jcoastres-d-10-00142%2E1

7/30/2019 jcoastres-d-10-00142%2E1

1/9

www.cerf-jcr.org

The Biological Flora of Coastal Dunes and Wetlands:Batis maritima C. Linnaeus

Robert I. Lonard{, Frank W. Judd{, and Richard Stalter{

{

Department of BiologyThe University of Texas-Pan American

Edinburg, TX 78539-2999, U.S.A.

[email protected]

{

Department of Biological SciencesSt. Johns University

8000 Utopia Parkway

Queens, NY 11439, U.S.A.

ABSTRACT

LONARD, R.I.; JUDD, F.W., and STALTER, R., 2011. The biological flora of coastal dunes and wetlands: Batis maritimaC. Linnaeus. Journal of Coastal Research, 27(3), 441449. West Palm Beach (Florida), ISSN 0749-0208.

Batis maritima C. Linnaeus (maritime saltwort) is a New World subtropical and tropical trailing subshrub that formsdense colonies in salt marshes, brackish marshes, and mangrove swamps and frequently is found on the margins of saltpans and wind-tidal flats. It typically occurs at elevations less than 1.0 m above mean sea level and at sites where salinityranges from 18 to 50 ppt. Leaf succulence increases significantly in the dry season and leaves are shed, thereby reducingsalt-induced stress. Batis maritima occurs in sites normally subject to minimal sand coverage. However, wrack depositsstimulate growth. Maritime saltwort provides cover and nesting sites for some species of birds, but glucosinolatecompounds in shoots make the plants unpalatable to most large vertebrates, with the exception of marine iguanas in the

Galapagos Islands. Recently, B. maritima is proposed to have a major role in reducing ozone levels in the stratosphere.

ADDITIONAL INDEX WORDS: Saltwort, morphology, geographical distribution, habitats, communities, populationbiology, reproduction, geomorphological interactions, interactions with other species.

INTRODUCTION

Batis maritima C. Linnaeus (maritime saltwort) is a

subtropical and tropical trailing subshrub found in rapidly

changing habitats in coastal sites (Carlquist, 1978). It forms

dense colonies in salt marshes, brackish marshes, and

mangrove swamps and frequently is found on the margins of

salt pans and wind-tidal flats. The monogeneric Batis in the

family Bataceae is represented by two species:B. maritima andBatis argillicola P. Royen. Batis maritima is a New World

species that has an amphi-Pacific tropical disjunct distribution

pattern (Carlquist, 1978; Thorne, 1972), whereas B. argillicola

occurs in Papua, New Guinea, northern Australia, and

Indonesia (Carlquist, 1978; Ronse de Craene, 2005). Both

species are similar, but B. maritima is dioecious, whereas

B. argillicola is monoecious (Ronse de Craene, 2005).

TAXONOMY AND VARIATION

Name

Batis maritima C. Linnaeus; family Bataceae, synonyms: B.

californica J. Torrey, B. fruticosa W. Roxburgh, B. spinosa W.Roxburgh,B. vermicularis W.J. Hooker; maritime saltwort, sea

samphire, beachwort, turtleweed, pickleweed, barilla, lechuga

de mar, vidrillos, camphere, planta de sal, pajoocsim, xpajooc-

sim, akulakuli kai, herbe-a-crabes, verdolaga rosada.

Taxonomic Description

The following taxonomic description has been assembled

from Britton and Millspaugh (1962), Burger (1977), Correll and

Johnston (1970), DArcy (1976), Johnson (1935), Logier (1985),

Radford, Ahles, and Bell (1968), Ronse de Craene (2005), and

Stone (1993) and from live plants (R.I. Lonard and F.W. Judd,

personal observations, unless otherwise cited).

Seed Morphology

Little is knownabout seed production. Seeds are hard-walled

and lenticular or oblong, are less than 1 mm long, and weigh

less than 0.5 mg (Marcone, 2003). Little endosperm is present,

and the embryo is straight (Carlquist, 1978). Seeds have

germinated after floating in seawater for several months.

Seedling Morphology

Maritime saltwort seedlings seldom are found in nature

(Johnson, 1935). Each of the pair of cotyledons is fleshy,

whereas young leaves are thick, glaucous, and succulent and

are similar to mature leaves. The primary root branches earlyin development and is unbranched until the shoot is 10 cm or

more in height (Johnson, 1935).

Shoot Morphology

Plants are dioecious, perennial subshrubs 0.11.5 m tall

which form dense colonies. Stems are glabrous, ribbed, pale

green to greenish yellow, and strong scented; bark is peeling at

maturity. Carlquist (1978) noted that growth rings are absent

DOI: 10.2112/JCOASTRES-D-10-00142.1 received 19 September2010; accepted in revision 12 November 2010.Published Pre-print online 21 March 2011. Coastal Education & Research Foundation 2011

Journal of Coastal Research 27 3 441449 West Palm Beach, Florida May 2011
http://-/?-http://-/?-

7/30/2019 jcoastres-d-10-00142%2E1

2/9

in woody stem tissues, and xylem vessels are solitary or

grouped. Branching is alternate, and adventitious rootsform at

thenodes. Flowering shootsare erect or decumbent. Leavesare

opposite and sessile, and the stipules are vestigial; blades are

succulent, linear, linear-clavate, or linear-oblanceolate, gla-

brous, 34 angled, and 12 cm long; margins are entire, apices

acute or mucronate, and the bases are tapered to a translucent

membrane that overlaps the node (Figure 1).

