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Overview

Comprehensive Description

White Mangrove (Laguncularia racemosa) is found along the shores of Florida and the West Indies, along both coasts of the American tropics, and in tropical West Africa (Little and Wadsworth 1964). It often occurs together with other mangrove species, such as Red Mangrove (Rhizophora mangle) and Black Mangrove (Avicennia germinans).

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The white mangrove, Laguncularia racemosa, is one of several species of trees known as mangroves that occur along coastlines worldwide. There are approximately 55 species of true mangroves in 20 genera (Hogarth 2007), and another 60 or more species of mangrove associates. Most species occur throughout the Indo-Pacific region. In the Indian River Lagoon, L. racemosa is one of three true species of mangroves commonly occurring along shorelines. The other two species are the red mangrove, Rhizophora mangle, and the black mangrove, Avicennia germinans.Laguncularia racemosa is a medium-sized tree or shrub, covered in thick, scaly bark, often reddish in color. The smooth, leathery leaves are up to 7 cm in length, opposite, with a silvery to yellow-green cast. Oval in shape and rounded at both apices, the leaves are often a distinguishing characteristic, differentiating L. racemosa from other mangrove species. White mangroves also exhibit unique glands called extra-floral nectaries found on either side of the stem at the leaf base. These structures excrete sugars which may attract ants that protect the plant from herbivorous insects (Hogarth 2007). Flowers are small and white, blooming at the leaf axils or branch tips. Fruits are about 2 cm in length, greenish with longitudinal ribs.
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Distribution

White Mangrove occurs along the shores of central and southern Florida, including the Florida Keys; in Bermuda and most of the West Indies; on both coasts of continental tropical America from Mexico south to Ecuador, northwestern Peru, and Brazil; and in western tropical Africa (Little and Wadsworth 1964)

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Range Description

The distribution of this species is restricted to the neotropics and West Africa (Tomlinson, 1986). It has been reported from the eastern tropical coasts of North and South America (ranging from Florida, U.S., 28°50', to Laguna, Brazil, 28°30') and all Caribbean Islands except Bermuda, Dominca and Netherlands Antilles; status on Anguilla is unknown (Wilkie and Fortuna 2003). It has been noted on the Pacific coast of South America from Estera Sargento, Mexico (29°17') south to Piura River, Peru (5°32') (de Lacerda, 2002). It is also noted from West Africa (Angola, Benin and Togo, Cameroon, Côte d'Ivoire, Democratic Republic of the Congo, Gabon, Gambia, Ghana, Guniea, Guinea-Bisau, Nigeria, Senegal, and Sierra Leone (Spalding et al. 1997). It is absent in the Galapagos Islands, Cocos, and Malpelo, and Canary Islands.

The south mid-Atlantic Island distributions should be confirmed.
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National Distribution

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

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Global Range: Shores of central and southern Florida including Florida Keys, Bermuda, and nearly throughout West Indies from Bahamas and Cuba to Trinidad and Tobago and Dutch West Indies. On both coasts of continental tropical America from Mexico south to Ecuador and northwestern Peru and to Brazil. The most widely distributed of the mangrove species in Puerto Rico (Little and Wadsworth 1964).

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Laguncularia racemosa occurs in tropical and subtropical regions throughout the world. Globally, the species ranges from Mexico, the West Indies to Brazil, through Central America to Peru, South America to Ecuador, and West Africa from Senegal to Angola (Exell 1958). In Florida, white mangroves share similar geographical limits with the red mangrove, Rhizophora mangle, having been reported as far north as Cedar Key on the west coast (Rehm 1976), and as far north as Ponce de Leon Inlet on the east coast (Teas 1977). Large populations can be found south of Cape Canaveral on Florida's east coast and around Tarpon Springs on the west coast (Odum & McIvor 1990). White mangroves occur throughout the Indian River Lagoon well above the high tide line, generally upland of other mangroves and associated species. However, they can be found intermingled with the black mangrove, Avicennia germinans. Their distribution is often patchy and predominantly occurs in the higher marsh areas (Ball 1980) along the lagoon, including spoil islands, tidal creeks and mosquito impoundments.
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Physical Description

