Rhizophora mangle, R. samoensis, R. racemosa, R.

× harrisonii 
(Atlantic–East Pacifc red mangrove)
Rhizophoraceae (mangrove family)
American mangrove (English, Australia); red mangrove (USA); mangle rojo (Central and Latin America, Pacifc and
Caribbean coasts); tiri wai (Fiji); togo (Samoa)
Norman C. Duke and James A. Allen
p
h
o
t
o
:

N
.

C
.

D
u
k
e
Atlantic–East Pacifc red mangrove Rhizophora mangle growing
along an estuarine shoreline near Braganza, Amazonian Brazil.
Species Profles for Pacifc Island Agroforestry
www.traditionaltree.org
April 2006
ver. 2.1
IN BRIEF
Distribution Native to American west and
east coasts and African west coast. One species,
Rhizophora mangle, was introduced to the cen-
tral Pacifc, including Hawai‘i and the Society
Islands. Closely allied with Indo–West Pacifc
stilt mangroves whose ranges naturally overlap
AEP mangroves only in the southern Pacifc.
Size Can reach 30–50 m (100–160 ft) in height,
although commonly attains 5–8 m (16–26 ft).
Habitat Inhabits the intertidal wetland zone,
0–6 m (0–20 ft) elevation between mean sea
level and highest tides, with variable rainfall.
Vegetation Commonly associated with other
mangrove species.
Soils Adapted to a very wide range of soils but
thrives best in fne mud sediments of down-
stream river estuaries.
Growth rate Grows less than 1 m/yr (3.3 ft/yr)
in height.
Main agroforestry uses Soil stabilization,
coastal protection, wildlife/marine habitat.
Main products Timber, fuelwood, charcoal,
dyes, and traditional medicine.
Yields Timber volume was estimated at 100–
150 m
3
/ha/yr (1400–2100 ft
3
/ac/yr).
Intercropping Recommended for planting to-
gether with other mangrove species.
Invasive potential Tese plants are ready col-
onizers of new mud banks, making them op-
portunistically invasive with a high potential to
invade alien environments; generally not rec-
ommended for planting outside of their natural
range.
2  Atlantic–East Pacifc Rhizophora species 
INTRODUCTION
Atlantic–East Pacifc red mangroves (AEP Rhizophora
species) are the most important and dominant mangrove
species of tropical coastal areas of the Atlantic Ocean, the
American Pacifc coast, and several islands in the south-
western Pacifc Ocean. Tis species group is one of two
that make up the genus Rhizophora and consists of three
species (two being closely allied) and one hybrid: R. mangle,
R. samoensis, R. racemosa, and R. × harrisonii, respectively.
Red mangroves, notably R. mangle, have also been intro-
duced into new sites in the Indo–West Pacifc (IWP) re-
gion during the past century. In the Hawaiian and Society
Islands, no mangroves were present until introductions
were made in the 1920s, since which time their presence has
become quite noticeable. In the Hawaiian Islands, man-
groves have reportedly overgrown channels, reduced tidal
fows, and overgrown archeological sites. Red mangroves
thrive under a range of intertidal wetland conditions, in-
cluding high salinity levels from greater than full strength
seawater to freshwater (Cintron et al. 1978), and they toler-
ate a range of fooding, soil types, and other physical site
factors. Typically, they are most common in the middle to
low intertidal zone above mean sea level, extending often
along the seaward margin of mangrove stands.
Many of the values of red mangroves are difcult to sepa-
rate from the larger role played by mangrove plants. As
such, the roles of particular species of Rhizophora are often
not distinguished from other members of the genus, in-
cluding the IWP stilt mangroves. Because Rhizophora spe-
cies dominate most tropical mangroves worldwide they are
generally believed to play a vital role for mangrove ecosys-
tems including shoreline protection, enhancement of water
quality in near-shore environments (plus coral reef areas),
and support of estuarine and marine food chains.
Red mangroves are generally considered non-native to
the IWP, not withstanding the disjoint but natural occur-
rence of R. samoensis in New Caledonia, New Hebrides,
Fiji, Tonga, and Samoa. However, where introductions of
R. mangle have been made, the uses of such plants need
to be weighed carefully against their efects as potentially
invasive species. In Hawai‘i, for example, several important
negative efects have been documented, including reduc-
tion in the habitat quality for endangered water birds such
as the Hawaiian stilt (Himantopus mexicanus knudensi),
colonization of habitats to the detriment of native species
(e.g., in anchialine pools), overgrowing native Hawaiian
archaeological sites, and causing localized drainage prob-
lems by reducing the fow through tidal creeks or drainage
channels.
