Invasive Species Compendium

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Rhizophora mangle
(red mangrove)

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Datasheet

Rhizophora mangle (red mangrove)

Summary

  • Last modified
  • 27 September 2018
  • Datasheet Type(s)
  • Invasive Species
  • Host Plant
  • Preferred Scientific Name
  • Rhizophora mangle
  • Preferred Common Name
  • red mangrove
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • Rhizophoramangle, a mangrove tree 5-10 m tall, is an important native species in coastal areas in tropical and subtropical America, with many environmental benefits, protecting coasts against storm and...

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Pictures

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PictureTitleCaptionCopyright
Rhizophora mangle (red mangrove); habit. Kealia Pond, Maui, Hawaii, USA. July, 2013.
TitleHabit
CaptionRhizophora mangle (red mangrove); habit. Kealia Pond, Maui, Hawaii, USA. July, 2013.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); habit. Kealia Pond, Maui, Hawaii, USA. July, 2013.
HabitRhizophora mangle (red mangrove); habit. Kealia Pond, Maui, Hawaii, USA. July, 2013.©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); habit. Waiohuli Keokea, Maui, Hawaii, USA. September, 2005.
TitleHabit
CaptionRhizophora mangle (red mangrove); habit. Waiohuli Keokea, Maui, Hawaii, USA. September, 2005.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); habit. Waiohuli Keokea, Maui, Hawaii, USA. September, 2005.
HabitRhizophora mangle (red mangrove); habit. Waiohuli Keokea, Maui, Hawaii, USA. September, 2005.©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); dense habit. Palaauwai, Molokai, Hawaii, USA. April, 2012.
TitleDense habit
CaptionRhizophora mangle (red mangrove); dense habit. Palaauwai, Molokai, Hawaii, USA. April, 2012.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); dense habit. Palaauwai, Molokai, Hawaii, USA. April, 2012.
Dense habitRhizophora mangle (red mangrove); dense habit. Palaauwai, Molokai, Hawaii, USA. April, 2012.©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); habit. Waiohuli Keokea, Maui, Hawaii, USA. September, 2005.
TitleHabit
CaptionRhizophora mangle (red mangrove); habit. Waiohuli Keokea, Maui, Hawaii, USA. September, 2005.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); habit. Waiohuli Keokea, Maui, Hawaii, USA. September, 2005.
HabitRhizophora mangle (red mangrove); habit. Waiohuli Keokea, Maui, Hawaii, USA. September, 2005.©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); large trees with aerial prop roots. Laie Kihei, Maui, Hawaii, USA. January, 2010.
TitleAerial roots
CaptionRhizophora mangle (red mangrove); large trees with aerial prop roots. Laie Kihei, Maui, Hawaii, USA. January, 2010.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); large trees with aerial prop roots. Laie Kihei, Maui, Hawaii, USA. January, 2010.
Aerial rootsRhizophora mangle (red mangrove); large trees with aerial prop roots. Laie Kihei, Maui, Hawaii, USA. January, 2010.©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); habit. Mangrove Park, Boynton Beach, Florida, USA. September, 2009.
TitleHabit
CaptionRhizophora mangle (red mangrove); habit. Mangrove Park, Boynton Beach, Florida, USA. September, 2009.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); habit. Mangrove Park, Boynton Beach, Florida, USA. September, 2009.
HabitRhizophora mangle (red mangrove); habit. Mangrove Park, Boynton Beach, Florida, USA. September, 2009.©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); leaves. Kealia Pond, Maui, Hawaii, USA. July 03, 2013
TitleLeaves
CaptionRhizophora mangle (red mangrove); leaves. Kealia Pond, Maui, Hawaii, USA. July 03, 2013
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); leaves. Kealia Pond, Maui, Hawaii, USA. July 03, 2013
LeavesRhizophora mangle (red mangrove); leaves. Kealia Pond, Maui, Hawaii, USA. July 03, 2013©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); leaves and propagules forming. Mangrove Nature Park, Boynton Beach, Florida, USA. September, 2009.
TitleLeaves
CaptionRhizophora mangle (red mangrove); leaves and propagules forming. Mangrove Nature Park, Boynton Beach, Florida, USA. September, 2009.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); leaves and propagules forming. Mangrove Nature Park, Boynton Beach, Florida, USA. September, 2009.
LeavesRhizophora mangle (red mangrove); leaves and propagules forming. Mangrove Nature Park, Boynton Beach, Florida, USA. September, 2009.©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); propagules. Waiohuli Keokea, Maui, Hawaii, USA. September, 2005.
TitleFruit
CaptionRhizophora mangle (red mangrove); propagules. Waiohuli Keokea, Maui, Hawaii, USA. September, 2005.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); propagules. Waiohuli Keokea, Maui, Hawaii, USA. September, 2005.
FruitRhizophora mangle (red mangrove); propagules. Waiohuli Keokea, Maui, Hawaii, USA. September, 2005.©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); propagules. Kealia Pond, Maui, Hawaii, USA. July, 2013.
TitleFruits
CaptionRhizophora mangle (red mangrove); propagules. Kealia Pond, Maui, Hawaii, USA. July, 2013.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); propagules. Kealia Pond, Maui, Hawaii, USA. July, 2013.
FruitsRhizophora mangle (red mangrove); propagules. Kealia Pond, Maui, Hawaii, USA. July, 2013.©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); prop roots. Laie Kihei, Maui, Hawaii, USA. January, 2010.
TitleProp roots
CaptionRhizophora mangle (red mangrove); prop roots. Laie Kihei, Maui, Hawaii, USA. January, 2010.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); prop roots. Laie Kihei, Maui, Hawaii, USA. January, 2010.
Prop rootsRhizophora mangle (red mangrove); prop roots. Laie Kihei, Maui, Hawaii, USA. January, 2010.©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); small plant. Laie Kihei, Maui, Hawaii, USA. January, 2010.
TitleSmall plant
CaptionRhizophora mangle (red mangrove); small plant. Laie Kihei, Maui, Hawaii, USA. January, 2010.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); small plant. Laie Kihei, Maui, Hawaii, USA. January, 2010.
Small plantRhizophora mangle (red mangrove); small plant. Laie Kihei, Maui, Hawaii, USA. January, 2010.©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); habitat, with young plants. Pukoo, Molokai, Hawaii, USA. May, 2005.
TitleHabitat, with young plants
CaptionRhizophora mangle (red mangrove); habitat, with young plants. Pukoo, Molokai, Hawaii, USA. May, 2005.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); habitat, with young plants. Pukoo, Molokai, Hawaii, USA. May, 2005.
Habitat, with young plantsRhizophora mangle (red mangrove); habitat, with young plants. Pukoo, Molokai, Hawaii, USA. May, 2005.©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); ex-habitat, with trees removed and chipped. Laie Kihei, Maui, Hawaii, USA. January, 2010.
TitleControl measures
CaptionRhizophora mangle (red mangrove); ex-habitat, with trees removed and chipped. Laie Kihei, Maui, Hawaii, USA. January, 2010.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); ex-habitat, with trees removed and chipped. Laie Kihei, Maui, Hawaii, USA. January, 2010.
Control measuresRhizophora mangle (red mangrove); ex-habitat, with trees removed and chipped. Laie Kihei, Maui, Hawaii, USA. January, 2010.©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); rebounding after control. Kealia Pond, Maui, Hawaii, USA. July, 2013.
TitleRegrowth after control
CaptionRhizophora mangle (red mangrove); rebounding after control. Kealia Pond, Maui, Hawaii, USA. July, 2013.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Rhizophora mangle (red mangrove); rebounding after control. Kealia Pond, Maui, Hawaii, USA. July, 2013.
Regrowth after controlRhizophora mangle (red mangrove); rebounding after control. Kealia Pond, Maui, Hawaii, USA. July, 2013.©Forest Starr & Kim Starr - CC BY 4.0