Morphology

White Mangrove has opposite, leathery, slightly fleshy, elliptic leaves 4 to 10 cm long and 2 to 5 cm wide, rounded at both ends, dull yellow green on both sides and borne on reddish petioles (leaf stalks) with two raised gland dots near the apex. The numerous small, stalkless, bell-shaped whitish flowers are about 0.5 cm long and borne in terminal and lateral clusters 5 to 10 cm long. The clustered fruits are velvety gray-green and slightly pear-shaped, 1.5 to 2 cm long, flattened and ridged. (Little and Wadsworth 1964)

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Size

Little is known regarding typical age to maturation for mangroves in south Florida, though the typical size of mature L. racemosa can reach or exceed 15 m in height (Odum & McIvor 1990).
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White Mangrove commonly reaches 12 meters in height, sometimes more (Little and Wadsworth 1964).

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Look Alikes

Lookalikes

In their large area of geographic overlap, White Mangrove (Laguncularia racemosa) and Black Mangrove (Avicennia germinans) both have erect breather roots (pneumatophores) protruding up out of the water, but those of White Mangrove are fewer in number, wider, and more often branched. White Mangrove grows landward of Red Mangrove (Rhizophora mangle), which has conspicuous prop roots, and Black Mangrove. It often occurs onshore with Buttonwood (Conocarpus erectus, like White Mangrove a member of the plant family Combretaceae), which may be easily distinguished from White Mangrove by the fact that Buttonwood has alternately arranged leaves and leafstalk glands that are less prominent than those of White Mangrove. (Petrides 1988) The position and appearance of the glands on both the petiole and on the leaf blade differ conspicuously between White Mangrove and Buttonwood (see images on this page and on the Buttonwood page).

An excellent resource for identifying the mangroves of Florida can be found at http://www.selby.org/

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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
This species tends to be found at the upper margins of the mangrove-upland interface, or high intertidal region, and not at the seaward margin (Tomlinson 1994, Sherman et al. 2001). Seedlings are less tolerant of salinity and changing hydroperiod than R. mangle (Cardona-Alarte et al. 2006). Although it is a pioneer and can establish in relatively open sites with low salinity and abundant nutrients, mortality of seedlings is nearly 100% (Tomlinson 1995).

However, more important is the role of mangroves as nurseries for juvenile phases of economically-important fish and crustaceans. Laguncularia, being a landward mangrove, contributes to this indirectly by buffering upland pollutants.

Systems
  • Terrestrial
  • Marine
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Depth range based on 4 specimens in 1 taxon.

Environmental ranges
  Depth range (m): 1 - 1
 
Note: this information has not been validated. Check this *note*. Your feedback is most welcome.

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Trophic Strategy

Mangrove forests typically show a wide range of productivity, depending on factors such as hydrological regimes, nutrient supply, etc., and are considered to be vital sources of organic matter for estuarine systems. An average of 2 to 3 g dry weight of leaf litter is produced by mature mangrove forests each day (Odum et al. 1982). This litter, consisting of twigs, leaves, bark, fruit and flowers, is broken down by bacteria and consumed by a wide variety of fauna inhabiting mangrove ecosystems. Litter fall occurs throughout the year in Florida, peaking at the beginning of the summer wet season and after periods of stress (Heald 1969, Pool et al. 1975, Twilley et al. 1986).Competitors: In addition to propagule dispersal, Ball (1980) suggested that competition among the three mangrove species may be partially responsible for the zonation observed in many mangrove areas. White mangroves thrive throughout intertidal areas in the absence of large numbers of red and black mangroves. However, white mangroves appear to dominate in higher areas because of some competitive advantage over red mangroves. Direct consumers of mangrove propagules in Florida include the mangrove root crab (Goniopsis cruentata), the swamp ghost crab (Ucides cordatus), the coffee bean snail (Melampus coffeus) and the ladder hornsnail (Cerithidea scalariformis). Consumers of mangrove leaves include G. cruentata, the mangrove tree crab (Aratus pisonii), the blue land crab (Cardisoma guanhumi) and various types of insects.
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Associations

