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The Biology of Limonium Carolinianum (Walt.) Britt. (Plumbaginaceae) in Coastal Wetlands

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The Biology of Limonium Carolinianum (Walt.) Britt. (Plumbaginaceae) in Coastal Wetlands International Biology Review The Biology of Limonium carolinianum (Walt.) Britt. (Plumbaginaceae) in Coastal Wetlands Richard Stalter*, Robert I. Lonard**, and Frank W. Judd** *Department of Biological ABSTRACT Sciences, St. John’s University, 8000 Utopia Limonium carolinianum (Walt.) Britt. is a New World humid Parkway, Queens, NY 11439 continental, temperate, subtropical, and tropical maritime taxon. It is a long-lived perennial that is found in wet sand **Department of Biology, and clay mineral soils. This species is a salt tolerant halophyte The University of Texas – Rio where salinity values approach 30 parts per thousand. Grande Valley, Edinburg, TX Reproduction is primarily vegetative. However, seed 78539 production is estimated at 1,000 to 10,000 per individual plant. Also known as sea lavender, it has been suggested as a Corresponding Author: species that can be used for re-vegetation efforts. Dr. Richard Stalter Inflorescences of this species are often harvested for dried ([email protected]) floral arrangements. Keywords: Sea lavender, morphology, habitats, communities, population ecology, physiological ecology, economic values. Copyright 2016 KEI Journals. All rights reserved 1 International Biology Review INTRODUCTION statice. The following taxonomic description has been derived from Correll and Correll The genus Limonium (Plumbaginaceae) 1972; Flora of North America Editorial includes about 300 to 350 species, and is Committee 2005; Hamilton 1997; Lehman primarily represented in maritime or in arid 2013; Luteyn 1976, 1990; and Radford et al. environments in the northern hemisphere 1968. (Correll and Correll 1972; Mabberley 1997). Eight species have been reported in North Limonium carolinianum has a woody taproot America including four species that have typical of dicots that extend up to 30.5 cm escaped from cultivation in California (Flora below the soil surface (R. Stalter, personal of North America Editorial Committee 2005; observation). Plants are long-lived Luteyn 1990). Limonium carolinianum perennials. Stems are acaulescent. Primary typically occurs in brackish marshes, salt leaves are in a basal rosette. Petioles are marshes, salt meadows, in irregularly glabrous, 2.5 to 15 cm long, and narrowly flooded tidal wetlands, and on barrier island winged at the base. Blades are 5 to 30 cm beaches (Duncan 1974; Stutzenbaker 1999). long and 0.5 to 7.5 cm wide, glabrous and It tolerates salinity conditions ranging from leathery, and are spatulate-elliptic, lance- freshwater to 30 parts per thousand(ppt) elliptic, oblanceolate, or obovate. The blade (Stalter 1968). Inflorescenes are often margins are entire. harvested for dried flower arrangements (Baltzer et al. 2002a; Baltzer et al., 2002b; The inflorescence is a diffuse panicle that Richardson and King 2011). Herein, we extends up to 90 cm above the basal rosette. summarize important aspects of this species. One to 3 flowers are borne on one side of the inflorescence axis, and are subtended by 1.0 TAXONOMY AND VARIATION papery, secund bracts. The bracts are 2 to 6 mm long. The funnel-form calyx consists of Limonium carolinianum (T. Walter) N. 5 united, pale white sepals, 4 to 6 mm long. Britton is a member of the family The calyx is persistent, dry, membranous, Plumbaginaceae P. Miller. The taxon was and glabrous, but pilose along the veins. The published under the name Statice caroliniana corolla consists of 5 slightly united petals Walt.in 1788. A partial list of synonyms or with elongated claws. The petals are rejected names includes Limonium lavender to purple or occasionally white, and angustatum (A. Gray) Small, L. carolini- are 1 to 2 mm longer than the calyx. Five anum var. angustatum (A. Gray) S.F. Blake, stamens are present, and the filaments are L. carolinianum var. compactum Shinners, L. adnate at the base of the corolla. The pistils carolinianum var. nashii (Small) B. Boivin, have 5 styles, and are free at the base (Figure L. carolinianum var. obtusilobum (S.F. 1). The fruit is a one-seeded, indehiscent Blake) H.E. Ahles, L. carolinianum var. utricle 3 to 7 mm long. Seeds are brown, trichogonum (S.F. Blake) B. Boivin, L. lustrous, reticulate, spindle-shaped, and 3 to nashii Small, L. nashii var. angustatum (A. 4 mm long. Gray) H.E. Ahles, L. obtusilobum S.F. Blake, L. trichogonum S.F. Blake (Flora of North The genus Limonium is characterized by America Editorial Committee 2005; Luteyn, having significant biodiversity, and is 1976). Common names include sea lavender, taxonomically complex (Cortinhas et al. Carolina sea lavender, marsh rosemary, 2015). Luteyn (1976) concluded that L. American thrift, marsh root, canker root, carolinianum is a complex of 6 to 7 taxa seaside thrift, lavender thrift, ink root, and formerly recognized as varieties or separate Copyright 2016 KEI Journals. All rights reserved 2 International Biology Review species. He noted that morphological (Duncan 1974; Hill1986; Radford et al. variation is almost continuous across the 