Identity

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Preferred Scientific Name

  • Rhizophora mangle L.

Preferred Common Name

  • red mangrove

Other Scientific Names

  • Bruguiera decangulata Griff.
  • Rhizophora americana Nutt.
  • Rhizophora racemosa G.F.W.MEY.

International Common Names

  • English: American mangrove; mangrove
  • Spanish: mangle; Mangle colorado; Mangle pinon; mangle rojo
  • French: manglier; manglier chandelle; Manglier rouge; Paletuvier rouge
  • Portuguese: manga-robeira; mangue-verdadeiro

Local Common Names

  • Brazil: manga robeira; mangue vermelho
  • Dominican Republic: mangle; mangle colorado; mangle rojo
  • Germany: Amerikanische Mangrove; Rote Mangrove
  • Haiti: mang chandel; mang nwa; mang wouj
  • Italy: mangrovia d'America
  • Netherlands: Wortelboom
  • Puerto Rico: mangle; mangle colorado; mangle de chifle; mangle rojo; mangle zapatero
  • Sweden: amerikansk mangrove

EPPO code

  • RHZMA (Rhizophora mangle)

Summary of Invasiveness

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Rhizophoramangle, a mangrove tree 5-10 m tall, is an important native species in coastal areas in tropical and subtropical America, with many environmental benefits, protecting coasts against storm and tidal damage and providing habitats needed for numerous marine and estuarine species. It can colonize areas rapidly and form dense, naturally monospecific stands that outcompete other species, and as such is an essential component of native ecosystems, though it is occasionally noted as weedy where native. It has only rarely been introduced, and is reported as an alien invasive species only in Hawaii where it was intentionally planted more than a century ago; there it has had a major negative environmental impact on native biodiversity, especially in coastal fish ponds, and has proven difficult to control. Further introduction is not recommended.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Rhizophorales
  •                         Family: Rhizophoraceae
  •                             Genus: Rhizophora
  •                                 Species: Rhizophora mangle

