The Biological Flora of Coastal Dunes and Wetlands: Schizachyrium littorale (G.

Nash) E. Bicknell
Author(s): Robert I. Lonard and Frank W. Judd
Source: Journal of Coastal Research, :654-662.
Published By: Coastal Education and Research Foundation
https://doi.org/10.2112/09-1180.1
URL: http://www.bioone.org/doi/full/10.2112/09-1180.1

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 Journal of Coastal Research 26 4 654–662 West Palm Beach, Florida July 2010

The Biological Flora of Coastal Dunes and Wetlands:

Schizachyrium littorale (G. Nash) E. Bicknell
Robert I. Lonard and Frank W. Judd

Department of Biology
The University of Texas–Pan American
Edinburg, TX 78541-2999, U.S.A.
[email protected]

ABSTRACT
LONARD, R.I. and JUDD, F.W., 2010. The biological flora of coastal dunes and wetlands: Schizachyrium littorale (G.
Nash) E. Bicknell. Journal of Coastal Research, 26(4), 654–662. West Palm Beach (Florida), ISSN 0749-0208.

Schizachyrium littorale (G. Nash) E. Bicknell is a New World tropical-, subtropical-, and temperate-zone rhizomatous
grass that is an important species on protected coastal beaches and in maritime grasslands. Herein, we present a
review of the biology of this important species. Schizachyrium littorale has low freshwater moisture requirements,
and it tolerates low nutrient levels, high sand temperature, and sand coverage. Also known as seacoast bluestem, it
inhabits mobile, semistable, and relatively stable dune systems where it plays an important role in stabilizing sec-
ondary dunes and barrier flats. Barrier island grasslands dominated by S. littorale recover quickly following wildfires.

ADDITIONAL INDEX WORDS: Seacoast bluestem, morphology, habitats, communities, population ecology, reproduc-
tion, geomorphological interactions, economic importance.

INTRODUCTION Gould, S. scoparium (A. Michaux) G. Nash subsp. littorale (G.

Nash) K. Gandhi & F. Smiens (Jones and Wipff, 2003); sea-
The grass genus Schizachyrium is represented by 60 spe-
coast bluestem, sea-coast little bluestem, maritime bluestem,
cies in the tropics, subtropics, and temperate zones. Nine spe-
shore bluestem, dune bluestem.
cies occur in North America north of Mexico (Wipff, 2003).
Hitchcock (1951) included the currently recognized genera Taxonomic Description
Schizachyrium, Bothriochloa, and Dichanthium as sections of
the genus Andropogon (Gould, 1967; Gould and Shaw, 1983). The following taxonomic description has been assembled
Schizachyrium littorale is distributed in coastal sand dunes from Gould (1975); Bruner (1987); Gandhi (1989); Lonard
adjacent to the Gulf of Mexico in Mexico and the United (1993); Hatch, Schuster, and Drawe (1999); Wipff (2003); and
States and in dune complexes on the Atlantic Coast (Castillo, from live plants (R.I. Lonard and F.W. Judd, personal obser-
Popma, and Moreno-Casasola, 1991; Gould, 1975; Wipff, vations, unless otherwise cited).
2003). The range of seacoast bluestem extends inland in
south Texas into a region referred to as the Coastal Sand Seed Morphology
Plain, where it is a dominant species in grasslands and sa- In S. littorale and in most other grasses, the seed coat and
vannas (Diamond and Fulbright, 1990). Seacoast bluestem is ovary wall (pericarp) are united at maturity, and they form
an important native forage grass, and it plays a significant a unique indehiscent, one-seeded fruit referred to as a cary-
role in stabilizing erosion-prone secondary dunes and vege- opsis. The caryopsis includes the embryo, a large volume of
tated flats. Many aspects of the ecology of S. littorale are endosperm, and the fused seed coat and pericarp (Gould and
unknown. Herein, we review the biology of this important Shaw, 1983).
species.
Seedling Morphology
TAXONOMY AND VARIATION Germination is initiated by the enlargement of the sheath-
Name like coleorhiza, which protects the primary root, and the ex-
pansion of the coleoptiles, which enclose the embryonic shoot
Schizachyrium littorale (G. Nash) E. Bicknell, family Po- system (Gould and Shaw, 1983). Initial leaf blades are similar
aceae (Gramineae), subfamily Panicoideae, tribe Andropogo- to those on mature culms; are linear-lanceolate, simple, al-
neae. Synonyms: Andropogon littoralis G. Nash, A. scoparius ternately arranged, and glabrous; and have a parallel vena-
A. Michaux var. littoralis (G. Nash) A.S. Hitchcock, S. sco- tion pattern with a prominent midvein.
parium (A. Michaux) G. Nash var. littorale (G. Nash) F.
Shoot Morphology
DOI: 10.2112/09-1180.1 received 1 January 2009; accepted in revision
3 February 2009. Plants cespitose and perennial with elongated rhizomes.
䉷 Coastal Education & Research Foundation 2010. Culms 40–150 cm, erect or decumbent, terete, but flattened
 The Biological Flora of Coastal Dunes and Wetlands 655