Inflorescence

Unisexual flowers are borne on separate plants in short,

rounded,axillary conelike catkins. The staminate inflorescence

consists of 1030 pairs of decussate flowers, each subtended bya 2-lobed perianth.Staminate flowers have 4 exserted stamens

alternating with 4 petaloid staminodes. Anthers are ovoid,

dorsifixed, versatile, introrse and dehisce longitudinally;

filaments are thick and glabrous. Pistillate flowers are 412

in the inflorescence; the ovary is superior and consists of 48

united pistils; a single erect anatropous ovule is present in a

locule; pistils have 2 styles.

Fruits

Fruits are 12 cm long, succulent, and drupaceous or

multiple because of the fused pistils at maturity. The endocarp

is bony.

Variability

No varieties or ecotypes of B. maritima are recognized.

Chromosome Number

Maritime saltwort has a chromosome number of 2n 5 22

(Goldblatt, 1976). A reported chromosome number of 2n 5 18

by Goldblatt (1976) is probably in error.

GEOGRAPHIC DISTRIBUTION

Batis maritima occurs on tropical and subtropical coastlines

of North America, Central America, northern South America,

and the Caribbean Islands (Anderson and Alexander, 1985;

Balick, Nee, and Atha, 2000; Barker and Dardeau, 1930;

Britton and Millspaugh, 1962; Burger, 1977; Cafferty and

Monro, 1999; Clewell, 1985; DArcy, 1976; Debrot et al., 1999;

Eleuterius, 1990; Eleuterius and McDaniel, 1978; Espejel,

1986; Funket al., 2007; Gonzalezet al., 2008; Hester, Spalding,

and Franze, 2005; Johnson, 1935; Lonard, Judd, and Smith,

2003; Long and Lakela, 1971; Luttge et al., 1989; Macdonald

and Barbour, 1974; Moreno-Casasola and Castillo, 1992;

Morrison, 2002; Nelson, Goetze, and Lucksinger, 2001; Nelson

Sutherland, 2008; Pennings and Richards, 1998; Rico-Gray,

1990; Ronse de Craene, 2005; Sanchez-Carillo et al., 2009;

Sauer, 1982; Schmalzer and Hinkle, 1990; Schofield, 1984;

Stalter, 1972, 1973; Stalter et al., 1999; Standley, 1930; Stone,

1993; Thom, 1967; Wikelski and Wrege, 2000; Wilder, Felger,

and Romero-Morales, 2008; Wunderlin, 1982.). Maritime

saltwort has been reported as an invasive species in Hawaii,

where it displaces native species (Rauzon and Drigot, 2002;Wagner, Herbst, and Sohmer, 1990) (Figure 2).

RANGE OF HABITATS

Batis maritima often is an important species in salt marshes

and similar landscape features throughout its geographic

range. Salt marshes are characterized by a broad spectrum of

environmental gradients, which are reflected by conspicuous

species zonation. Mild conditions typically occur on the

terrestrial margins, and progressively more severe conditions

exist in salt pans and water-logged sites at lower elevations, to

the point where rooted vegetation does not survive. Pennings

and Callaway (2000) reported that B. maritima, Salicornia

virginica, and Distichlis spicata are dominant species in mid-

and higher level salt marshes in Georgia. Choi et al. (2001)

noted a similar zonation pattern in salt marshes in Florida.

Theyfound maritime saltwort in hypersaline salt barrens in the

transition zone between mid- and higher levels of the marsh.

Mangrove swamps, about 0.5 m above mean sea level and

subject to frequent tidal inundation and water-logged condi-

tions, often support a ground layer of dense colonies of B.

maritima on tropical coastlines (Lewis, 2005; Long and Lakela,

1971). In this harsh environment, maritime saltwort is absent

from sites where Avicennia germinans, Rhizophora mangle,

Conocarpus erecta, or Laguncularia racemosa form a closed

upper story canopy (U.S. Fish and Wildlife Service, 1993).

Monotypic stands ofB. maritima often occur on the margins

of barrensalt flats that are subject to flooding in thewet seasonof northern Venezuela (Medinaet al., 1989). Seasonal variation

in salinity is appreciable in this habitat.

Upper reaches of tidal flats, including mangrove margins to

occasionally inundated sandy or muddy soils in Sonora,

Mexico, and southern California, are characterized by succu-

lent halophytes, including B. maritima, S. virginica, and

Salicornia bigelovii (Morzaria-Luna et al., 2004; Wilder,

Felger, and Romero-Morales, 2008; Zedler, 1977). This habitat

is present about 60 to 70 cm above mean sea level.

Figure 1. Batis maritima: pistillate plants on the margins of a salt marsh,

Cameron County, Texas.