Notes on Taxonomy and Nomenclature

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R. mangle is one of approximately 35 species of true mangroves, with another 60 or more species of mangrove associates (Hill, 2001). Most of these species occur in the Indo-Pacific region, with R. mangle being one of the three species that commonly occur in the Americas. R. mangle is one a of small number of species in this pantropical genus, with six species of Rhizophora noted in the Plant List (2013) and seven by USDA-ARS (2015). Two varieties are noted by the Missouri Botanic Garden (2015), var. racemosa and var. samoensis, whereas both are given species rank by The Plant List (2013) and USDA-ARS (2015). They are sometimes called ‘sibling species’ to R. mangle, noting their close morphological resemblance and overlapping native ranges in part, and further taxonomic clarification based on molecular analysis is required (Duke and Allen, 2006). The hybrid R. x harrisonii (The Plant List 2013), sometimes given specific rank (e.g. USDA-ARS, 2015), is considered by Duke and Allen (2006) and others as a hybrid between R. mangle and R. racemosa. Duke and Allen (2006) include a map and much detail regarding the taxonomy of this species group. But noting the taxonomic issue, it is possible that some reports and records refer to misidentifications, especially in the Pacific and West Africa.

Description

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R. mangle is normally a small evergreen tree 5-10 m tall with a trunk diameter of 20 cm, but can grow to 20-30 m (and even 50 m) tall, with diameters of 20–50(-70) cm, with arching stilt roots 2–4.5 m high. Bark grey or grey-brown, smooth and thin on small trunks, becoming furrowed and thick on larger ones. Inner bark reddish or pinkish, with a slightly bitter and salty taste. Twigs stout, grey or brown, hairless, ending in a conspicuous narrow pointed green bud 2.5–5 cm long, covered with 2 green scales (stipules) around pairs of developing leaves, and making a ring scar around the twig when shedding. Leaves opposite, crowded at end of twig, hairless, with slightly flattened leafstalks 13–22 mm. Blades elliptical, 6-10 cm long, blunt at apex and short-pointed at base, slightly rolled under at edges, slightly leathery and fleshy with side veins not visible, shiny green above, yellow green beneath. Flowers usually 2-4 together at leaf bases on forked green stalks, 4-7.5 cm long, slightly fragrant, pale yellow, about 2 cm across. The bell-shaped pale yellow base (hypanthium) less than 6 mm long bears four widely spreading narrow pale yellow sepals almost 13 mm long, leathery and persistent; four narrow petals 10 mm long, curved downward, whitish but turning brown, white woolly or cottony on inner side; eight stamens; pistil of two-celled ovary mostly inferior but conical at apex, with two ovules in each cell, slender style, and two-lobed stigma. Fruits dark brown, conical, about 3 cm long and 13 mm in diameter, with enlarged curved sepals, remaining attached. The single seed germinates inside the fruit, forming the long narrow first root (radicle), green except for brown enlarged and pointed end, up to 13 mm in diameter. The propagules fall when they are 20-30 cm long (adapted from Duke, 1983; Little and Skolmen, 1989; Hill, 2001; Duke and Allen, 2006).

Distribution

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R. mangle is a tropical and subtropical American species, native to a wide area on both eastern and western coasts: on the Pacific, from Baja California, Mexico, to northern Peru, and on the east, from North Carolina to Florida (USDA-NRCS, 2015) and around the Gulf of Mexico, the Caribbean, and along northern South America to the north east of Brazil (USDA-ARS, 2015). However, the common name American mangrove is somewhat of a misnomer, as the species is also reported as native to West Africa, from Senegal to Nigeria (Duke, 1983; USDA-ARS, 2015). A further distinct record from Angola (Duke, 1983) requires verification, and may refer to R. racemosa. Steele et al. (1999) reported R. mangle as native in American Samoa, but this is considered a misidentification, and refers to R. samoensis. There is some confusion among authorities about whether R. mangle in the Pacific (Micronesia, Polynesia) is native or introduced (ISSG, 2013; USDA-ARS, 2015).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasivePlantedReferenceNotes

Asia

MalaysiaUnconfirmed recordCAB Abstracts

Africa

AngolaPresentNativeDuke, 1983Record may refer to R. racemosa
BeninPresentNativeDuke, 1983
Côte d'IvoirePresentNative Not invasive
GambiaPresentNativeDuke, 1983
GhanaPresentNativeDuke, 1983
GuineaPresentNativeUSDA-ARS, 2015
Guinea-BissauPresentNativeDuke, 1983
LiberiaPresentNativeDuke, 1983
NigeriaPresentNativeUSDA-ARS, 2015
SenegalPresentNativeUSDA-ARS, 2015
Sierra LeonePresentNative Not invasive
TogoPresentNativeDuke, 1983