Figure 1. Schizachyrium littorale on the secondary dunes at South Padre Island, Texas, with Croton punctatus in the foreground.

near the base of the culms, glaucous, branching, and usually goneae tribe. A rame consists of a sessile and pedicellate
forming adventitious roots at the lower nodes. Leaf blades spikelet pair and the associated inflorescence axis or rachis.
15–30 cm long, 3–5 mm wide, flat or folded, glabrous, glau- Disarticulation occurs below the sessile spikelets. Therefore,
cous, and often purple tinged. Ligules 1.5–2 mm long, mem- the rame consists of a sessile fertile spikelet, a sterile pedi-
branous; auricles yellowish. Sheaths glabrous proximally but cellate spikelet, and the associated rachis. Peduncles are 0.5–
pubescent distally adjacent to the ligules. 5.0 mm long bearing a raceme, and rame units are delimited
by a sheathlike leaf at the base of the culm node.
Root Morphology Inflorescence axis or rachis with 10–20 spikelets, inter-
nodes densely pubescent with hairs 4–7 mm. Sessile fertile
Schizachyrium littorale has a fibrous root system typical of
spikelets 6–8 mm long, lance-ovate, glabrous; awns 1–2 cm;
monocots. Adventitious roots are formed at the lower nodes
pedicels pubescent. Glumes as long as the spikelets, with nu-
in plants that are partially covered by moving sand.
merous indistinct veins, glabrous, firm, and enclosing 2 mem-
branous florets. Pedicellate spikelets reduced to an axis that
Inflorescence
appears to be an extension of the pedicel. Stamens 3, anthers
The spikelet containing two florets is the basic unit of the 3.8–4.2 mm, linear, reddish purple (Figure 1).
numerous racemelike inflorescences in S. littorale (Gould and
Shaw, 1983). Terminal and numerous axillary racemes typi- Caryopsis
cally are produced in later summer and early fall. Complex
disarticulation units referred to as rames are present in the The caryopsis is tightly enclosed by the firm lower and up-
genus Schizachyrium and in other members of the Andropo- per glumes of the sessile spikelet.

Journal of Coastal Research, Vol. 26, No. 4, 2010

656 Lonard and Judd

Variability
Schizachyrium littorale was formerly recognized as a vari-
ety of the wide-ranging S. scoparium (A. Michaux) G. Nash
in North America. The S. scoparium complex is represented
by three intergrading varieties including var. divergens (E.
Hackel) F. Gould, var. scoparium, and var. stoloniferum (G.
Nash) J.K. Wipff (Wipff, 2003). Schizachyrium littorale differs
from S. scoparium by the presence of well-developed rhi-
zomes.

Chromosome Number
Schizachyrium littorale has a chromosome number of 2n ⫽
40 (Wipff, 2003).