DISTRIBUTION
Native range
Atlantic–East Pacifc red mangroves are native to tropical
regions along the American east and west coasts to the
African west coast, as well as in isolated occurrences in the
southwestern Pacifc islands.
Rhizophora mangle is the species that most characterizes
AEP Rhizophora. It occurs naturally and dominates tropi-
cal tidal areas along both sides of the Atlantic. Te closely
related and almost identical “sibling species,” R. samoen-
sis (=R. mangle sensu lato Tomlinson 1986; =R. mangle var.
samoensis Hochr.), is native to the American west coast and
islands in the southwestern Pacifc, notably New Caledonia,
New Hebrides, Fiji, Tonga, and Samoa. Te taxonomic sta-
tus of these sibling taxa is arguable based on morphological
data alone. Te status of observed diferences in like forms
R. mangle and R. samoensis, however, will only be resolved
in genetic studies and selected breeding programs.
Rhizophora racemosa is less common than R. mangle and
R. samoensis but occurs as a distinct co-inhabitant of man-
WHAT IS A MANGROVE?
Mangroves form a unique and dominant ecosystem
comprised of intertidal marine plants, mostly trees,
predominantly bordering margins of tropical coast-
lines around the world. Tese halophytic (salt toler-
ant) plants thrive in saline conditions and daily inun-
dation between mean sea level and highest astronomi-
cal tides, and they provide vital structure as habitat
and food for similarly adapted resident and transient
fauna. Mangrove plants exchange gases from exposed
roots using special lenticels, while fooding tides allow
uptake of river-borne nutrients and frequent dispersal
by their buoyant propagules. Te ecological limits de-
fned by the diurnal tidal range explain the setting and
why just 70 species around the world are considered
to be mangroves (Tomlinson 1986, Duke et al. 1998),
compared with adjacent rainforests that may have
hundreds of tree species per hectare. Specialized mor-
phological and physiological characteristics largely
defne and characterize mangrove plants, such as but-
tress trunks and roots providing support in soft sedi-
ments, aboveground roots allowing vital gas exchange
in anaerobic sediments, and physiological adaptations
for excluding or expelling salt. Fewer than 22 plant
families have developed such essential attributes, rep-
resenting independent instances of co-evolution over
millions of years to form today’s mangrove habitats.
 Species Prohles for Pacihc Island Agroforestry (www.traditionaltree.org)  3
grove stands in wetter areas and larger catchment estuaries
of the Atlantic. Te species favors riverine estuaries, and
it is restricted in the AEP to a few stands along the Pa-
cifc coast of the Americas and does not occur in the IWP.
At least one other possible species, R. × harrisonii, occurs
across the same range. Given this and its intermediate
characters, the taxon is considered the putative hybrid of
R. mangle and R. racemosa. Te distribution of R. racemosa
and the putative hybrid R. × harrisonii appears restricted
mostly to equatorial estuaries of larger river systems with
more continuous freshwater fows.
Of great interest also is the natural presence of red man-
grove in the southwestern Pacifc islands. Also occurring
in this region is another hybrid taxon, R. selala (Salvoza)
Tomlinson, whose putative parents include R. samoensis
(=R. mangle var. samoensis) and R. stylosa (=R. mucronata
var. stylosa). Tis hybrid taxon is special because R. stylosa
is a dominant member of the IWP stilt mangroves. Te oc-
currence of this hybrid means that there appears to be very
little genetic separation between these defning and most
divergent of Rhizophora species.
Current distribution
Because red mangroves are recognized as valuable timber
producers, benefcial to shoreline stabilization and fsher-
ies, it is feasible that their dispersal westward in the Pa-
cifc may have been assisted by aboriginal travelers in the
past. However, there is no evidence for this happening, so
currently there is no agreed explanation for the presence
of R. samoensis in the southwestern Pacifc. In the absence
of human intervention, and accepting that long-distance
dispersal was not possible, one explanation is island-hop-
ping by natural dispersion across an ancient archipelago
of seamounts that crossed the southeastern Pacifc during
early formation of the Pacifc Plate in the Late Cretaceous
period (Duke 1992, 1995). Any migration would have been
westward, with distances between the putative ancient is-
lands large enough to restrict all mangrove species except
the dispersal-specialist red mangroves. In support of this
theory, there are tantalizing similarities in cross-ocean
linkages for several shallow-water reef fsh.
Red mangroves extend notably beyond their native range.
Rhizophora mangle has been introduced to the Hawaiian
and Society Islands from Atlantic populations in Florida.