442 Lonard, Judd, and Stalter

Journal of Coastal Research, Vol. 27, No. 3, 2011

7/30/2019 jcoastres-d-10-00142%2E1

3/9

In the landscape of Padre Island, Texas, where freshwater

inflow is restricted, salt marshes are replaced by mostly barren

wind-tidal flats (Lonard, Judd, and Smith, 2003). Margins of

these wind-tidal flats and the margins of hurricane washover

fans provide a habitat for succulent halophytes, including B.

maritima, S. bigelovii, S. virginica, Sesuvium portulacastrum,

Suaeda linearis, and Blutaparon vermiculare (Lonard, Judd,

and Sides, 1978; Lonard, Judd, and Smith, 2003).

A salt pan habitat occurs inland on barrier islands in

Louisiana. Batis maritima and most of the succulent species

listed above occur on margins of these barren sites where

salinity exceeds 18 ppt (Hester, Spalding, and Franze, 2005).

Salt marshes and brackish marsh habitats on the Texas

mainland adjacent to South Padre Island are distinctly zoned

along a salinity gradient. Batis maritima is the dominant

species in the lowest elevations of both types of marsh habitats,where it occurs with Sporobolus virginicus. Monanthochloe

littoralis occurs in a narrow belt at intermediate elevations,

and Spartina spartinae and Borrichia frutescens are co-

dominants at higher elevations (Judd and Lonard, 2004).

Substrate Characteristics

Maritime saltwort occurs in low high marshes and middle

marshes on the South Carolina coastline, from Cape Island

southward, where the pH ranges from 5.2 to 7.0 (Stalter,

personal observations). Substrates consist of siliceous and

calcareous sands to wet mineral clay and sandy soils (Espejel,

1986; Stutzenbaker, 1999). On South Padre Island, Texas,

Judd, Lonard, and Sides (1977) found that 80.2% of sandparticle sizes ranged from 0.18 to 0.25 mm on the margins of

wind-tidal flats where maritime saltwort occurs and the depth

to the water table was 1315 cm below the soil surface.

Climatic Requirements

The geographic distribution of B. maritima extends from

33uN latitude at Cape Island, South Carolina, to about 3uS

latitude in northern Brazil. The northern distribution appears

to be influenced by the severity, duration, and frequency of

freezing temperatures. Many sites where maritime saltwort

occurs are subject to severe tropical storms.

PLANT COMMUNITIES

Batis maritima is an important species in a wide variety of

coastal plant communities in the tropics and subtropics at

elevations typically less than 1 m above mean sea level in the

transition zone from the mid- to high marsh. On Padre Island,

Texas, the low-growing vegetation that characterizes this zone

is referred to as a Succulent Halophyte Community. This plant

community includes a small set of genera that includes Batis,

Salicornia, Suaeda, and Blutaparon.

In the intertidal zone in the Bahamas, B. maritima often is a

conspicuous species in the Mangrove Community, where it

occurs with R. mangle, A. germinans, L. racemosa, and the

succulent S. virginica (Morrison, 2006). Morrison (2006) noted

that this community is frequently inundated at high tide.In Florida, the Intertidal Salt Marsh Community, High Salt

Marsh Community, and Buttonwood (Conocarpus erecta)

Community have species similar to those listed above but also

include A. germinans, S. virginica, R. mangle, B. frutescens,

and other salt-tolerant species (Anderson and Alexander,1985;

Rey, Crossman, and Kain, 1990).

In some sites in Louisiana, maritime saltwort occurs in the

Salt Marsh Community formerly dominated by Spartina

alterniflora. Spartina alterniflora has been eliminated in some

Figure 2. Geographic distribution ofBatis maritima in the United States

and Mexico (A) and Central America, South America, and Caribbean

Islands (B). Shaded states (A) and shaded countries (B) have documented

populations of the species. Caribbean islands supporting B. maritima are

identified with shading and in writing.

Batis maritima, Coastal Dunes, and Wetlands 443


7/30/2019 jcoastres-d-10-00142%2E1

4/9

7/30/2019 jcoastres-d-10-00142%2E1

5/9

Luttge et al. (1989) noted that succulence and sulfate concen-

trations doubled in maritime saltwort in the dry season.

Seasonal variation in the halophytic succulent zone is great

because of temporary flooding in the wet season followed by

extended drought during the dry season (Medinaet al., 1989). In

arid environments in south Texas, prolonged rainfall increases

root: shoot ratios in maritime saltwort, B. frutescens, and S.

linearis but decreases these ratios in the competing S. virginica(Dunton, Hardegee, and Whitledge, 2001). No allelopathic

chemicals have been reported for maritime saltwort (Ungar,

1998). Betacyanins (red-violet pigments) and betaxanthins

(yellow pigments) characteristic of the order Centrospermae

are not present in this species (Mabry and Turner, 1964).