GEOGRAPHIC DISTRIBUTION
Schizachyrium littorale occurs on tropical, subtropical, and
temperate coastlines of the eastern United States and the
Gulf of Mexico (Dahl et al., 1975; Dowhan, 1979; Fernald,
1950; Gandhi, 1989; Gillispie, 1976; Gleason and Cronquist,
1991; Hill, 1986; Hitchcock, 1951; Judd, Lonard, and Sides,
1977; McAvoy and Bennett, 2001; Negrete et al., 1999; Nel-
son, Goetze, and Lucksinger, 2001; Nelson et al., 2000; Oost-
ing and Billings, 1942; Schmalzer and Hinkle, 1990; Stalter,
2004; Stalter and Lamont, 1997, 2002; Tatnall, 1946; Wag-
ner, 1964) (Figure 2A). In south Texas, S. littorale occurs in-
land in sandy coastal prairies and in wooded openings (Dia-
mond and Fulbright, 1990; Gould, 1975; Hatch, Schuster, and
Drawe, 1999; Schuster and Hatch, 1990). Various authors
have reported seacoast bluestem from sand dunes adjacent
to Lake Michigan and Lake Erie in Illinois, Indiana, Penn-
sylvania, Ohio, and Ontario, Canada (Cooperrider, Cusick,
and Kartesz, 2001; Gould, 1975; Hitchcock, 1951; Morton and
Venn, 1990; Rhoads and Block, 2000; Wipff, 2003). However,
Gandhi (1989) concluded that populations referred to as S. Figure 2. The distribution of Schizachyrium littorale in (A) the United
littorale in dune sites adjacent to the Great Lakes are more States and (B) Mexico.
closely related to S. scoparium. On the Gulf Coast of Mexico
it ranges as far south as Tabasco (Castillo, Popma and Mo-
reno-Casasola, 1991; Loera and Moreno-Casasola, 1982; Mar-
barrier islands, and often it is the dominant or codominant
tı́nez and Moreno-Casasola, 1996; Martı́nez, Vázquez, and
species in the secondary dunes and vegetated flats (Carls,
Sánchez-Colón, 2001; Moreno-Casasola, 1988; Moreno-Casa-
Lonard, and Fenn, 1991; Castillo, Popma, and Moreno-Casa-
sola and Espejel, 1986; Moreno-Casasola et al., 1982; Sauer,
sola, 1991; Dahl et al., 1975; Judd, Lonard, and Sides, 1977;
1967) (Figure 2B).
Judd et al., 2007; Lonard and Judd, 1997; Lonard et al., 1999;
Martı́nez and Moreno-Casasola, 1996; Moreno-Casasola and
RANGE OF HABITATS
Espejel, 1986; Nelson et al., 2000). At Veracruz, Mexico, sea-
Schizachyrium littorale is an important species of the veg- coast bluestem is an important species in coastal dunes dom-
etation in unstable and stable dunes of New World tropical, inated primarily by perennial herbaceous vegetation. Sea-
subtropical, and temperate shorelines. In south Texas, sea- coast bluestem occurs in shifting, semimobile, and relatively
coast bluestem extends inland in the nearly level to gently stable dunes adjacent to diverse habitats that include tropical
rolling Coastal Sand Plain region of the Gulf Coast Prairies rainforests, mangroves, pastures, and agricultural fields
and Marshes vegetation zone, where it is the dominant spe- (Valverde and Pisanty, 1999).
cies in the savanna (Diamond and Fulbright, 1990; Diamond, The vegetation of South Padre Island, Texas, is topograph-
Riskind, and Orzell, 1987; Gould, 1975). ically zoned in conjunction with exposure to wind, aerosol salt
On the Gulf of Mexico coastlines of Texas and Mexico, S. spray, and depth to the water table (Judd, Lonard, and Sides,
littorale is a dominant species on the barrier islands in the 1977). Judd, Lonard, and Sides (1977) noted that seacoast
United States and on the barrier islands and mainland coast- bluestem was a minor species in the narrow backshore to-
al dunes in Mexico. It frequently grows in the backshore and pographic zone prior to Hurricane Allen in 1980. The back-
on the windward and leeward slopes of primary dunes on shore extends from the high tide line of the bare foreshore to