In each case, founder populations have increased and ex-
panded dramatically, especially in the Hawaiian Islands.
Plants introduced in the early 1900s to Moloka‘i and O‘ahu
now extend to most islands of the group, and the expecta-
World distribution of red mangroves, the AEP Rhizophora species. Given the apparent hybrid status of R. × harrisonii, its dis-
tribution is likely where the distributions R. racemosa and R. mangle/R. samoensis overlap. Coastlines marked in red indicate the
distribution of all mangroves. image: N.C. Duke
4  Atlantic–East Pacifc Rhizophora species 
tion is that they will spread further. Accordingly, R. mangle
is treated as an invasive species in these islands.
Similar introductions of R. mangle elsewhere are appar-
ently less invasive, possibly because other mangroves
that already occupy the habitat reduce establishment op-
portunities for any new introductions. In the Townsville,
Australia, area, local authorities were not prepared to take
the risk, however, so when a small introduced stand of R.
mangle was discovered in the upper part of Ross River, it
was destroyed.
BOTANICAL DESCRIPTION
Preferred scientifc names
Rhizophora mangle L.
Rhizophora samoensis (Hochr.) Salvoza (=R. mangle var.
samoensis Hochr.)
Rhizophora racemosa G.F.W. Meyer
Rhizophora × harrisonii Leechman (=R. mangle × R.
racemosa)
Family
Rhizophoraceae (mangrove family)
Common names
Atlantic–East Pacifc red mangroves, American mangrove
(English)
red mangrove (USA)
mangle rojo (Central and Latin America, Pacifc and
Caribbean coasts)
tiri wai (Fiji)
togo (Samoa)
Size
Atlantic–East Pacifc red mangroves are medium to tall
trees. Tey may reach 30–50 m (100–160 ft) in height, al-
though they are commonly much shorter, around 5–8 m
(16–26 ft). Stem diameters are about 15–35 cm (6–14 in)
taken just above the highest prop root. Tis method of
measurement difers fundamentally from the standard di-
ameter at breast height (dbh) used for most forest surveys,
as diameter height above the substrate varies from 0.5–7
m (1.5–23 ft) (consider the tree in the photo on the frst
page).
Form
Red mangroves are often multi-stemmed rambling to co-
lumnar trees with distinct aboveground prop roots. Tey
tend to be of shorter stature and more spreading in shape
on the seaward edge of stands or in sites of higher salinity.
Taller, single-stemmed trees are often found just behind
the seaward edge of stands downstream in major river es-
tuaries. Multi-stemmed trees occur in frontal areas but are
more common in upper intertidal regions. Prop roots are
sturdy even when relatively thin.
Flowering
Flowers are perfect, containing both male and female parts.
Inforescences have few to many joints with 1,2,3,4–choto-
mous branching and one to many buds possible per in-
forescence. Open fowers are located within or below leaf
axils at leaf nodes below the apical shoot, depending on
species. For R. mangle, R. × harrisonii, and R. racemosa,
mature buds and fowers are located at 1–2, 3–5, and 7–9
nodes down from the apical shoot, respectively. Te ca-
lyx is typically waxy yellow to creamy white and green at
maturity, with four lobes. Buds elongate to ovate, green
when immature to lighter colors as they mature, smooth,
dimensions 1–2 cm (0.4–0.8 in) long, ~0.5 cm (0.2 in) wide.
Petals, usually four, are lanceolate to linear, creamy white,
with woolly to sparsely hairy margins, ~12 mm (0.5 in) long
and ~4 mm (0.16 in) wide. Stamens number eight, pale
yellow, to golden brown at maturity. Style is pale green, te-
rete, fliform, 0.5–4 mm (0.02–0.16 in) above ovary base,
1.5–3 mm (0.06–0.12 in) wide, dichotomous tip, pale yel-
low. Bracts and bracteoles are distinct. Peduncle is 3–4 cm
long (1.2–1.6 in), ~0.3 cm (0.01 in) wide. Flowering period
is chiefy (but not exclusively) August–December in the
southern hemisphere and February–June in the northern
hemisphere.
Leaves
Leaves are opposite, simple, bright green, obovate, leathery,
margins revolute, generally curved surface, obtuse blunt
apex with a minute lip folded under. Cork wart spots occur
on most species’ leaf undersurfaces, scattered evenly, not
raised, reddish brown (may be distinguished from infec-
tions and wounds by their uniform cover). Note that non-
spotted leaves are found on an unusual form of R. racemosa
in northern Brazil. Leaf emergence occurs chiefy during
November–February in the southern hemisphere and May–
August in the north. Leaf fall occurs chiefy over the wet
summer period, October–February in the southern hemi-
sphere and April–August in the northern hemisphere.