Mangroves form intertidal forests in which red mangrove prop roots, black mangrove pneumatophores, and their associated peat banks serve as the dominant intertidal substrata for other members of the mangrove community. All three species are commonly found in association with one another. However, segregation of the species does occur, with red mangroves typically occupying the lowest intertidal position. Black and white mangroves occur at slightly higher tidal elevations. White mangroves can be distinguished from the other species by leaf shape, the presence of extra-floral nectaries and the lack of either pneumatophores or prop roots that occur in black and red mangroves, respectively. In addition to other mangrove species, the buttonwood, Conocarpus erecta, can be found in the landward edge of L. racemosa stands. Several species of flora and fauna, including epiphytic plants, insects, birds, reptiles and mammals occur in and around white mangroves.
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Gilbert and Sousa (2002) studied the host-associations of wood-decaying basidiomycete polypore fungi on three mangrove species (Rhizophora mangle, Avicennia germinans, and Laguncularia racemosa) in a Panamanian mangrove forest. They note that the pattern typically observed for these fungi in diverse tropical forests is that there are a large number of rare species, with the smaller number of common species necessarily being nonspecialists due to the challenge of host rarity. In contrast, the authors found that in the tropical mangrove forest they studied, the polypore assemblage was strongly dominated by a few host-specialized species. Three fungal species, each with a strong preference for a different mangrove host species, comprised 88 percent of all fungi collected (the authors note, however, that these fungi are all reported from other hosts outside of mangrove forests as well). At least for polypore fungi within tropical mangrove forests, where host diversity is low and the abundance of individual host species is high, the restriction against host specialization typically imposed by host rarity in tropical forests may be relaxed, resulting in a polypore community dominated by a few common host-specialist species. (Gilbert and Sousa 2002) One polypore found to be locally specialized on White Mangrove in the study by Gilbert and Sousa (2002), Datronia caperata, was found only on this host in several other Panamanian mangrove forests as well (Parrent et al. 2004).

Smith et al. (2009) studied the effects of fiddler crabs (Uca rapax and Uca pugilator) on the growth of White Mangroves in a restored Florida marsh. They found that the presence of crab burrowing increased final tree height by 27%, final basal trunk diameter by 25%, and final leaf production by 15% over mangroves growing where crabs were removed and excluded. They also observed significant positive associations between mangrove production and crab burrow density. Crab burrows accounted for 24, 29, and 16% of the variation in mangrove height, trunk diameter, and leaf production, respectively.

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Population Biology

Laguncularia racemosa is surpassed in abundance by both the red mangrove, R. mangle, and the white mangrove, A. germinans in most areas of the lagoon. However, large stands of white mangroves can be found in patches, often in disturbed higher marsh areas.
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Life History and Behavior

Reproduction

White mangroves are semiviviparous, with germination of seedlings starting while propagules are still attached to the parent plant. The species is androdioecious, with both hermaphroditic and male plants in a population (Tomlinson 1980, Landry & Rathcke 2007). Flowering occurs from May to December in Florida, peaking in June and July (Tomlinson 1980). Male flowers are typically open for one day, and hermaphroditic flowers remain viable for two days (Landry & Rathcke 2007). Both flower types produce nectar and are pollinated by a wide variety of insects. However, hermaphroditic flowers have the ability to self-pollinate (Rathcke et al. 2001, Landry 2005). Fruits, or propagules, mature within a few months. Germination continues to completion after the propagule drops from the parent tree and is dispersed in the water.Dispersal: Propagules of the white mangrove are approximately 2 cm in length, flattened and lens-shaped. Original coloration is pea-green, turning brown within days after ripening and falling from the tree. Dispersal is facilitated by the outer tissue layer, or pericarp, which acts as a float for the propagule. To fully germinate, propagules must remain in the water for a period of about eight days, and have a lifespan of approximately 35 days. Roots often begin to develop on floating propagules after five days (Rabinowitz 1978).
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The slightly flesht fruit (a drupe) of White Mangrove floats and is dispersed by water. The fruit contains a single large seed which starts to enlarge (and sometimes to germinate) within the fruit while still on the tree or floating in the water. White Mangrove grows rapidly and may flower and fruit when less than 2 years old. (Little and Wadsworth 1964)

