S. patens, are sympatric.
None of the populations Mobberley (1956) examined was growing in undisturbed land.
Mobberley's (1956) investigations led him to conclude that
the populations of S. ×caespitosa are polythetic in origin.
Part of the evidence for his conclusion was the variability he observed.
It is this variability that makes it necessary to bring out the hybrid
at several locations in the key. Its distribution is, however, very limited,
a fact that may be more useful for identification than any of the morphological
characteristics examined.c
13. Spartina pectinata Link
Prairie Cordgrass, Spartine Pectinée
Plants strongly rhizomatous; rhizomes elongate,
(2)3-8 mm thick, purplish-brown or light brown (drying white), scales closely
imbricate. Culms to 250 cm tall, 2.5-11 mm thick, solitary or in small
clumps, indurate. Sheaths mostly glabrous, throats often pilose; ligules
1-3 mm; blades 20-96 cm long, 5-15 mm wide, flat when fresh, becoming
involute when dry, glabrous on both surfaces, margins strongly scabrous, blade
of the second leaf below the panicles 32-96 cm long, 5-14 mm wide, usually involute.
Panicles 10-50 cm, not smooth in outline, with 5-50 branches; branches
1.5-15 cm, appressed to somewhat spreading, with 10-80 spikelets. Spikelets
10-25 mm. Glumes shortly awned, glabrous or sparsely hispidulous; lower
glumes 5-10 mm, from 3/4 as long as to equaling the adjacent lemmas, keels
hispid, apices awned; upper glumes 10-25 mm (including the awn), exceeding
the florets, glabrous or sparsely hispid, keels scabrous to hispid, trichomes
about 0.3 mm, lateral veins usually glabrous (rarely hispid), on either side
of, and close to, the keels, apices awned, awns 3-8 mm; lemmas glabrous,
keels pectinate distally, apices bilobed, lobes 0.2-0.9 mm; anthers 4-6
mm, well-filled, dehiscent. 2n = 40, 40+1,
80 [42, 70, 84].
Spartina pectinata is native to Canada and the United States, but it
has been introduced at scattered locations on other continents. On the Atlantic
coast, it grows in marshes, sloughs, and flood plains, being a common constituent
of ice-scoured zones of the northeast and growing equally well in salt and fresh
water habitats. In western North America, it grows in both wet and dry soils,
including dry prairie habitats and along roads and railroads.
Spartina pectinata is thought to be one of the parents
of S. ×caespitosa, the other parent
being S. patens.