Fruit
Fruits, when mature, are pear-like, elongate, waist constric-
tion, smooth brown surface, calyx lobes elongate spreading
(when the hypocotyl is ready to emerge). For R. mangle,
R. × harrisonii, and R. racemosa mature fruit located in leaf
 Species Prohles for Pacihc Island Agroforestry (www.traditionaltree.org)  5
axils 3, 7 (rare) and 10–11 nodes down from apical shoot,
respectively.
Seeds/hypocotyls
Like all Rhizophora species, red mangroves are viviparous,
meaning that they produce seeds hidden in the mature
fruit, and these germinate while the fruit is still attached to
the parent plant. Te dispersal unit, a viviparous seedling,
is called a hypocotyl. One hypocotyl is usually produced
from each fruit, although on rare occasions twins may be
observed.
Hypocotyls are narrowly cylindrical, elongate, green,
smooth with irregular small brown lenticels, distal half
wider, distal tip pointed. Rhizophora mangle and R. samo-
ensis both have distinct brown distal portions of other-
wise green, relatively short hypocotyls (see photo). For R.
mangle, R. × harrisonii, and R. racemosa, mature hypocotyls
are located in leaf axils 3–8, ~8 (rarely observed), and 13–15
nodes down from the apical shoot, respectively.
“Fruiting,” when mature hypocotyls fall, occurs chiefy (but
not exclusively) in November–January in the southern
hemisphere and May–July in the northern hemisphere.
Bark
Bark is smooth and red-brown in seaward and exposed lo-
cations (rocky and sandy substrates), to gray-fssured with
smooth, red-brown prop roots in sheltered locations (mud
substrates). Tere is total coverage of gray-fssured bark in
some localities.
Top left: Te blunt apex of leaves is characteristic of all AEP red mangroves, as in this R. samoensis from Fiji. Top right: Leafy ro-
sette of Rhizophora mangle (similar to R. samoensis) showing open fowers with elongate, refexed calyx lobes. Bottom left: Open
fower of Rhizophora racemosa showing fat, slightly hairy petals and stif-erect, non-refexed calyx lobes. Bottom right: Mature
fruit of Rhizophora mangle (similar to R. samoensis) showing its elongate pear-shape, prior to emergence of the hypocotyl. Note
the persistent style at the distil tip of the fruiting body. Atlantic Panama. photos: N.C. Duke
6  Atlantic–East Pacifc Rhizophora species 
Rooting habit
Mature trees have distinctive, sturdy, aboveground prop
roots surrounding the stem base that anchor only shal-
lowly in the sediments to 1–2 m (3.3–6.6 m) depth. Tis
conforms to the anoxic conditions commonly observed in
mangrove sediments.
Similar species
Red mangroves are distinguished from IWP stilt man-
groves principally by the leaf apex: red mangroves have
a blunt leaf apex and lack the spiked, mucronate leaf tip
present in stilt mangroves.
Rhizophora mangle and R. samoensis, with mostly 0–3 in-
forescence joints, are distinguished from R. racemosa and
R. × harrisonii, which have 3–8 inforescence joints.
Rhizophora racemosa is distinguished from R. × harrisonii
by the node position of mature buds and fowers in leaf
axils, down from the apical shoot, being 7–9 nodes and
3–5 respectively. Te possible hybrid character is shown in
R. × harrisonii, where it has characters intermediate be-
tween R. racemosa and R. mangle.
Rhizophora mangle and R. samoensis, the sibling species, are
distinguished by minute/absent bracts on pedicles at the
calyx base in R. mangle, while R. samoensis has bracts twice
as wide as the pedicle. As noted, R. mangle and R. samoen-
sis appear very closely related and are likely to be the same
species. Discriminating between them reliably is not pos-
sible in many instances without detailed examination of
key morphological and genetic characteristics.
GENETICS
Variability of species
Atlantic–East Pacifc red mangroves are those Rhizophora
species that occur naturally along the east and west coasts
of the Americas, as well as along the west African coast
and those island populations in the southwestern Pacifc.
In this group there appear to be four relatively distinct taxa,
although at times their morphological and taxonomic dif-
ferences are questionable. Te uncertainty is chiefy based
on: 1) the presence of one intermediate individual that is
recognized as a distinct hybrid, namely R. × harrisonii, and
2) the occurrence of two sibling species, R. mangle and R.
samoensis, which may, on closer examination, be shown to
be the same species.