Journal of Coastal Research, Vol. 26, No. 4, 2010

 The Biological Flora of Coastal Dunes and Wetlands 657

the base of the primary dunes and is inundated infrequently Substrate Characteristics
by spring tides and storms. The storm surge from the hurri-
cane removed all vegetation from the backshore (Judd and Schizachyrium littorale occurs on sandy coastlines that
Sides, 1983). Seacoast bluestem has not been observed in have a narrow pH range of 8.4–8.6 (Moreno-Casasola et al.,
these narrow sites since 1980. However, S. littorale was found 1982). In the secondary dunes and vegetated flats on South
in swales in a relatively undisturbed backshore site (113.3 m, Padre Island where seacoast bluestem was the dominant spe-
n ⫽ 3 transects; see Judd et al., 2007). Therefore, it appears cies, 85.9% of the sand particle sizes ranged between 0.18 and
that the width and the occurrence of shallow microenviron- 0.25 mm. At a depth of 25 cm, the mean water content of
ments in the backshore zone are important factors in the dis- sand samples was 11%, and the mean depth to the water
tribution of this species in this harsh environment. Schiza- table was only 34 cm (Judd, Lonard, and Sides, 1977). Gandhi
chyrium littorale was a species of minor importance on the (1989) indicated that seacoast bluestem may occur in clay
salt spray–impacted windward faces of the primary dune loam and gravelly soils as well as in deep sands.
complex (Judd, Lonard, and Sides, 1977). On the more shel-
tered leeward primary dunes, seacoast bluestem was either Climatic Requirements
the dominant or codominant species (Judd, Lonard, and The geographical distribution of S. littorale extends from
Sides, 1977; Judd et al., 2007; Judd et al., 2008; Lonard et al., about 18.5⬚ N latitude at Frontera, Tabasco, Mexico (Sauer,
1999). In the stable and semistable dunes, the depressions, 1967) to about 41⬚ N latitude in Connecticut and Massachu-
and the nearly level topography of the secondary dunes and setts (Dowhan, 1979; Gleason and Cronquist, 1991). The
vegetated flats topographic zone, seacoast bluestem was the northern distribution appears to be influenced by the sever-
dominant species (Judd, Lonard, and Sides, 1977; Judd et al., ity, duration, and frequency of freezing temperatures. Aerial
2008; Lonard et al., 1999). Judd, Lonard, and Sides (1977) shoots are dormant during the winter months in the temper-
reported seacoast bluestem as an important species on the ate zone, and the perennial subterranean intercalary meri-
margins of tidal flats and hurricane washover sites where the stems in a rhizome system are protected by scalelike leaves.
vegetation typically consisted of salt-tolerant species. It oc- Gandhi (1989) found rhizomes up to 15 cm deep in Texas
curred also on the margins of disturbed roadsides (Lonard et coastal populations. In the subtropics, episodic freezes mea-
al., 1991). sured to be a few hours in duration have little or no effect on
Seacoast bluestem occurs in similar habitats on North Pa- exposed plant surfaces. Lonard and Judd (personal observa-
dre Island, Texas, due to the close proximity to South Padre tions) have noted vegetative growth during all seasons on
Island. Nelson et al. (2000) noted the importance of S. littor- South Padre Island, Texas.