Some White Mangrove populations are androdioecious (i.e., with separate male and hermaphrodite individuals), while others lack male plants. Landry et al. (2009) surveyed 65 populations in Florida and the Bahamas. Because White Mangrove fruits are water-dispersed, the observed distribution of breeding systems was compared to local and regional water currents in order to determine whether dispersal could be important to the maintenance of male plants in androdioecious populations. Twenty-two of the 36 populations surveyed in Florida were androdioecious, with male frequencies that ranged from 1 to 68%. On the east coast of Florida, all populations north of latitude 26 degrees 30' N lacked males, while all populations south of this latitude were androdioecious, suggesting that northern populations may lack males due to dispersal limitation. The pattern of distribution on the west coast of Florida suggests that males may be maintained in some populations via dispersal. Nine islands in the north-central Bahamas were surveyed. and androdioecious populations were found only on San Salvador island, where male frequencies ranged from 5 to 28%. Landry et al. discuss the possible roles of dispersal, fragmentation, and selection in explaining the observed pattern of distribution of androdioecious and pure hermaphrodite populations.

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Physiology and Cell Biology

Physiology

Gilbert et al. (2002) studied the possible role of salt excretion by mangroves as a defense against pathogenic fungi in a mangrove forest in Panama. Although presumably evolved for other reasons, salt excretion by leaves of some mangrove species may serve as an important defense against fungal attack, reducing the vulnerability of typically high-density, monospecific forest stands to severe disease pressure. In their study, Gilbert et al. found that Black Mangrove (Avicennia germinans) suffered much less fungal leaf damage from than did White Mangrove (Laguncularia racemosa) or Red Mangrove (Rhizophora mangle). Black Mangrove leaves also supported the least fungal growth on the leaf surface, the least endophytic colonization, and the lowest fungal diversity, followed by White Mangrove and Red Mangrove.

Host specificity of leaf-colonizing fungi was greater than expected at random. The fungal assemblage found on Black Mangrove appears to be a subset of the fungi that can grow on the leaves of Red and White Mangrove. The authors suggested that the different salt tolerance mechanisms in the three mangrove species may differentially regulate fungal colonization. The mangroves differ in their salt tolerance mechanisms such that Black Mangrove (which excretes salt through leaf glands) has the highest salinity of residual rain water on leaves, White Mangrove (which accumulates salt in the leaves) has the greatest bulk salt concentration, and Red Mangrove (which excludes salt at the roots) has little salt associated with leaves. The high salt concentrations associated with leaves of Black and White Mangrove, but not the low salinity of Red Mangrove, were sufficient to inhibit the germination of many fungi associated with mangrove forests. The authors suggest that efficient defenses against pathogens may be especially important in natural communities, such as mangrove forests, where host diversity is low and the density of individual hosts is high – ideal conditions for diseases to have strong impacts on plant populations.

Mangrove forests are unusual among tropical forests for their low tree species diversity and associated high population density of individual species. Mangrove species are unusual in their ability to grow in flooded, saline soils and for the array of mechanisms they have evolved to tolerate high salt concentrations. The work by Gilbert et al. suggests that some mangrove species may also be unusual in their escape from strong disease pressures, even when growing at high densities, through the inhibitory effects of
high foliar (leaf) salt concentration on fungal infection. (Gilbert et al. 2002)

The leaves of mangroves in general have abundant and diverse glands and their function and physiological and ecological importance appear to be poorly known. The glands on the petioles (leaf stalks) of White Mangrove and Buttonwood, for example are extrafloral nectaries (i.e., structures outside of flowers that secrete nectar), but the ecological significance of these structures seems not to have been investigated (these glands are not salt-secreting structures, as is sometimes claimed). The glands on the leaf blade of White Mangrove accumulate salt, but it is not secreted to the leaf surface, so in contrast to a Black Mangrove leaf, a White Mangrove leaf does not taste salty when licked. It is sometimes stated that the glands on the leaf blade of White Mangrove serve as domatia (special shelters) for beneficial mites. While this may be true, this appears to be another intriguing topic that remains unstudied.

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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
LC
Least Concern

Red List Criteria

Version
3.1

Year Assessed
2010

Assessor/s
Ellison, A., Farnsworth, E. & Moore, G.