Te relationship of R. mangle and R. samoensis is perhaps
the most contentious. Tey appear very closely related, as
they are distinguished by only a few morphological charac-
ters, as well as their distinct geographic ranges. Te sibling
Mature hypocotyls of Rhizophora mangle (similar to R. samo-
ensis), showing distinctive brown distal end, Hawai‘i. photo:
N.C. Duke
Expended fruits of Rhizophora samoensis (left) and R. stylosa
(right), Fiji. photo: N.C. Duke
 Species Prohles for Pacihc Island Agroforestry (www.traditionaltree.org)  7
species separate naturally across the American land barrier,
with R. mangle in the Atlantic and R. samoensis in the Pa-
cifc. Te situation became more complex during the 1900s
when R. mangle (mostly from around Atlantic Florida)
was introduced to the Pacifc in several isolated instances,
notably to Hawai‘i, the Society Islands, and northeastern
Australia. Te Australian introduction was eradicated a
few years ago, but in both other locations the spread of
R. mangle has proceeded unchecked, and the species now
dominates most intertidal wetland areas of each location.
In Hawai‘i changes in shoreline ecology have been dra-
matic, especially because no mangroves grew in this iso-
lated location previously. A similar situation is expected in
the Society Islands, but no detailed reports are available.
Tese introductions highlight the unexplained disconti-
nuity of R. samoensis in the southwestern Pacifc islands.
Tis is the only example where AEP and IWP Rhizophora
species naturally co-exist. Curiously, the human introduc-
tions of R. mangle demonstrate that island habitats with-
out mangroves existed between Samoa and South America,
and that these are suitable for mangrove colonization. Te
question raised by this natural discontinuity remains unre-
solved. Tis observation emphasizes the profound limita-
tions of long-distance dispersal for this mangrove group of
otherwise long-distance dispersal specialists.
Recognizing the morphological diferences between R.
mangle and R. samoensis is considered useful in distinguish-
ing between introduced populations of R. mangle in the
Hawaiian and Society Islands, as compared with natural
populations of R. samoensis where they occur elsewhere in
the Indo–West Pacifc.
By contrast, R. racemosa is readily distinguished from R.
mangle and R. samoensis. Tis species appears more com-
mon in estuaries infuenced by larger and more continu-
ous freshwater fows in equatorial regions of three distinct
regions: the east Pacifc (Costa Rica to Ecuador), west At-
lantic (Venezuela to Brazil), and east Atlantic (western Af-
rica). Tere are no reports of diferences within this taxon
between these areas, but recently (N.C. Duke, pers. obs.),
two forms of R. racemosa were found to co-exist in north-
ern Brazil. One fts the type description in every respect
including the presence of cork wart spots on leaf under-
surfaces, while the other form lacked these spots. Tis was
particularly notable since all red mangroves until now have
been reported to possess such spots on their leaf under-
surfaces. Tis character is present in IWP stilt mangroves
and in a similar way defnes the two forms of R. apiculata
occurring in Australasia and the northwestern Pacifc.
Known varieties and hybrids
Rhizophora harrisonii is the apparent hybrid of R. mangle
and R. racemosa based on its intermediate and shared mor-
phological characteristics. Further investigations are need-
ed to adequately describe Rhizophora taxa and their dis-
tributions throughout the AEP region. Current evidence
indicates the situation may be more complex, with another
possible hybrid where R. racemosa apparently occurs on the
eastern Pacifc coast of the Americas with R. samoensis, the
allied partner to R. mangle.
Furthermore, as noted above, two forms of R. racemosa
were observed in Brazil. Te same recent investigation
also discovered two intermediate forms, recognized as po-
tential hybrids between the two R. racemosa forms and R.
mangle. Key questions arise from these new observations
including: what is the distribution of the two forms of R.
racemosa, and how do these compare with populations in
western Africa? Tere are clearly more questions than an-
swers concerning genetic variation in red mangroves.
Aberrant individuals are reported to produce chlorophyll-
defcient propagules, called albinos. Tis is particularly no-
table and common in R. mangle stands throughout the Ca-
ribbean area to northern Brazil. Yellow or red propagules
can be observed hanging in afected trees alongside normal
green propagules. Teir relative numbers on an individual
tree are believed to quantify the amount of outcrossing
that occurs among neighboring normal trees. Other trees
have also been observed with variegated foliage.
Culturally important related species
All Rhizophora species are closely similar in tree form, and
cultural groups in the Pacifc region may not always dis-
tinguish between them. Other mangrove genera, such as
Bruguiera, are considered close in form and value also, and
these together are often used in similar ways by indigenous
peoples.