ale on the protected slopes of the primary dune system. The
broad topographic facet of the secondary dunes and barrier
PLANT COMMUNITIES
flats, which are protected from aerosol salt spray, are domi-
nated by S. littorale on drier sites and by Paspalum monos- Plant communities on sandy tropical, subtropical, and tem-
tachyum in moister shallow depressions (Carls, Lonard, and perate shorelines in Mexico and the United States where sea-
Fenn, 1991; Dahl et al., 1975; Lonard et al., 2004; Nelson et coast bluestem occurs are influenced by offshore winds and
al., 2000). The codominance of S. littorale and P. monostach- sandy soils. These communities are present in moist depres-
yum indicates recovery from long-term, heavy grazing pres- sions or on nutrient-deficient elevated dry sites.
sure and a successional trend toward a midgrass prairie (Nel- From the literature relative to shoreline vegetation, it is
son et al., 2000). difficult to ascertain the position of seacoast bluestem in var-
Schizachyrium littorale is an important constituent of the ious topographic sites. Table 1 includes major species or in-
maritime grasslands of the mid-Atlantic states and North dicator species associated with S. littorale in selected sites
Carolina. At Bogue Bank, North Carolina, S. littorale is the without designation of topographic zones or plant communi-
dominant species on the lee slopes of primary dunes to the ties.
base of the secondary dunes (Wagner, 1964). It is common in Schizachyrium littorale is an important species in dunas
topographic facets that include dry foredunes, back dunes, costeras (coastal dunes) covered by herbaceous perennial veg-
and dry interdunes, and it is common in sites where there is etation on the coastlines of Mexico from Tamaulipas to Cam-
sparse vegetation (Maryland Department of Natural Resourc- peche and Tabasco (Moreno-Casasola and Espejel, 1986). The
es, 2004; Oosting and Billings, 1942; Stalter and Lamont, species is common in dry coppice dune and foredune com-
1997). At Beaufort Harbor near Fort Macon, North Carolina, munities dominated by Sporobolus virginicus and in humid
Oosting and Billings (1942) reported that S. littorale requires and damp interdune depressions where Fimbristylis spatha-
little moisture to survive and is confined to sheltered swales cea is the community indicator species (Moreno-Casasola and
where protection from salt spray is afforded by fore and rear Espejel, 1986).
dunes. Seacoast bluestem is the dominant species in shrub- The vegetation of North and South Padre Island, Texas, is
herbaceous zones of old leveled interdunes and backdunes on similar and is included in the Gulf Prairies and Marshes veg-
the Maryland coastline (Maryland Department of Natural etational area of Texas (Schuster and Hatch, 1990). The re-
Resources, 2004). Stalter and Lamont (2002) recorded S. lit- gion is part of the Gulf Coastal Plain Grassland Ecosystem
torale in fields on Long Island, New York, and Stalter (2004) that extends northward to the Mississippi Delta in Louisiana
has cataloged the species along an abandoned elevated rail and southward to the barrier island grasslands in northern
line in New York City. Tamaulipas. Schizachyrium littorale is a species of limited