North America

BermudaPresentNativeMissouri Botanical Garden, 2015
MexicoPresentNative Not invasive USDA-ARS, 2015
USAPresent
-FloridaPresentNative Natural USDA-NRCS, 2015
-HawaiiPresentIntroduced Invasive Duke and Allen, 2006; USDA-ARS, 2015
-North CarolinaPresentNativeUSDA-NRCS, 2015
-South CarolinaPresentNativeUSDA-NRCS, 2015

Central America and Caribbean

BahamasPresentNativeMissouri Botanical Garden, 2015
BelizePresentNativeUSDA-ARS, 2015
Costa RicaPresentNativeUSDA-ARS, 2015
Dominican RepublicPresentNativeUSDA-ARS, 2015
El SalvadorPresentNativeMissouri Botanical Garden, 2015
GuatemalaPresentNativeMissouri Botanical Garden, 2015
HaitiPresentNativeTimyan, 1996, publ. 1997
HondurasPresentNativeUSDA-ARS, 2015
JamaicaPresentNativeUSDA-ARS, 2015
NicaraguaPresentNativeUSDA-ARS, 2015
PanamaPresentNativeUSDA-ARS, 2015
Puerto RicoPresentNativeUSDA-NRCS, 2015
Trinidad and TobagoPresentNativeMissouri Botanical Garden, 2015

South America

BrazilPresentNativeUSDA-ARS, 2015
-BahiaPresentNativeMissouri Botanical Garden, 2015
-CearaPresentNativeMissouri Botanical Garden, 2015
-MaranhaoPresentNativeMissouri Botanical Garden, 2015
-ParaPresentNativeMissouri Botanical Garden, 2015
-Rio de JaneiroPresentNativeMissouri Botanical Garden, 2015
-Santa CatarinaPresent
-Sao PauloPresentNativeMissouri Botanical Garden, 2015
ColombiaPresentNativeUSDA-ARS, 2015
EcuadorPresentNative Natural USDA-ARS, 2015
-Galapagos IslandsPresentNativeMissouri Botanical Garden, 2015
French GuianaPresentNativeUSDA-ARS, 2015
GuyanaPresentNativeUSDA-ARS, 2015
PeruPresentNativeUSDA-ARS, 2015
SurinamePresentNativeUSDA-ARS, 2015
VenezuelaPresentNativeUSDA-ARS, 2015

Oceania

American SamoaAbsent, invalid recordNativeSteele et al., 1999Considered a misidentification; record refers to R. samoensis
Australia
-QueenslandEradicatedIntroduced Invasive Ross RIver, Townsville
French PolynesiaPresentIntroducedUSDA-ARS, 2015
Micronesia, Federated states ofPresentNative Not invasive

History of Introduction and Spread

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In its native range, Holm et al. (1991) note R. mangle as a weed in Nicaragua and the USA, but with no information on impacts.

As an exotic, several reports indicate that R. mangle was introduced to Hawaii in 1902, and there were no mangrove species present prior to this date. Duke and Allen (2006) note that introductions to Hawaii and the Society Islands (French Polynesia) were made from populations in Florida. It was intentionally planted on the island of Molokai, apparently by the American Sugar Company, primarily for the purpose of stabilizing coastal mudflats, and was planted in salt marshes on Oahu in 1922 (Little and Skolmen, 1989). The introductions of 1922 as well as 1960 involved the planting of an estimated 3000 propagules (Allen, 1998). At least five other mangroves or associated species were introduced at the same time, although none of these spread to the extent that R. mangle has, though Bruguiera gymnorrhiza (now identified as B. sexangula (Allen et al. (2000)) and Conocarpus erectus have naturalized (Allen, 1998). In the absence of native mangrove species, R. mangle invaded coastal habitats on all the main islands except Kahoolawe and Niihau, and it continues to spread (Csurhes and Shanahan, 2012). It is now very well established throughout the Hawaiian archipelago, and in some places densities of more than 24,000 trees per hectare have been reported (Cox and Allen, 1999).

R. mangle has also been reported as present in Queensland, Australia. However, Csurhes and Shanahan (2012) detail the true situation there, where it should be recorded as eradicated. It has only ever been recorded from the Townsville Palmetum in coastal northern Queensland, the first and only record in Queensland or Australia, where two trees were reported planted, thought to have been ‘gifts’. These were destroyed in 2002, and surveillance in surrounding areas did not find any other specimens, and as such, it is considered that it may have been eradicated.