ASSOCIATED PLANT SPECIES
AEP red mangroves can be found growing with other man-
grove species, preferably within intertidal wetland condi-
tions above mean sea level elevations in tropical regions. R.
mangle and R. samoensis grow in marine saline conditions,
while R. racemosa forms and hybrid grow in proximity to a
regular freshwater source. R. racemosa are commonly found
in lower to middle tidal reaches of rivers and streams with
regular, frequent freshwater runof, while R. mangle and R.
samoensis occur in coastal embayments and coral islands. In
Fiji, however, R. samoensis occurs together with R. stylosa (a
stilt mangrove). In this case, R. samoensis occurs more fre-
8  Atlantic–East Pacifc Rhizophora species 
quently in upstream locations of the freshwater dominated
streams, while R. stylosa remains the marine specialist, pre-
ferring downstream and exposed coastal locations.
Red mangroves often extend up the tidal profle to the
terrestrial fringe in areas of higher rainfall. In lower-rain-
fall areas, red mangroves occur on estuarine margins just
above mean sea level. Best development has been observed
in lower tidal reaches and on soft, fne, muddy sediment
substrates.
Associated species commonly found
In Atlantic and east Pacifc populations, red mangroves
occur commonly with Avicennia germinans and Laguncu-
laria racemosa. In the southwestern Pacifc, R. samoensis is
associated with IWP mangroves, including R. stylosa and
Bruguiera gymnorrhiza predominantly. In other locations,
red mangroves are associated with other mangrove spe-
cies. Te types of associated species are notably variable
where this species has been introduced to the IWP region.
In Hawai‘i, while the species frequently occurs in nearly
mono-specifc stands, it does sometimes occur alongside a
mangrove introduced from the IWP (Bruguiera sexangula)
and species that typically occur in lower-salinity environ-
ments, such as Hibiscus tiliaceus.
ENVIRONMENTAL PREFERENCES
AND TOLERANCES
Climate
Atlantic–East Pacifc red mangroves thrive in tropical and
subtropical environments characterized by moderately
high and well distributed rainfall. Mangrove plants appear
to depend on groundwater to sustain optimal growth, es-
pecially during drier months. In drier locations, such as
the Baja Peninsula on the northern Pacifc coast of Mexico,
the stunted but dense thickets of R. samoensis (=R. man-
gle?) attest to the great adaptability of red mangroves to a
wide range of climatic types.
Ilevation range 
0–6 m (0–20 ft), in reference to mean sea level.
Mean annual rainfall 
Tese mangroves grow in all rainfall conditions. Teir ex-
tent, form, and biomass refect the diferent rainfall condi-
tions.
Rainfall pattern 
Tey grow in climates with summer or uniform rainfall
patterns.
Ðry season duration (consecutive months with <40 
mm l1.6 in| rainfall) 
Because of access to groundwater, red mangroves grow in
regions with up to 8 months of drought. Across a wide va-
riety of climatic regions, mangrove cover expands and con-
tracts through time. Tis has been evident in correlations
between El Niño events and reduced growth as possible
causes of some damage to mangroves, presumably due to
decreases in freshwater availability.
Mean annual temperature 
20–30°C (68–86°F) (estimate)
Mean maximum temperature of hottest month 
32–38°C (90–100°F) (estimate)
Mean minimum temperature of coldest month 
0–5°C (0–41°F)
Minimum temperature tolerated 
0°C (32°F) (estimate)
Soils
Trees develop greatest stature and columnar growth form
in estuaries of larger tropical rivers, characterized by fne
clay, black mud sediments with relatively high loads of or-
ganic carbon. Tey are also anaerobic with high concentra-
tions of sulfde. Trees also grow well in sites with aerobic
sediments consisting of fne sands to coarse stones and
rocks, and coral ramparts.
Soil texture 
Plants grow in light, medium, and heavy texture soils
(sands, sandy loams, loams, and sandy clay loams, sandy
clays, clay loams, and clays).
Soil drainage
Te trees grow in soils with free and unimpeded drainage,
as well as waterlogged soils.
Soil acidity 
pH 6–8.5
Special soil tolerances 
Plants grow best in saline soils but survive well in fresh wa-
ter. Te optimal salinity range is reported to be from 8 to
26 ppt (parts per thousand), compared with approximately
34–36 ppt for seawater.
 Species Prohles for Pacihc Island Agroforestry (www.traditionaltree.org)  9
Tolerances
Ðrought
Red mangroves can tolerate drought periods well, although
trees have apparently been killed by drought in some sites
on Moloka‘i, Hawai‘i where groundwater sources appear
to be limited. Drought conditions presumably cause soil
salinity to increase in excess of tolerable limits for these
plants.