Reviewer/s
Polidoro, B.A., Livingstone, S.R. & Carpenter, K.E. (Global Marine Species Assessment Coordinating Team)

Contributor/s

Justification
This species is widespread and tolerant of a variety of habitats. Relative to the other mangrove species within the wider Caribbean, the conservation status of this species appears to be more stable. However, this species is threatened by the loss of mangrove habitat throughout its range, primarily due to extraction and coastal development. There has been an estimated 17% decline in mangrove area within this species range since 1980. Mangrove species are more at risk from coastal development and extraction at the extremes of their distribution, and are likely to be contracting in these areas more than in other areas. It is also likely that changes in climate due to global warming will further affect these parts of the range. Therefore there are overall range declines in many areas, but not enough to reach any of the threatened category thresholds. This species is listed as Least Concern.
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National NatureServe Conservation Status

United States

Rounded National Status Rank: N2 - Imperiled

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NatureServe Conservation Status

Rounded Global Status Rank: G5 - Secure

Reasons: Shores of central and southern Florida including Florida Keys, Bermuda, and nearly throughout West Indies from Bahamas and Cuba to Trinidad and Tobago and Dutch West Indies. On both coasts of continental tropical America from Mexico south to Ecuador and northwestern Peru and to Brazil. The most widely distributed of the mangrove species in Puerto Rico.

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Population

Population
Although there is no species specific population information, it can be assumed that there are areas of population decline throughout its range due to coastal development.

Individual population sizes are highly variable through time, as mortality of seedlings can be quite high due to competition with other mangroves (Sherman et al. 2001, Ross et al. 2006). Seedlings recruit in thousands following a disturbance (Sherman et al. 2001). Saplings and trees can number in thousands in a few areas, but Laguncularia tends to be less populous in a given mangal than the other Afrotropical species, with low importance values and basal area (Murray et al. 2003).

Eastern and western Atlantic provenances of Laguncularia show significant genetic differentiation, as indicated by leaf chemistry (Dodd et al. 1998). There have been no other genetic studies to date.

Population Trend
Decreasing
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Threats

Major Threats
Laguncularia is likely to be susceptible to increasing tidal height or salt intrusion, especially under conditions of sea-level rise (Ross et al. 2000). The invasive alien species, Schinus terebrinthifolius, is directly impacting populations in Florida (Schmalzer 1995, Herwitz et al. 1996, Gordon 1998, Ewe and Sternberg 2005). The species tends to be outcompeted by Rhizophora mangle; the native fern, Acrostichum aureum, may also affect seedling establishment. In addition, altough marine aquaculture is not documented specifically from the Caribbean, on the Pacific coast it is negatively impacting Laguncularia (Kovacs 1999). Although local estimates are uncertain due to differing legislative definitions of what is a 'mangrove' and to the imprecision in determining mangrove area, current consensus estimates of mangrove loss in the last quarter-century report an approximately 17% decline in mangrove areas in countries within this species range since 1980 (FAO 2007).

Laguncularia has a higher requirement for freshwater inputs than other mangrove species, so may be vulnerable to drought. Storms/floods/hurricanes have extensively damaged Laguncularia stands (Roth 1992, McCoy et al. 1996, Sherman et al. 2001, Piou et al. 2006, Ross et al. 2006). Laguncularia experiences high mortality and leaf loss during freeze events at the northern edge of its range in Florida (Ellis et al. 2006).

Selective logging is also a possible threat as this species is regarded as suitable for polewood construction on Pacific coast of Mexico (Kovacs 1999). Clear-cutting is occurring in certain areas (Suman 1994); loss rates are estimated at 1.4%/year for Laguncularia/Avicennia-dominated mangal in western Mexico (Ramirez-Garcia et al. 1998). Clearing of mangal for settlement and agriculture cited as major cause of decline in Latin America (Lacerda 1993) without a compensating economic return from agriculture or fast mangrove recovery (Tovilla-Hernandez et al. 2001). Subsistence use of Laguncularia for fuelwood occurs (Kovacs 1999).