Succulence in halophytes is a mode of countering deleterious

effects of salts by sequestering high salt concentrations in

vacuoles in leaves. The fresh weight of maritime saltwort is

99% water (Kuramoto and Brest, 1979). In Puerto Rico, leaf

succulence increases by as much as 1.5 times during the dry

season. Subsequently, leaves are shed, thereby effectively

reducing salt-induced stress. Zedler, Winfield, and Williams

(1980) noted that B. maritima leaves and stem segments

typicallyare dropped in late August in southern California saltmarshes as a result of aging and stress-related conditions.

Water balance is influenced further by nighttime transpiration

rates that significantly reduce predawn xylem pressure

potential (Donovan, Linton, and Richards, 2001).

Carbon isotope ratios indicate that Batis maritima is a C3plant in its mode of carbon fixation in the light-independent

reactions of photosynthesis (Luttge et al., 1989). Maritime

saltwort has year-round photosynthetic activity, but the winter

rate of carbon dioxide fixation is reduced. Although water

constitutes almost the entire fresh weight of leaves, water

usage for photosynthesis is relatively inefficient. Water

movement is accompanied by accumulation of salts (Antlfinger

and Dunn, 1983). With less water-containing dissolved salts

being transpired in winter, reduced transpiration rates play animportant role in balancing the energetics of maritime saltwort

in a hypersaline environment. Therefore, dilution of salts and

improved moisture relations probably account for overall

photosynthetic efficiency (Antlfinger and Dunn, 1979).

Batis maritima is proposed to have a major role in reducing

ozone levels in the stratosphere (Bill et al., 2002; Manley et al.,

2006; Rhew, et al., 2002). Maritime saltwort produces methyl

halides, which are the chief carriers of bromide and chloride

ions from the marine and terrestrial environments to the

stratosphere (Manley et al., 2006; Ni and Hager, 1998; Rhewet

al., 2002). Halide emission by B. maritima is temperature

influenced, and a small area of salt marsh dominated by

maritime saltwort can make an appreciable contribution to

total methyl halides emitted from a salt marsh (Manley et al.,2006).

Phenology

Flowering and fruiting cycles for maritime saltwort range from

April to November on South Padre Island, Texas (Lonard and

Judd, 1989).Massive amounts of fruits aredeposited in the strand

line of the Gulf of Mexico and bay-shore lines during winter on

South Padre Island (Lonard and Judd, personal observations).

POPULATION BIOLOGY

Perennation

Batis maritima is a long-lived perennial that roots at the

nodes. Shoots are intolerant to heavy burial by sand but

tolerate frequent to infrequent flooding by high tides. Plants

survive mild winters in the subtropics.

Population Dynamics

Natural disturbance and anthropogenic perturbations are

significant features of population dynamics in coastal ecosys-

tems (Judd and Lonard, 1987). Judd and Lonard (personal

observations) have noted occasional damage to maritime

saltwort colonies on South Padre Island on the margins of

wind-tidal flats because of off-road vehicular traffic. Batis

maritima populations decline as a result of shading by taller

mangrove species (Rey, Crossman, and Kain, 1990). Pennings

and Callaway (2000) noted that severanceof clonal segments of

maritime saltwort significantly reduced growth.

REPRODUCTION

Sexual Reproduction

Pollination and Fertilization

Batismaritima is self-incompatible because of thenature of its

unisexual flowers, which are borne on separate plants. Pollen is

disseminated as monads and is tricolporate, tricolporoidate, or

stephanocolporate with a granular surface (Rowe, 2006; Tobe

and Takahashi, 1995; Willard et al., 2004). The pertectate

feature of exine ultrastructure is likely related to wind

pollination (Carlquist, 1978). Flowers have no detectable odor.

Seed Production

Little is known about seed production. Apparently, germi-

nation can occur in propagules that have been floating in

seawater for several months.

Dispersal

Batis maritima is a widely distributed New World tropical

and subtropical species. Propagules are dispersed by water

(Lonard and Judd, 1980; Thorne, 1972). However, mammals,

birds, reptiles (iguanas), and crustaceans may disperse fruits

and seeds. Propagules arebuoyant in seawater and arecapable

of floating for a few months (Sauer, 1982).

Seed Bank and Seed SizeNo information is available on maritime saltwort seed banks.

LaPeyreet al. (2005) noted that seed banks often are unreliable

sources for salt marsh restoration projects. Seeds are less than

1 mm long and weigh less than 0.5 mg (Marcone, 2003).

Germination Ecology and Establishment of Seedlings

Little is known about germination requirements and the

establishment of B. maritima seedlings. However, Ungar



7/30/2019 jcoastres-d-10-00142%2E1

6/9

(1978) stated that seeds of most halophytes usually are

dormant at high salinities, with germination occurring at a

later time when salinities are lower. Martinez, Valverde, and

Moreno-Casasola (1992) reported that no dormancy mecha-

nisms other than light requirements and thermoperiod are

required for germination of most tropical maritime species in

Mexico. Johnson (1935) did not find seedlings in established

colonies of maritime saltwort in Jamaica. However, he foundthat seeds planted in equal mixtures of sand and potting soil

germinated quickly in a greenhouse. Growth was rapid when

seedlings were watered with brackish water (Johnson, 1935).