Iull sun
Tey grow best in full sun.
Shade
Tey appear to have low tolerance of shade. However, re-
cent evidence shows this more likely due to weevils that
infest and kill cooler, shaded seedlings (Brook 2001, Sousa
et al. 2003).
Iire
Red mangroves have no tolerance of fre in close proxim-
ity.
Irost
Tolerance of sub-freezing temperatures is low to none.
Waterlogging
Red mangroves are tolerant of daily tidal fooding up to
depths of up to 1.5 m (5 ft). While tolerating permanently
saturated soils, they are intolerant of drying soils.
Salt spray
Te trees are highly tolerant of salt spray.
Wind
Red mangroves are typically found in seaward areas subject
to wind and salt spray but largely protected from waves.
Planting in such highly wind-prone locations is recom-
mended, but only where required and within the tree’s na-
tive range.
Waves
In general, exposed, wave-prone coastlines are inhospitable
to mangroves. In areas where wave action is infrequent, red
mangroves are believed to provide signifcant bufering of
coastal areas during storm and tsunami surge events (Dah-
douh-Guebas 2005).
Abilities
Self-prune
Red mangroves self-prune well in dense stands, but they
commonly maintain lower branches in more open-grow-
ing locations.
Coppice
Te trees have notably poor coppice ability. Generally, if
50% or more of the leaves are removed from a tree, it will
die.
GROWTH AND DEVELOPMENT
Growth rates vary with age. Tey generally grow less than
MANGROVES AND CLIMATE
CHANGE
Atlantic–East Pacifc red mangroves, like other man-
grove species, are afected by climate change. Te
unique physiological characteristics of each species
defne its capacity for survival in the face of change.
Mangroves are expected to respond rapidly and deci-
sively to shifts in key factors, like temperature, rainfall,
and sea level, as each species has defned ranges of tol-
erance for each factor. For instance, because mangroves
are characteristically restricted to elevations between
mean sea level and highest tides, as sea level rises their
communities must move upland to survive. Since man-
groves have narrow optimal temperature ranges, ris-
ing temperatures will cause their distributions to shift
north or south to areas where temperature conditions
are most suitable, and they will die of in areas where
they are not suited. Of course, their success in making
these shifts depends on their successful dispersal and
re-establishment, and the availability of suitable new
space. Clearly, such changes have occurred throughout
history, so the distribution of mangroves today repre-
sents the survivors of all past changes.
Key indicators of change can be identifed and mapped
as incremental shifts and responses of mangrove com-
munities. Tese might be observed as shifts in vegeta-
tion, for example: 1) in the total tidal wetland habitat
zone, as expected with changes in sea level; and 2) in
the salt marsh–mangrove ecotone, as expected with
changes in longer-term rainfall patterns as this afects
moisture stress in saline environments. In both cases,
the response zones will follow elevation contours.
Changes along contours can be quantifed from long-
term spatial assessments over decade- and century-
long time periods, depending on the rates of change.
Knowledge of these changes and their causes allows
better prediction of future change.
10  Atlantic–East Pacifc Rhizophora species 
1 m/yr (3.3 ft/yr) but can exceed this in favorable circum-
stances. Height growth is rapid shortly after establishment
while food reserves are taken up from the hypocotyls of
established seedlings. Growth rate slows when trees ap-
proach a site-specifc maximum canopy height. When near
maturity and maximum height, trees broaden their cano-
pies and increase stem diameter rather than grow taller.
Flowering and fruiting
Flowering and fruiting periods of red mangroves are dis-
tinctly seasonal. Peak pheno-events are expected to shift
into later months with cooler temperatures and higher
latitudes. Trees have notable and relatively long periods of
reproductive development, taking 18–30 months from frst
emergence of fower bud primordia until maturation and
drop of mature hypocotyls. Te duration depends on spe-
cies, with the longest being R. racemosa with an expected
period of around 30 months for each reproductive cycle.
Yields/growth rates
Growth rates vary with species, spatial position in the stand,
competition, vigor, and age. For northern Pacifc sites in
Costa Rica, Jimenez (in Chong 1988) reported annual di-
ameter growth increments for R. mangle and R. racemosa
trees of 0.14–0.19 cm (0.06–0.07 in) for diameter size
classes less than 10 cm (4 in), and 0.08–0.17 cm (0.03–0.07
in) for diameter size classes greater than 10 cm.