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
French Polynesia Florida   Duke and Allen (2006)
Hawaii Florida 1902 Habitat restoration and improvement (pathway cause) Yes Duke and Allen (2006)

Risk of Introduction

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R. mangle is opportunistically invasive, with a high potential to invade alien environments and is not recommended for planting outside its natural range (Duke and Allen, 2006). An invasive species risk assessment for Australia indicated that R. mangle has the potential to invade mangrove communities in Queensland, competing or hybridizing with native Rhizophora species; it is thus classified as a ‘high risk’ species for preventative control (Csurhes and Shanahan, 2012).

Habitat

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R. mangle is the dominant neotropical mangrove species, and is commonly found from low intertidal swamp margins to shaded sites at the highest high water mark. It is always a coastal species, mainly inhabiting tidal flats and estuarine areas, and is noted by Hill (2001) to be found in five different topographical communities in subtropical to tropical coastal regions. These are fringe, riverine, overwash, basin and supra-tidal flats, the differences between these being mainly in elevation and the effects this has on changes in the water-level and proximity to the ocean. R. mangle can survive under permanent submersion, resulting from flooding, due to its unique anatomy and physiology (Hill, 2001).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial – ManagedDisturbed areas Present, no further details Natural
Terrestrial ‑ Natural / Semi-naturalRiverbanks Present, no further details Natural
Wetlands Present, no further details Natural
Littoral
Coastal areas Principal habitat Harmful (pest or invasive)
Coastal areas Principal habitat Natural
Mangroves Principal habitat Natural
Mud flats Principal habitat Natural
Intertidal zone Principal habitat Natural
Salt marshes Present, no further details Natural

Biology and Ecology

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Genetics

The chromosome number of R. mangle is 2n = 36 (Yoshioka et al., 1984). Natural stands of R. mangle tend to form single species monocultures with little genetic diversity (Lowenfeld and Klekowski, 1992), and being self-pollinating, this leads to inbreeding depression within the population (Proffitt et al., 2006).

Reproductive Biology

R. mangle flowers are hermaphroditic, containing both male and female sex organs, and pollen is wind dispersed (Mehlig, 2006), both of which may increase the high incidence of self-pollination, leading to inbreeding depression within populations (Proffitt et al., 2006). Vegetative reproduction through coppicing and suckers is possible but not common for red mangrove (Proffitt et al., 2006).

R. mangle usually begins flowering before 6 years old, sometimes from when as young as 3 years old, and flowering has been reported in saplings as small as 0.5-1 m in height (Allen, 2002). Some planted R. mangle trees were seen not to set fruit until the second flowering episode, about 2.5 years after planting (Ellison and Farnsworth, 1997). Species of the genus Rhizophora have been described as ever-flowering, as flowers and fruits at all stages of development can be found throughout the year in any stand or even on a single tree, although flowering on a single shoot may be periodic (Gill and Tomlinson, 1969). Flower production in R. mangle is continuous throughout the year in equatorial regions, with a peak in flower production during the wet season (Mehlig, 2006), and during early spring and summer in more temperate regions (Gill and Tomlinson, 1971). Flower production is not dependent on day-length, and embryo development can continue albeit reduced in colder climates and periods (Mehlig, 2006).

R. mangle is viviparous. Following fertilization, propagules exhibit continuous development from flower to germinated seedling while still attached to the parent plant, with no dormant or seed phase, and whereas the propagules resemble seed pods, they are actually embryonic root structures. High rates of propagule production are observed, and population levels can rise quickly (Krauss and Allen, 2003; Chimner et al., 2006). The abundance of fruit set is noted to be markedly seasonal in southern Florida, with maximum fruit production occurring in the autumn (Gill and Tomlinson, 1969). The propagules eventually fall from the parent plant, but if submerged at the time, they can float for more than a year in seawater until they settle on suitable strata (Hill, 2001).

Physiology and Phenology

As a facultative halophyte, R. mangle can tolerate waterlogged soils and high salinity in both water and sediment up to 90 ppt, but salinity is not a requirement for growth (Hill, 2001), and higher salinity can reduce establishment (Elster et al., 1999). Mangrove species exhibit different types of mechanisms for tolerating such high salt concentrations. R. mangle roots exclude the uptake of salt, whereas the black mangrove (Avicennia germinans) and the white mangrove (Laguncularia racemosa) take up seawater through their roots but excrete excess salt through pores or salt glands on leaf surfaces. In fully submerged conditions there is an absence of oxygen, and one of the most visible adaptations are root adaptations, with R. mangle developing prop roots from the stem or trunk of the plant from 2 to 4.5 m above the substrate (Duke, 1983), whereas other species rely on roots growing upwards from under the substrate. In either case, prop roots help to stabilise the tree, and allow oxygen to be supplied to the underground root system via lenticels or pores in the aerial roots (Hill, 2001).