GEOMORPHOLOGICAL INTERACTIONS

Response to Burial

Sand accretion and sand scouring play a major role in

population dynamics of numerous coastal zone species. How-

ever, maritime saltwort colonies occur in areas normally

subject to minimal sand coverage. This species is absent in

unstable dune systems.

Wrack deposited by high tides normally kills low-growing

vegetation and initiates secondary succession. However, wrack

deposited in the high salt marsh in Georgia stimulates growth

ofB. maritima. Plants aretaller, leavesare more abundant and

larger, and total biomass is several times higher than in

maritime saltwort populations found in lower elevations in the

salt marsh (Pennings and Richards, 1998).

Role in Geomorphology

Vegetation on muddy tropical and subtropical shorelines and

the margins of salt flats and depressions is limited to a small

number of species. Batis maritima is an important colonizing

species after tropical storms and hurricanes in sites formerly

occupied by mangroves. This has implications related to global

warming and subsequent sea level rise. Maritime saltwort

colonies might be important in re-establishing mangroves in

the coastal zone of Florida. A trailing growth form anchored by

an extensive adventitious root system serves to stabilize

substrates. Avicennia germinans seedlings planted in the

mudflats among maritime saltwort colonies had significantly

lower mortality rates than seedlings planted in barren

mudflats. The dense root system of B. maritima increases

elevation slightly, favoring mangrove seedling survival (Mil-

brandt and Tinsley, 2006).

INTERACTION WITH OTHER SPECIES

Competition

Competition, disturbance, and the harsh environment are

the parameters that define spatial patterns in salt marsh plant

communities. Disturbance associated with impoundment of

salt marshes for mosquito control in Florida results in damage

to high-marsh vegetation. Avicennia germinans, B. maritima,

and S. virginica are replaced by monotypic stands ofR. mangle

(Rey, Crossman, and Kain, 1990). Recovery ofB. maritima and

S. virginica stands is associated with reduced water levels and

increased substrate aeration (Rey, Crossman, and Kain, 1990).

Salicornia virginica is a competitive dominant over B.

maritima and S. bigelovii in disturbed habitats in which soil

salinities are high (Rey, Crossman, and Kain, 1990). However,

S. virginica is limited by competition with B. maritima or S.

bigelovii, or a combination of both, in sites subjected to flooding

and reduced salinity (Zedler, 1977; Zedler, Winfield, and

Williams, 1980).

Maritime saltwort shows a broad tolerance range toenvironmental variation. Competition plays an important role

in zonation ofA. germinans mudflats in Mexico (Lopez-Portillo

and Ezcurra, 1987). As the life form of A. germinans changes

from a shrub to a tree, B. maritima decreases its presence.

Avicennia germinans displaces maritime saltwort because of

shading androot zone competition. Every unit of cover removed

from the overstory canopy results in nearly equal recovery of

maritime saltwort in the understory (Lopez-Portillo and

Ezcurra, 1987).

Batis maritima and R. mangle are invasive species in

Hawaiiancoastal wetlands, where they displace native species.

They colonize unexploited habitats and negatively affect water

quality and hydrology(Rauzonand Drigot, 2002). Dense stands

of maritime saltwort exclude shorebirds, including the feder-ally endangered Hawaiian stilt (Himantopus mexicanus

knudseni), from foraging. Control measures include plowing

heavily infested sites with amphibious assault vehicles

(Rauzon and Drigot, 2002).

Predation and Symbiosis

Salt marshes and mangrove swamps where maritime

saltwort is an important species provide habitat for 28 species

of mosquitoes in Florida (Hribar, 2005). Morrison (2002, 2006)

documented high populations of several ant species associated

with B. maritima colonies in the Bahamas. These sites were

frequently submerged by high tides.

Glucosinolates have been isolated from the Brassicaceae(mustards) and B. maritima (Goldblatt et al., 1976). These

organic compounds contain sulfur and nitrogen compounds,

which produce a bitter or sharp taste that is a natural defense

against herbivores. However, Johnson (1935) found that

rabbits browsed young shoots of maritime saltwort that had

been transplanted into a garden in Maryland. Seeds for these

transplants had been collected in Jamaica.

Koske (1988) found that obligate-symbiotic vesicular-arbus-

cular mycorrhizae (VAM) colonize B. maritima roots. VAM

improve phosphate nutrition and indirectly reduce water

stress in host plants. Koske (1988) suggested that the presence

of VAM fungi might have been important in the successful

colonization of numerous vascular plants in Hawaii, including

the invasive B. maritima.

RESPONSE TO WATER LEVELS

Batis maritima colonies are adversely affected by increased

flooding depth. Rey, Crossman, and Kain (1990) found that

high marsh vegetation in Florida including maritime saltwort

was damaged whenwater was impounded for mosquitocontrol.

Avicennia germinans, B. maritima, and S. virginica were

eliminated from the high marsh and replaced by R. mangle.

446 Lonard, Judd, and Stalter


7/30/2019 jcoastres-d-10-00142%2E1

7/9

Reduced water levels and increased aeration of the substrate

contributed to recovery of the original set of halophytes (Rey,

Crossman, and Kain, 1990).