Journal of Coastal Research, Vol. 26, No. 4, 2010

658 Lonard and Judd

Table 1. Species associated with Schizachyrium littorale. stands in moist depressions. Heterotheca subaxillaris is con-
spicuous on rolling dunes and in dry disturbed sites (Dia-
Species VCM SPI NPI TSP ATL
mond and Fulbright, 1990). These three species are consis-
Ambrosia psilostachya ⫻ ⫻ tent with the dominant species reported in the secondary
Ammophila breviligulata ⫻ dunes and vegetated flats on South Padre Island (Judd et al.,
Andropogon virginicus ⫻
Aristida sp. ⫻
2008).
Baccharis hamilifolia ⫻ Schizachyrium littorale is an important species in maritime
Centella asiatica ⫻ dune grassland plant communities in the Mid-Atlantic Coast-
Chamaecrista chamaecristoides ⫻ al Plain, including the barrier islands of North Carolina and
Cirsium horridulum ⫻
in the maritime vegetation of New England and New York.
Commelina erecta ⫻
Conoclinium betonicifolium ⫻ Seacoast bluestem occurs in communities dominated by pe-
Croton punctatus ⫻ ⫻ ⫻ ⫻ rennial grasses or perennial grasses and herbaceous dicots,
Dichanthelium acuminatum ⫻ or in a complex of shrubs and perennial grasses. Bunch grass-
Dichanthelium scoparium ⫻ es, including S. littorale, Andropogon virginicus, Panicum
Diospyros virginiana ⫻
Eupatorium hyssopifolium ⫻
amarum var. amarulum, Dichanthelium scoparium, and D.
Euthamia tenuifolia ⫻ acuminatum, typically are dominant species in the landscape.
Fimbristylis castanea ⫻ Representative herbaceous dicot species include Solidago
Fimbristylis spathacea ⫻ sempervirens, Cirsium horridulum, Eupatorium hyssopifolia,
Heterotheca subaxillaris ⫻ ⫻ ⫻ ⫻
Euthamia tenuifolia, and Opuntia humifusa. Sparsely occur-
Hudsonia tomentosa ⫻
Indigofera miniata ⫻ ring woody species may include Morella pensylvanica, Bac-
Ipomoea imperati ⫻ ⫻ charis halimifolia, Diospyros virginiana, and Prunus mariti-
Ipomoea pes-caprae ⫻ ⫻ ma (Maryland Department of Natural Resources, 2004).
Morella pensylvanica ⫻
Oenothera drummondii ⫻
Oenothera humifusa ⫻ PHYSIOLOGICAL ECOLOGY
Palafoxia lindenii ⫻
Palafoxia texana ⫻ Physiology
Panicum amarum ⫻
Panicum amarum subsp. amarulum ⫻ Schizachyrium littorale occurs where aerosol salt spray de-
Panicum capillarioides ⫻ position is low. Glaucous abaxial and adaxial leaf epidermal
Paspalum monostachyum ⫻ ⫻ ⫻
Paspalum plicatulum ⫻ ⫻
surfaces also may be an important factor in inhibiting expo-
Pectis saturejoides ⫻ sure to aerosol salts (Gandhi, 1989). The waxy leaf surfaces
Prunus maritimus ⫻ may be significant in reflecting high-intensity sunlight and
Schoenoplectus pungens ⫻ in reducing excessive evapotranspiration (Gandhi, 1989).
Sesuvium portulacastrum ⫻
Solidago sempervirens ⫻ ⫻
Oosting and Billings (1942) reported that daily spraying with
Spartina patens ⫻ seawater kills seacoast bluestem. However, S. littorale has
Sporobolus virginicus ⫻ low freshwater moisture requirements. It tolerates low nu-
Trachypogon plumosus ⫻ trient levels, high sand temperatures, and sand coverage.
Triplasis purpurea ⫻ ⫻
Long internodes form in mature plants under low nutrient
Uniola paniculata ⫻ ⫻ ⫻
Vitis rotundifolia ⫻ conditions (Valverde and Pisanty, 1999), and seedling growth
was slow in nutrient-deficient sands (Valverde, Pisanty, and
VCM ⫽ Vera Cruz, Mexico (Moreno-Casasola et al., 1982); SPI ⫽ South
Padre Island, Texas (Judd, Lonard, and Sides, 1977); NPI ⫽ North Padre
Rincón, 1997).
Island, Texas (Nelson et al., 2000); TSP ⫽ Texas Coastal Sand Plain (Di- Seacoast bluestem is a C4 plant in its mode of carbon fix-
amond and Fulbright, 1990); ATL ⫽ Atlantic coast (Maryland Depart- ation in the light-independent reactions of photosynthesis.
ment of Natural Resources, 2004). Leaf anatomy features are consistent with Kranz anatomy
(Gandhi, 1989).
importance in the Cenicilla–Beach Morning Glory (Sesuvium
portulacastrum–Ipomoea imperati) plant community in the Phenology
backshore topographic zone (Diamond, Riskind, and Orzell,
1987). However, seacoast bluestem is the dominant or codom- Flowering and fruiting cycles for seacoast bluestem on the
inant species in the secondary dunes and vegetated flats or coastline of Maryland and Virginia are from August through
barrier flats (Judd, Lonard, and Sides, 1977; Lonard et al., September (Hill, 1986). Spikelets are produced sporadically
1999; Lonard et al., 2004; Nelson et al., 2000). The community from April to September, and peak flowering follows rainfall
type is identified as the Seacoast Bluestem (Schizachyrium in October on South Padre Island. Flowering and fruiting
littorale) Series (Diamond, Riskind, and Orzell, 1987). continue through mid-December (Gandhi, 1989; Lonard and
In south Texas, the seacoast bluestem community occurs Judd, 1989). On the Mexican Gulf Coast at Campeche, S. lit-
in the Coastal Sand Plain subregion of the Gulf Prairies and torale produces caryopses from November to January, coin-
Marshes and extends 97 km inland in a broad arc from Baffin ciding with the winter storm season, and seedlings are es-
Bay to Port Mansfield. Seacoast bluestem is the dominant tablished in January and February after the storm season
species, whereas P. monostachyum usually forms dense (Martı́nez and Moreno-Casasola, 1996).