All mangrove ecosystems occur within mean sea level and high tidal elevations, and have distinct species zonations that are controlled by the elevation of the substrate relative to mean sea level. This is because of associated variation in frequency of elevation, salinity and wave action (Duke et al. 1998). With rise in sea-level, the habitat requirements of each species will be disrupted, and species zones will suffer mortality at their present locations and re-establish at higher elevations in areas that were previously landward zones (Ellison 2005). If sea-level rise is a continued trend over this century, then there will be continued mortality and re-establishment of species zones. However, species that are easily dispersed and fast growing/fast producing will cope better than those which are slower growing and slower to reproduce.

In addition, mangrove area is declining globally due to a number of localized threats. The main threat is habitat destruction and removal of mangrove areas. Reasons for removal include cleared for shrimp farms, agriculture, fish ponds, rice production and salt pans, and for the development of urban and industrial areas, road construction, coconut plantations, ports, airports, and tourist resorts. Other threats include pollution from sewage effluents, solid wastes, siltation, oil, and agricultural and urban runoff. Climate change is also thought to be a threat, particularly at the edges of a species range. Natural threats include cyclones, hurricane and tsunamis.
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Management

Conservation Actions

Conservation Actions
There are no conservation measures specific to this species, but its range may include some marine and coastal protected areas. More research is needed on the effects of ongoing sea level rise on Laguncularia, given its low salinity tolerance. Likewise, the impacts of invasive species (and consequences of their removal) on Laguncularia need to be quantified.

The effectiveness of habitat restoration and success of replantings with Laguncularia needs to be assessed. New Landsat and IKONOS technology should be used to do species-based, landscape-level monitoring of deforestation (Kovacs et al. 2005). More research needed is on Laguncularia influences on water quality, erosion control, and pollution buffering.

Restoration of Laguncularia is being pursued in Florida (Milano 1999, McKee and Faulkner 2000), Costa Rica (Lewis and Marshall 1998) and Colombia (Elster and Perdomo 1999, Elster 2000). See valuable general review by Lewis (2005). More information and forestry trials are needed to optimize silvicultural techniques and management.
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Relevance to Humans and Ecosystems

Benefits

Economic Uses

Uses: MEDICINE/DRUG, Building materials/timber, Fuelwood, Tannin/dye

Comments: The wood is used locally for fuel. The bark is rich in tannin and is occasionally employed in tanning and for medicinal purposes (Little and Wadsworth 1964). In Brazil, aside from fuelwood and charcoal, the wood is used to some extent in carpentry and for small beams and rafters (Santos 1987).

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Mangrove forest ecosystems are vital as sources of energy, provide nursery habitat for juvenile fishes and invertebrates, and are important as buffers in decreasing storm impacts along coastlines. Additionally, they provide roosting and nesting habitat for wading birds and serve as a source for timber production.
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Uses

Bees make good honey from the flowers of White Mangrove (Elias 1980).

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Wikipedia

Laguncularia racemosa

Laguncularia racemosa (white mangrove; syn. Conocarpus racemosa) is a species of flowering plant in the leadwood tree family, Combretaceae. It is native to the coasts of western Africa from Senegal to Cameroon, the Atlantic coast of the Americas from Bermuda, Florida, the Bahamas, Mexico, the Caribbean and south to Brazil; and on the Pacific coast of the Americas from Mexico to northwestern Peru, including the Galápagos Islands.

Laguncularia racemosa flowers

It is a mangrove tree, growing to 12–18 metres (39–59 ft) tall. The bark is gray-brown or reddish, and rough and fissured. Pneumatophores and/or prop roots may be present, depending on environmental conditions. The leaves are opposite, elliptical, 12–18 centimetres (4.7–7.1 in) long and 2.5–5 cm (0.98–1.97 in) broad, rounded at both ends, entire, smooth, leathery in texture, slightly fleshy, without visible veins, and yellow-green in color. The petiole is stout, reddish, 10–13 mm (0.39–0.51 in) long, with two small glands near the blade that exude sugars. The white, bell-shaped flowers are mostly bisexual and about 5 mm (0.20 in) long. The fruit is a reddish-brown drupe, about 12–20 mm (0.47–0.79 in) long, with longitudinal ridges. The single seed is sometimes viviparous.

It grows in coastal areas of bays, lagoons, and tidal creeks, typically growing inland of other mangroves, well above the high tide line.

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