ECONOMIC IMPORTANCE

Coastal Protection

Batis maritima can tolerateconditionsthat make it a pioneerspecies on newly available habitats. Harrington (2002) sug-

gested thatB. maritima is a good substrate-stabilizing species.

She found high root densities and greater root masses in

greenhouse-grown plants watered with 300 mg/L potassium

chloride. Establishment of colonies in tropical and subtropical

mudflatsresults in slightlyhigher elevations that are favorable

to colonization of A. germinans, which provides greater

protection from coastal erosion (Milbrandt and Tinsley, 2006).

Restoration ofB. maritima in southern California salt marshes

is enhancedby constructionof tidal creek networks.Tidal creek

margins are favorable sites for transplanting and growing

maritime saltwort clones (OBrien and Zedler, 2006).

Potential Agronomic Values

Maritime saltwort has been investigated as a potential crop

for brackish and saline soils (El-Haddad and Noaman, 2001;

Miyamoto, Glenn, and Olsen, 1996). As a halophyte, it produces

less biomass than conventional agricultural crops (El-Haddad

and Noaman, 2001). Seeds contain 17.3% crude protein and

25%oil, similar to safflower oil (Marcone,2003). Roots areused

as a sweetener for coffee by Comiaac Indians (Wilder, Felger,

and Romero-Morales, 2008). Leaves occasionally are added to

salads in Puerto Rico (Logier, 1990).

Wildlife Values

Batis maritima provides nutrition and cover for wildlife. Onone island in the Galapagos Island chain, a subpopulation of

the marine iguana (Amblyrhynchus cristatus) supplements its

diet of macroscopic marine algae withmaritime saltwort. These

marine lizards nearly double their body size when they are

compared with other marine iguanas that feed only on algae

(Wikelski and Wrege, 2000). The Cuban iguana (Cyclura

nubila) feeds on maritime saltwort leaves. Leaf fragments

have been identified in their scat (Beovides-Casas and

Mancina, 2006).

Maritime saltwort is an important species in the salt marsh

habitat of the marsh rice rat (Oryzomys palustris) in Texas and

in the habitat of the endangered silver rice rat (O. palustris

natator) in southern Florida (Abuzeineh et al., 2007; U.S. Fish

and Wildlife Service, 1993).Batis maritima provides cover and nesting sites for migra-

tory and wading birds. Endangered whooping cranes forage in

salt marshes and brackish marshes where maritime saltwort

occurs at the Aransas National Wildlife Refuge in Texas

(Campbell, 1996). Laughing gulls (Larus atricilla) nest in low-

growing maritime saltwort and S. portulacastrum colonies in

Florida (Kushland and White, 1977). In the Dutch West Indies,

B. maritima is the larval host of the butterfly (Brephidium

exilis) (Debrot et al. 1999).

Medicinal Uses

Batis maritima has been used in folk herbal medicine in

Puerto Rico to treat gout, eczema, psoriasis, rheumatism, blood

disorders, and thyroid disorders(Logier,1990). Standley (1930)

indicated that it is used to treat cutaneous infections in the

Yucatan peninsula of Mexico.

Potential Biological Control Agents

No biological control agents have been isolated from

maritime saltwort.

ACKNOWLEDGMENTS

We thank Kem Lowry, Glennis Lonard, and David Lonard

for technical assistance.

LITERATURE CITED

Abuzeineh, A.A.; Owen, R.D.; McIntrye, NE.; Dick, C.W.; Strauss,R.E., and Holsomback, T., 2007. Response of marsh rice rat(Oryzomys palustris) to inundation of habitat. Southwestern

Naturalist, 25(3), 313322.Anderson, L.C. and Alexander, L.L., 1985. The vegetation of Dog

Island, Florida. Florida Scientist, 48(4), 1534.

Antlfinger, A.E. and Dunn, E.L., 1979. Seasonal patterns of CO2 andwater vapor exchange of three salt marsh succulents. Oecologia, 43,249260.

Antlfinger, A.E. and Dunn, E.L., 1983. Water use and salt balance inthree salt marsh succulents. American Journal of Botany, 70(4),561567.

Balick, M.J.; Nee, M.H., and Atha, D.E., 2000. Checklist of theVascular Plants of Belize. Bronx, New York: New York BotanicalGarden Press, 246p.

Barker, H.D. and Dardeau, W.S., 1930. Flore DHat. Port-Au-Prince,Haiti: Departement de L Agriculture, 456p.

Beovides-Casas, K. and Mancina, C.A., 2006. Natural history andmorphometry of the Cuban iguana (Cyclura nubila Gray, 1831) inCayo Siju, Cuba. Animal Diversity and Conservation, 29(1), 18.

Bill, M.; Rhew, R.C.; Weiss, R.F, and Goldstein, A.H., 2002. Carbonisotope ratios of methyl bromide and methyl chloride emitted from acoastal salt marsh. Geophysical Research Letters, 29(4), 4p.

Britton, N.L and Millspaugh, C.M., 1962. The Bahama Flora. NewYork: Hafner Pu blishing Company, 695p.