Journal of Coastal Research, Vol. 26, No. 4, 2010

 The Biological Flora of Coastal Dunes and Wetlands 659

POPULATION BIOLOGY and Judd, 1980). Rame units are dispersed during the dry
season (December–January) at Vera Cruz, Mexico (Valverde
Perennation
and Pisanty, 1999). Ghost crabs (Ocypode albicans) harvest
Schizachyrium littorale is a rhizomatous perennial that can and bury seacoast bluestem rames in their burrows. These
be long lived. Shoots tolerate heavy burial by sand. Rhizome sites are conducive for germination and establishment of
lengths vary between 10 cm and 65 cm in Texas populations seedlings (Ehrenfeld, 1990).
(Gandhi, 1989) and between 10 cm and 1.0 m in populations
from southern Mexico (Valverde and Pisanty, 1999). Adven- Seed Bank and Rame Weights
titious roots are present at the lower nodes. Aerial shoots die
No information is available on S. littorale seed banks. The
back during winter in temperate regions. However, plants in
mean weight of Texas seacoast bluestem rames is 1.4 mg
protected areas survive mild winters in the tropics and sub-
(Gandhi, 1989).
tropics.
Germination Ecology and Establishment of Seedlings
Population Dynamics
At Vera Cruz, Mexico, germination of S. littorale caryopses
Natural disturbance is a significant feature of population
occurs during the rainy season (June–September) during a
dynamics in coastal ecosystems. Judd et al. (1991) investi-
period of minimal sand movement (Martı́nez and Moreno-
gated resilience of seacoast bluestem communities on South
Casasola, 1996).
Padre Island to simulated hurricane effects. In vegetation re-
moval experiments, reestablishment of cover was 39.4% per
Vegetative Reproduction
year. However, the rate of recovery in sand coverage experi-
ments was much slower with increasing depths of sand cover. Rhizome formation is the mode of vegetative reproduction
Sand coverage at a depth of 30 cm resulted in only 6% recov- in cespitose populations of seacoast bluestem. Aerial shoots
ery per year. Saltwater exposure had only a slight impact on are produced at the subterranean nodes. Daughter colonies
S. littorale populations (Judd et al., 1991). produced from rhizomes were more prevalent in stable and
Fires initiated by lightning or by human activities occur semistable dunes than in highly disturbed mobile dunes at
annually on North Padre Island (Drawe and Kattner, 1978). Vera Cruz, Mexico. However, more aerial shoots (tillers) were
Lonard et al. (2004) reported that seacoast bluestem was the produced in unstable dunes (Valverde and Pisanty, 1999).
dominant species in both burned and nonburned sites two
years following a wildfire. It took only 106 days for S. littorale GEOMORPHOLOGICAL INTERACTIONS
to be restored as the dominant species.
Response to Burial
Seacoast bluestem populations also are resilient to all-ter-
rain vehicle (ATV) perturbations on the barrier islands. All Populations of S. littorale are resistant to sand coverage.
traces of ATV tracks through seacoast bluestem communities Martı́nez and Moreno-Casasola (1996) experimentally buried
on South Padre Island resulting from 1, 5, 10, and 20 repet- juvenile seacoast bluestem plants at depths where 50% and
itive passes were healed within one year (Judd et al., 1989). 100% of the meristems were covered. In the latter treatment
only the upper leaves extended above the sand. Biomass and
REPRODUCTION leaf area increased in both treatments over the controls
where no sand was added. Root/shoot ratios decreased as
Sexual Reproduction
more biomass was allocated to culms and leaves. It is unclear
Pollination and Fertilization how sand accretion stimulates shoot growth (Martı́nez and
Moreno-Casasola, 1996). No data are available on the impact
Schizachyrium littorale pollen is disseminated by wind.
of full plant coverage. Martı́nez, Moreno-Casasola, and Váz-
Seacoast bluestem, as well as other perennial grasses, are
quez (1997) reported that sand movements decreased with
cross-fertilized and self-sterile (Gould and Shaw, 1983).
increasing amounts of vegetative cover by S. littorale and as-
sociated species.
Caryopsis Production
Spikelets are paired in rame units. One pedicellate spikelet Role in Geomorphology
is reduced to an axis that appears to be an extension of a
Schizachyrium littorale plays a significant role in coastal
pubescent pedicel. The sessile spikelet contains a rudimen-
geomorphology by stabilizing barrier flats and secondary
tary floret and a bisexual floret enclosed within two firm
dunes. Seacoast bluestem inhabits mobile, semistable, and
walled glumes. The caryopsis is indehiscent at maturity.
stable dune systems (Day, Crawford, and Dilustro, 2001; Val-
verde and Pisanty, 1999).
Dispersal
The dispersal unit in S. littorale is a rame that consists of INTERACTION WITH OTHER SPECIES
a sessile fertile spikelet, the pedicel and rudimentary sterile
Competition
pedicellate spikelet, and the internode that extends between
two sessile spikelets (Barkworth, 2003). The villous-pubes- The harsh environment of North and South Padre Island
cent internode and pedicel allow for wind dispersal (Lonard has excluded competition of invasive species with seacoast