Trees prefer full sun but seedlings can survive in the understorey until a gap forms (Hill, 2001). However, Farnsworth and Ellison (1996) found that R. mangle shows both light demanding and shade tolerant properties, observing changes in the mode and magnitude of its adaptability to light throughout its growth stages. Seedlings in the shade or full sun were seen to grow at different rates, and to differ in a range of morphological characteristics including leaf number, size and shape, specific leaf area, internode length, blade petiole angles, stomatal density and in the ratio of height to crown diameter.

R. mangle trees are not tolerant of fire, and have poor coppicing ability (Duke and Allen, 2006).

Associations

R. mangle commonly forms dense monospecific stands in its native range, or is associated with the black mangrove (Avicennia germinans) and the white mangrove (Laguncularia racemosa) amongst other species. Batis maritima is another common associate.

Environmental Requirements

R. mangle is a fast growing species that can tolerate the wide range of environmental conditions found in coastal areas, although Krauss and Allen (2003) report that low salinity combined with reduced light, or simply low sunlight alone, appear to favour R. mangle growth. R. mangle tends to prefer full sun, but seedlings can survive in the understorey until a gap forms (Hill, 2001). Optimal salinity is given as 8-26 ppt or about 35 ppt for seawater (Duke and Allen, 2006).

R. mangle seedlings cannot tolerate dry conditions during establishment, so communities often develop around areas where water is not limited, at least during certain periods (Elster et al., 1999). Mean temperature ranges for optimal growth of R. mangle are 21.6°C to 25.6°C (Duke, 1983) and cold temperatures limit the native range to below the latitudes of 28°N and 28°S (Hill, 2001). Substrate type can vary widely, with pH ranges of 5.3-8.5 being common (Duke and Allen, 2006). Limiting nutrients tend to be nitrogen and phosphorous (Davis et al., 2003).

Climate

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ClimateStatusDescriptionRemark
Af - Tropical rainforest climate Preferred > 60mm precipitation per month
Am - Tropical monsoon climate Preferred Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))
As - Tropical savanna climate with dry summer Preferred < 60mm precipitation driest month (in summer) and < (100 - [total annual precipitation{mm}/25])
Aw - Tropical wet and dry savanna climate Preferred < 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])
BS - Steppe climate Preferred > 430mm and < 860mm annual precipitation
BW - Desert climate Tolerated < 430mm annual precipitation
Cs - Warm temperate climate with dry summer Tolerated Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
28 28

Air Temperature

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Parameter Lower limit Upper limit
Absolute minimum temperature (ºC) 0
Mean annual temperature (ºC) 20 30
Mean maximum temperature of hottest month (ºC) 32 38
Mean minimum temperature of coldest month (ºC) 0 5

Rainfall

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ParameterLower limitUpper limitDescription
Dry season duration06number of consecutive months with <40 mm rainfall
Mean annual rainfall5004000mm; lower/upper limits

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Aratus pisonii Herbivore Leaves Hill, 2001
Cardisoma guanhumi Herbivore Leaves Hill, 2001
Cercospora Pathogen not specific McMillan RT Jr, 1984
Cerithidea scalariformis Herbivore Fruits/pods/Seedlings/Seeds Hill, 2001
Coccotrypes rhizophorae Herbivore Roots Timyan, 1996, publ. 1997
Cylindrocarpon didymum Pathogen Roots/Stems Timyan, 1996, publ. 1997
Cytospora rhizophorae Pathogen Leaves Wier et al., 2000
Diaprepes abbreviatus Herbivore N
Goniopsis cruentata Herbivore Fruits/pods/Leaves/Seedlings/Seeds Hill, 2001
Melampus coffea Herbivore Fruits/pods/Seedlings/Seeds Hill, 2001
Pseudocercospora rhizophorae Pathogen Leaves Little and Skolmen, 1989
Pterosporidium rhizomorphae Pathogen Leaves Little and Skolmen, 1989
Sphaeroma terebrans Herbivore Roots Timyan, 1996, publ. 1997
Ucides cordatus Herbivore Fruits/pods/Seedlings/Seeds Hill, 2001

Notes on Natural Enemies

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A number of pests and diseases have been reported in the native range. Pests include the wood borers Poecilips rhizophorae [Coccotrypes rhizophorae] and Sphaeroma terebrans that attack prop roots and cause extensive damage in Florida (Timyan, 1996), and the citrus weevil (Diaprepes abbreviatus). Diseases include Pseudocercospora rhizophorae, Anthostomella rhizomorphae [Pterosporidiumrhizomorphae] on leaves (Little and Skolmen, 1989), Cytospora rhizophorae in Puerto Rico (Wier et al., 2000), Cercospora leaf spot in Florida (McMillan, 1984), and a gall disease from Cylindrocarpon didymum in Florida causing malformation of trunks and prop roots, with heavy infestations proving fatal (Timyan, 1996).