Burger, W.C.,1977. FloraCostarricensis.Fieldiana,Botany, 40, 5455.

Cafferty, S. and Monro, A.K., 1999. Typification of Batis maritima L.(Bataceae). Novon, 9(4), 490.

Campbell, L., 1996. Endangered and Threatened Animals of Texas:Their Life History and Management. Austin, Texas: University ofTexas Press, 140p.

Carlquist, S., 1978. Wood anatomy and relationships of Bataceae,Gyrostemonaceae, and Stylobasiaceae. Allertonia, 5, 297330.

Choi, Y.P; Wang, Y.; Hsieh, Y-P., and Robinson, L., 2001. Vegetation

succession and carbon sequestration in a coastal wetland innorthwest Florida: evidence from carbon isotopes. Global Biogeo-chemical Cycles, 15(2), 311319.

Clewell, A.F., 1985. Guide to the Vascular Plants of the FloridaPanhandle. Tallahassee, Florida: University Presses of Florida,605p.

Correll, D.C. and Johnston, M.C., 1970. Manual of the VascularPlants of Texas. Renner, Texas: Texas Research Foundation, 1881p.

DArcy, W.G., 1976. Flora of Panama, Part IV, Family 55. Bataceae.Annals of the Missouri Botanical Garden. 63(3), 385388.

Debrot, A.O.; Miller, J.Y.; Miller, L.D., and Leysner, B.T., 1999. Thebutterfly fauna of Curacao, West Indies: 1996 status and long-termspecies turnover. Caribbean Journal of Science, 35(34), 184189.

Donovan, L.A.; Linton, M.J., and Richards, J.H., 2001. Predawn plantwater potential does not necessarily equilibrate with soil water



7/30/2019 jcoastres-d-10-00142%2E1

8/9

7/30/2019 jcoastres-d-10-00142%2E1

9/9

maritima L.) in regeneration of degraded mangrove forests.

Hydrobiologia, 568, 369377.

Miyamoto, S.; Glenn, E.P., and Olsen, M.W., 1996. Growth, water useand salt uptake of four halophytes irrigated with highly saline

water. Journal of Arid Environments, 32, 141159.

Moreno-Casasola, P. and Castillo, S., 1992. Dune ecology on theeastern coast of Mexico. In: Seeliger, U. (ed.), Coastal PlantCommunities of Latin America. San Diego, California: Academic

Press, pp. 309321.

Morrison, L.W. 2002. Island biogeography and metapopulationdynamics of Bahamian ants. Journal of Biogeography, 29, 387394.

Morrison, L.W., 2006. The ants of small Bahamian cays. BahamasNaturalist & Journal of Science, 1(2), 2732.

Morzaria-Luna, H.; Callaway, J.C.; Sullivan, G., and Zedler, J.B.,2004. Relationship between topographic heterogeneity and vegeta-tion patterns in a California salt marsh. Journal of VegetationScience, 14, 523530.

Nelson, A.; Goetze, J., and Lucksinger, A., 2001. A comparison of the

flora of North Padre Island to that of Matagorda Island, MustangIsland, and southern Padre Island, Texas. Occasional Papers,

Museum of Texas Tech University, 209, 123.

Nelson Sutherland, C.R., 2008. Catalogo de las Plants Vasculares deHonduras. Espermatofilas. Tegucigalpa, Honduras: Secretara deResources Naturales, 1576p.

Ni, X. and Hager, L.P., 1998. cDNA cloning of Batis maritima methylchloride transferase and purification of the enzyme. Proceedings of

the National Academy of Science, USA. 95, 12,86612,871.OBrien, E.L. and Zedler, J.B., 2006. Accelerating the restoration of

vegetation in a southern California salt marsh. Wetlands Ecologyand Management, 14, 269286.

Pennings, S.C. and Callaway, R.M., 2000. The advantages of clonal

integration under different ecological conditions: a community widetest. Ecology, 81(3), 709716.

Pennings, S.C. and Richards, C.L., 1998. Effects of wrack burial insalt-stressed habitats: Batis maritima in a southwest Atlantic salt

marsh. Ecography, 21(6), 630638.

Radford, A.E.; Ahles, H.E., and Bell, C.R., 1968. Manual of the

Vascular Plants of the Carolinas. Chapel Hill, North Carolina:University of North Carolina Press, 1183p.

Ramsey, E. III and Rangoonwala, K., 2005. Leaf optical property

changes associated with the occurrence of Spartina alternifloradieback in coastal Louisiana related to remote sensing mapping.

Photogrammetric Engineering and Remote Sensing, 71(3), 299311.

Rauzon, M.J. and Drigot, C., 2002. Red mangrove eradication and

pickleweed control in a Hawaii wetland, waterbird responses, andlessons learned. In: Veitch, C.R. and Clout, M.N. (eds.), Turning theTide: The Eradication of Invasive Species. Gland, Switzerland, and

Cambridge, U.K.: IUCNSSC Invasive Species Specialist Group.IUCN, pp. 240248.