Journal of Coastal Research, Vol. 26, No. 4, 2010

660 Lonard and Judd

bluestem. Composition and structure of plant communities in low depressions. Exposed plants are killed by aerosol salt
coastal dunes are influenced by a mosaic of microenviron- spray. Therefore, this species affords little or no protection
ments with varying degrees of nutrients and organic detritus. from storm surges. However, seacoast bluestem is a domi-
In the closely related S. scoparium, a dominant species in the nant species in the protected secondary dunes and barrier
Mid-grass Prairie of North America, biomass and growth oc- flats, where it is important in stabilizing dunes and in pre-
cur in an optimum pH range of 5.44 to 6.07 and in response venting wind erosion.
to higher levels of phosphate ions (Jurena and Van Auken,
1996). Seacoast bluestem grows in more alkaline conditions. Medicinal Uses
Higher levels of phosphate ions may account for a competitive
No information is available about medicinal uses of this
advantage for S. littorale, but data are unavailable to support
species.
this hypothesis.

Predation and Symbiosis Potential Biological Control Agents

Reports of predation on S. littorale are limited. An unde- No biological control agents have been isolated from S. lit-
scribed subspecies referred to as the crystal skipper torale.
(Atrytonopsis loammi) feeds on seacoast bluestem leaves (A.
Leidner, personal communication). This butterfly taxon is en- ACKNOWLEDGMENTS
demic on barrier islands and on dredged material islands in
We thank David Lonard and Glennis Lonard for providing
Carteret and Onslow counties, North Carolina (H. Legrande,
assistance in the preparation of the manuscript. Ms. Jeanne
personal communication). Grasshoppers also may damage
Harrell, Acquisitions Librarian at Texas A&M University,
leaves in these coastal sites (A. Leidner, personal communi-
was helpful in locating reference materials. Bryan Connolly,
cation).
Massachusetts Natural Heritage and Endangered Species
Grazing animals readily consume seacoast bluestem. A
Program, furnished information about the distribution of sea-
large cattle industry was supported on North Padre Island
coast bluestem in Massachusetts and Connecticut. We thank
from the early 19th century until 1972, when all livestock
anonymous reviewers for their helpful suggestions.
were removed (Dahl et al., 1975; McAtee and Drawe, 1981).
Baumgardner (1991) and Goetze, Nelson, and French
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