1968; Richards et al. 2005; Stalter, 1974; geographical range of the taxon. Hamilton Stalter et al. 1999a, 1999b). (1997) reported that genetically isolated populations occur within 5 km of each other In Mississippi, sea lavender also occurs in on the coastline of Rhode Island. He brackish, saline, and hypersaline coastal sites recognized the role of isolation by distance (Eleuterius and McDaniel 1978). In Texas in influencing ecological variation. Sea coastal wetlands, it occurs on the margins of lavender has a chromosome number of 2n = mostly bare wind tidal flats, in irregularly 36 (Flora of North America Editorial flooded salt marshes, on beaches, in wet Committee 2005). depressions in the secondary dune topographic zone, and on the margins of 2.0 GEOGRAPHIC DISTRIBUTION and hurricane washover channels (Judd et al. RANGE of HABITATS 1977; Lehman 2013; Nelson et al. 2000). Limonium carolinianum is distributed from 2.1 SUBSTRATE CHARACTERISTICS about 47.5°N latitude in Newfoundland and and CLIMATIC REQUIREMENTS Labrador, Canada, to Florida, and from the Gulf coast of Florida to about 24.5°N in Limonium carolinianum is found in highly Tamaulipas, Mexico (Lehman, 2013; Luteyn, stressful physical environments in coastal 1976, 1990; Rachlin et al. 2012; Stalter wetlands. Substrate conditions include wet, 1994; Stalter et al. 1986, 1990). This species tidal-influenced, sandy and clay mineral soils is also reported from Bermuda (Luteyn that are deficient in organic material, and in 1976). nitrogen, phosphorus, and potassium (Drawe et al. 1981; Proffitt et al. 2005; Stutzenbaker Sea lavender typically occurs in regularly or 1999). In South Carolina tidal marshes, irregularly flooded saline habitats 1.4 to Stalter (1968) found that sites where L. 1.7m above mean sea level at Georgetown, carolinianum occurs are flooded for one South Carolina (Stalter 1968). It is primarily hour daily by spring tides. Under more xeric found in saturated soil conditions in brackish conditions in irregularly flooded sites on marshes, salt marshes, and on the margins of South Padre Island, Texas, the water table on salt pans or salt flats. It is rarely a dominant the margins of the wind-tidal flats is 13 to 15 species. In Rhode Island and New cm below the soil surface (Judd et al. 1977). Hampshire, L. carolinianum occurs in The pH typically ranges between 6.0 to 8.5, regularly inundated upper middle and high and the salinity ranges from 0 to 30 ppt on marsh habitats, in saturated depressions in North Padre Island, Texas, and in South the secondary dune complex, and Carolina in saline wetlands (Drawe et al. occasionally on rocky beaches (Dunlop and 1981; Stalter 1968). Soil temperatures can Crow 1985; Hacker and Bertness 1999). reach 52.8 °C in North Carolina sites (Oosting 1954). In the mid-Atlantic states of Virginia and Maryland to subtropical sites in Florida, L. This species occurs over broad climatic carolinianum occurs in similar habitats listed conditions ranging from a humid continental above, but it is also found in irregularly climate in Newfoundland and Labrador to a flooded higher elevations in the salt marsh, subtropical climate in Tamaulipas, Mexico. on the margins of tidal flats, and on the In the broad continuum of climatic zones for margins of hurricane washover channels this species, plants survive unfavorable Copyright 2016 KEI Journals. All rights reserved 3 International Biology Review temperatures by producing a basal rosette, and by having a perennial caudex. Cold Dunlop and Crow (1985) and Stalter and tolerance studies have not been conducted. Lamont, (1990, 2000, 2002) recognized plant communities associated with habitats; 2.2 PLANT COMMUNITIES namely tidal marshes and salt marshes. All communities are in saline and hypersaline Plant communities in saline coastal wetlands sites and are subject to periodic flooding. where sea lavender is present are referred to Table 1 includes common species associated by a variety of inconsistently used with L. carolinianum throughout its range. terminology. Most workers name plant communities according to topographic zones 3.0 PHYSIOLOGICAL ECOLOGY in which they occur or by listing a habitat. For example, Judd et al. (1977) associated Limonium carolinianum is a salt tolerant plant communities of South Padre Island, halophyte that occurs where salinity values Texas, with topographic facets including approach 30ppt (Stalter 1968). Little is backshore, primary dunes, secondary dunes known about the physiology of this species. and vegetated flats, and tidal flats. They also Anatomical features include highly identified washovers as a site for a plant developed aerenchyma tissue in the root community. Judd et al. (1977) reported L. system and petioles (Lehman 2013). carolinianum on the margins of washover Aerenchyma tissue aids in diffusion of channels and in the mostly barren wind tidal oxygen to tissues that are otherwise flats associated with Salicornia bigelovii, subjected to anaerobic conditions. Salt Sarcocornia utahensis = Salicornia virgini- glands are present on leaves and stems. ca, Blutaparon vermiculare, Sporobolus Aerosol salt spray and salt crystals are often tharpii, and Sesuvium portulacastrum. found on leaves and stems.
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