Rey, J.R.; Crossman, R.A., and Kain, T.R., 1990. Vegetation dynamicsin impounded marshes along the Indian River Lagoon, Florida,USA. Environmental Management, 14(3), 397407.

Rhew, R.C.; Miller, B.R.; Bill, M.; Goldstein, A.H., and Weiss, R.F.,

2002. Environmental and biological controls of methyl halideemissions from a southern California coastal salt marsh. Biogeo-chemistry, 60, 141161.

Rico-Gray, V., 1990. Bataceae de la peninsula de Yucata n, Mexico.

Phytologia, 68(2), 108112.

Ronse de Craene, L.P., 2005. Floral developmental evidence for the

systematic position of Batis (Bataceae). American Journal ofBotany, 92(4), 752760.

Rowe, C., 2006. The Australasian Pollen and Spore Atlas User

Document. PalaeoWorks. Canberra, Australia: Australian NationalUniversity Technical Report 8, 30p.

Sanchez-Carillo, S.; Sanchez-Andres. R.; Alatorre, L.C; Angeler, D.G.;A lvarez-Cobelas, M, and Areola-Lizarraga, J.A., 2009. Nutrientfluxes in a semi-arid microtidal mangrove wetland in the Gulf ofCalifornia. Estuarine, Coastal and Shelf Science, 82, 654662.

Sauer, J.D., 1982. Cayman Islands Seashore Vegetation: A Study inComparative Biogeography. Berkeley, California: University ofCalifornia Press, 161p.

Schmalzer, P.A. and Hinkle, C.R., 1990. Flora and Threatened Plantsof John F. Kennedy Space Center, Florida. Washington, DC:

National Aeronautics and Space Administration, 68p.Schofield, E.K., 1984. Field Guide and Travel Journal: Plants of the

Galapagos Islands. New York: University Books, 160p.Stalter, R., 1972. The flora of Outer Otter Island, Colleton County,

South Carolina. Castanea, 37(4), 298300.Stalter, R., 1973. The flora of Turtle Island, Jasper County, South

Carolina, Castanea 38(1), 3537.Stalter, R.; Tamory, J.; Lynch, P., and Lockwood, B., 1999. The

vascular flora of Biscayne National Park, Florida. Sida, 18(4),12071226.

Standley, P.C., 1930. Flora of the Yucatan. Chicago, Illinois: FieldMuseum of Natural History, 492p.

Stone, W.J., 1993. Bataceae, saltwort family. In: Hickman, J.C. (ed.),

The Jepson Manual, Higher Plants of California. Berkeley,California: University of California Press, 1400p.

Stutzenbaker, C.D., 1999. Aquatic and Wetland Plants of the WesternGulf Coast. Austin, Texas: University of Texas Press, 465p.

Thom, B.G., 1967. Mangrove ecology and deltaic geomorphology:Tabasco, Mexico. Journal of Ecology, 55(2), 301343.

Thorne, R.F., 1972. Major disjunctions in the geographic ranges ofseed plants. Quarterly Review of Biology. 47(4), 365411.

Tobe, H. and Takahashi, M., 1995. Pollen morphology of Gyrostemo-naceae, Bataceae, and Koeberlinia. Journal of Plant Research, 108,283288.

Ungar, I.A. 1978. Halophyte seed germination. Botanical Review,44(2), 233264.

Ungar, I.A., 1998. Are biotic factors significant in influencing thedistribution of halophytes in saline habitats? Botanical Review,64(2), 176199.

U.S. Fish and Wildlife Service, 1993. Endangered and threatenedwildlife and plants; designation of critical habitat for the silver ricerat. Washington, DC: Federal Register, 58(167), 46,03046,034.

Wagner, W.L.; Herbst, D.R., and Sohmer, S.H., 1990. Bataceae.Saltwort Family. In: Manual of the Flowering Plants Hawaii,Volume I. Honolulu, Hawaii: University of Hawaii Press andBishop Museum Press, pp. 381382.

Wikelski, M. and Wrege, P.H., 2000. Niche expansion, body size, andsurvival in Galapagos marine iguanas. Oecologia, 124, 107115.

Wilder, B.T.; Felger, R.S., and Romero-Morales, H., 2008. Succulent

plant diversity of the Sonoran Island, Gulf of California, Mexico.

Haseltonia, 14, 127160.Willard, D.A.; Bernhardt, C.E.; Weimer, L.; Cooper, S.R.; Gamez, D.,

and Jensen, J., 2004. Atlas of pollen and spores of the FloridaEverglades. Palynology, 28, 175227.

Wunderlin, R.P., 1982. Guide to the Vascular Plants of CentralFlorida. Tampa, Florida: University Presses of Florida, 472p.

Zedler, J.B., 1977. Salt marsh community structure in the Tijuanaestuary, California USA. Estuarine and Coastal Marine Science,5(1), 3953.

Zedler, J.B.; Winfield, T., and Williams, P., 1980. Salt marshproductivity with natural and altered tidal circulation. Oecologia,44(2), 236240.



jcoastres-d-10-00142%2E1

Documents

Transcript of jcoastres-d-10-00142%2E1