Hill (2001) reports that R. mangle propagules in Florida are consumed directly by the spotted mangrove crab (Goniopsis cruentata), the mangrove land crab (Ucides cordatus), the coffee bean snail (Melampus coffea) and the ladder horn snail (Cerithidea scalariformis), and leaves are eaten by the mangrove crab (Aratus pisonii), the spotted mangrove crab (G. cruentata), the blue land crab (Cardisoma guanhumi), and various insects.

It is considered that the lower effectiveness of non-indigenous predators in Hawaii may be one reason for the high rate of R. mangle establishment there (Steele et al., 1999).

Means of Movement and Dispersal

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Natural Dispersal

Natural dispersal of R. mangle propagules is facilitated by tides and seasonal flooding (Murray et al., 2003) and severe storms (Proffitt et al., 2006). As propagules were found to float for 8-12 months and still remain viable, R. mangle is likely to be effective at inter-island dispersal, and should be capable of at least occasionally dispersing across the relatively short distances between most of the main Hawaiian Islands (Allen and Krauss, 2006). Egler (1948) describes the processes involved in the dispersal and establishment of mangrove seedlings.

Vector Transmission (Biotic)

There is no evidence of dispersal by animals (Hill, 2001).

Intentional Introduction

R. mangle propagules have been intentionally introduced. Propagules were released in Hawaii in 1902 (and later) to stabilize mudflats that were forming as a result of erosion of agricultural fields (Cox and Allen, 1999). In Mexico it has been planted to improve coastal beach habitats for native fauna and reduce the effects of erosion from severe weather or tidal forces (Mendez Linares et al., 2007), and has also been planted by the aquaculture industry to improve coastal habitat for fisheries and shellfish harvest (Kovacs, 1998).

Impact Summary

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CategoryImpact
Economic/livelihood Positive
Environment (generally) Positive and negative

Environmental Impact

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The environmental impacts of R. mangle and its effects on native biodiversity in Hawaii are reported by many authors (e.g., Allen, 1998; Pratt, 1998; Cox and Allen, 1999; Rauzon and Drigot, 2003; Fronda et al., 2008; Chimner et al., 2006; Demopoulos and Smith, 2010; Siple and Donahue, 2013). It is also worth noting that in its native range, R. mangle stands are also being invaded by other exotic species, for example by exotic mangroves in Florida (Fourqurean et al., 2010).

Many of the habitats typically occupied by mangroves elsewhere in the tropics, as in Hawaii, had no tree cover at all, and as such they represent an entirely new life form in the ecosystems they invade, causing dramatic effects on plant community structure. R. mangle has invaded many coastal areas, transforming sandy habitats into heavily vegetated areas with low water velocity, high sedimentation rates and anoxic sediments (Siple and Donahue, 2013).

Known negative impacts include reduction in habitat quality for endangered waterbirds, affecting the nesting habitats of some endangered waterbirds, including the Hawaiian duck (Anas wyvilliana), Hawaiian coot (Fulica alai), Hawaiian stilt (Himantopus mexicanus knudseni) and Hawaiian moorhen (Gallinula chloropus sandvicensis) (Chimner et al., 2006), reducing native species in fish ponds (Pratt, 1998) and anchihaline pools that have a large algal component and a highly specialized and vulnerable fauna, overgrowing native Hawaiian archaeological sites (Fronda et al., 2008), and causing drainage and aesthetic problems (Allen, 1998). R. mangle forests also provide habitat for other exotic species including burrowing predators which impact native benthic communities (Siple and Donahue, 2013). Invasive mangroves also facilitate the persistence and spread of introduced species, which may ultimately impact the 500 or so endemic estuarine and marine species in Hawaii (Demopoulos and Smith, 2010).

R. mangle is also likely to have significant negative effects on water quality. Net increases in leaf litter input can decrease dissolved oxygen concentrations, and may compound these impacts in areas with limited tidal flushing by blocking channels and water control structures, further restricting the already limited exchange of water. The most direct impact they have had on endangered waterbirds is the invasion of foraging and nesting habitat where none of the species will forage or nest (Cox and Allen, 1999). R. mangle was observed, however, to be colonized by black-crowned night-herons (Nycticorax nycticorax hoactli) and cattle egrets (Bubulcus ibis) (Rauzon and Drigot, 2003). Also, MacKenzie and Kryss (2013) suggest that exotic mangroves in Hawaii do not have adverse effects on native fish assemblages in tide pools, providing nursery habitat for both native and exotic fishes, and the presence of vegetation may actually be influential on exotic fish assemblages.

R. mangle removal results in gradual changes in community composition and an increase in total faunal abundance, a decrease in subsurface deposit feeders, and an increase in suspension-feeding worms (Siple and Donahue, 2013). However, the long-term impact of mangrove removal on Hawaiian stilt populations is still not clear (Rauzon and Drigot, 2002).

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Pioneering in disturbed areas
  • Highly mobile locally
  • Long lived
  • Fast growing
  • Has high reproductive potential
  • Has propagules that can remain viable for more than one year
  • Reproduces asexually
Impact outcomes
  • Ecosystem change/ habitat alteration
  • Monoculture formation
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Rapid growth
Likelihood of entry/control
  • Highly likely to be transported internationally deliberately
  • Difficult/costly to control

Uses

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R. mangle has a variety of uses, but is used primarily as a source of timber for construction, fencing, firewood and charcoal, for medicinal uses, as a source of tannin, and as a habitat for commercial fisheries and aquaculture (Kovacs, 1998).

The timber is durable and very hard with a specific gravity of 0.89 (0.7-1.2). The sapwood is yellowish to light brown, greyish or pinkish, the heartwood dark red to reddish brown with a fine to medium texture, having a straight to irregular grain, low lustre and no odour, and radial, tangential and volumental shrinkage values of 5.0, 10.7 and 14.3, respectively (Timyan, 1996). The wood is, however, susceptible to attack by dry-wood termites (Little, 1983). It is used as roundwood for posts and poles, for marine piling and wharves, shipbuilding, and in cabinetwork; it also produces excellent firewood and charcoal (Little and Skolmen, 1989), and is also occasionally exploited as a source of pulp wood (Duke, 1983). Timber production is estimated at 150 m3/ha per year (Duke and Allen, 2006).

Cattle will eat the leaves if lime is added to raise the pH, and leaves could serve as a valuable source of feed but this potential has yet to be realized (Duke and Allen, 2006). Leaves contain around 10% protein. Various human uses have been reported in the literature, including an alcoholic drink, a famine food from the fruit and young stems, and dried leaves have been used in Florida as a tobacco substitute. Fibres from the branches and roots have been used to make fishing lines. The bark is important commercially in tanning leather, the leaves are also rich in tannin, and dyes are obtained from the bark (Little and Skolmen, 1989). Dry bark contains 10-40% tannin and aerial roots around 10% (Duke, 1983).

Extracts from R. mangle are reported to have various medicinal uses, including as a treatment for diarrhea, dysentery, fevers, eye ailments, skin disorders and a range of other diseases, though their effectiveness is not verified, but research does show that bark extracts reduce gastric ulcers, and have antimicrobial and antioxidant properties (Berenguer et al., 2006). Plant extracts have also proved effective against the pest Cylasformicarius (Williams, 1999).

R. mangle is also used for ecological and landscape maintenance and stabilization, and is commonly used to stabilize coastal mudflats and reduce erosion (Cox and Allen, 1999).

Similarities to Other Species/Conditions

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R. mangle is easily recognized from most other species by the mass of peculiar, branching, curved and arching stilt roots that enable the trees to spread in shallow salt and brackish water, forming dense, impenetrable thickets. Similar species include Avicennia germinans, Laguncularia racemosa, Rhizophora racemosa, Rhizophora samoensis and Rhizophora harrisonii. However, R. mangle has somewhat larger and shinier leaves than these other mangroves, and is further distinguished by its long and pencil-shaped ‘fruits’. For more detail on the morphological separation of R. mangle from closely related R. racemosa, R. samoensis and R. harrisonii, see Duke and Allen (2006).

Prevention and Control

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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Physical/Mechanical Control

Duke and Allen (2006) note that seedlings may not be able to survive on sites where there is a high presence of grazing animals, and often trees will die if more than 50% of the leaves are removed for any reason.

Physical control options are expensive but have proved to be moderately to highly effective in Hawaii, where cutting by hand or with machines and dredging all reduce tree numbers and allow native species to return (Allen, 1998). Cutting trees 10 cm above substrate level or to the waterline in stands is likely to kill plants outright. Rauzon and Drigot (2003) report the results of 20 years of work using thousands of volunteer hours and more than US$2.5 million of contracted labour to remove 8 hectares of R. mangle in a fishpond complex in Hawaii, using hand tools, chainsaws and heavy equipment.

Biological control

A possible biological control agent for R. mangle in Hawaii suggested by Wier et al. (2000) is the fungal pathogen Cytospora rhizophorae which forms cankers on the stem tissue and causes 33% mortality in seedlings when inoculated in the field.

Chemical control

Walsh et al. (1973) found that a combination of 2,4 D and picloram killed all seedlings, and noted previous reports using bark applications of 2,4-D to related species. Basal treatments of triclopyr are also reported to be effective on R. mangle.

References

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Contributors

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17/04/2015 Original text by:

Nick Pasiecznik, Consultant, France

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