Chong (1988) provides information on potential yields
from Playa Garza Pilot Area in an investigation of the fea-
sibility of managing mangrove forests the Terraba-Sierpe
Forest Reserve on the southern (wetter) Pacifc coast of
Costa Rica. Te estimated mean annual increment (MAI)
was 6 m
3
/ha/yr (86 ft
3
/ac/yr) with some stands as high as
11–14 m
3
/ha/yr (157–200 ft
3
/ac/yr).
Reaction to competition
Rapid early growth of red mangrove seedlings in full sun-
light ensures their success and dominance in preferred es-
tuarine and intertidal conditions. Newly established seed-
lings grow best in close proximity with their same species
cohort. Tis afords them maximal protection from physi-
cal damage by drift logs and erosive waves. Since competi-
tion is high between neighboring seedlings, slower plants
die and decompose quickly, leaving faster competitors the
beneft of not only the space they occupied but also their
nutrients.
PROPAGATION
Although natural regeneration is generally relied upon
from Pacifc to Atlantic regions, these species are relative-
ly easy to propagate. Propagation is simple and relies on
the special feature of the genus in having large viviparous
propagules. Planting simply entails gently pushing the dis-
tal end of the 20–60 cm (8–24 in) long hypocotyl one third
of its length into the sediment, spaced at about 1–1.5 m
(3.3–5 ft) intervals. No holes need be dug, and neither nurs-
ery preparation nor stakes are needed. Low maintenance is
generally sufcient for maximizing seedling establishment
success in sheltered areas. However, substantial protection
is required in more exposed coastal locations during the
frst decade of establishment. Such protection methods
may include encasement of individual seedlings in PVC
piping (Riley and Kent 1999), or installation of temporary
structures to dampen wave action and reduce debris drift
across restoration sites, as observed in Vietnam and China
(Field 1996).
Propagule collection
Propagules may be available throughout the year, but peak
production occurs around July–August in northern parts
of the range and around January–February in the southern
hemisphere. Propagule maturation occurs later in relatively
higher latitude locations north or south beyond the tropic
zone (e.g., July–October in Hawai‘i).
Mature propagules may be collected from the ground after
they have fallen or picked directly from trees. Only healthy
looking propagules should be selected. Propagules that
are shrunken or desiccated in appearance or that exhibit
signifcant physical damage should be rejected. Although
propagules with only minor borer damage may survive and
grow, selection of propagules with no signs of borer or crab
damage are strongly preferred. Propagules that already
have some root or leaf development can be used in most
cases, but should not be stored for long.
Propagule processing
Processing of mature propagules is not required for red
mangroves, although damaged and insect-infested indi-
viduals should be removed. Te following pretreatment
section gives possible additional steps. Propagules can be
sown in nursery beds, or preferably planted directly in the
feld soon after collection.
Propagule storage
Propagules can be kept viable for at least 6–7 days by stor-
ing them in brackish water or by wrapping them in wet
burlap bags and keeping them out of direct sunlight. Pre-
treatment is considered essential if considering such stor-
age (see below). It is likely that propagules can be kept
 Species Prohles for Pacihc Island Agroforestry (www.traditionaltree.org)  11
longer, but storage beyond 2 weeks is not recommended,
and long-term storage is not feasible.
Propagule pretreatment
Pretreatment of propagules is generally considered unnec-
essary. However, a study of scolytid beetle larvae (Cocco-
trypes fallax) infections on Rhizophora propagules in Aus-
tralia found at least 18% were heavily infested (Brook 2001).
Infested propagules were established under canopy-shaded
areas. Te study went further to fnd that pretreatment in
a 50°C (122°F) water bath for 5 minutes killed the beetles
and removed the risk of establishment losses due to borer
damage. Heat treatment might be easily achieved by leav-
ing the collected propagules in the sun for a short period
(a few hours) before planting.
Growing area
Red mangroves should be grown in full sunlight.
Seedling establishment
Leaves and roots may begin to develop within a week or
two of sowing.
Media
Although a wide variety of soil media are acceptable, a mix
of sand and peat in equal parts has been recommended for
mangrove nurseries.
Time to outplanting
Seedlings are ready for outplanting at the six-leaf (three-
node) stage if grown up in nursery conditions, which can
take up to 6 months to achieve. Older seedlings up to 0.5
m (1.5 ft) tall have also been successfully transplanted, but
this is not recommended. Direct planting of large numbers
of propagules is restricted by their peak seasonal availabil-
ity, as propagules do not keep for extended periods.
Guidelines for outplanting
Outplanting anytime of the year is suitable. Propagules or
nursery-grown seedlings usually have excellent survival in
sites correctly selected and, if appropriate, protected from
disturbance. Survival rates of 90% or greater are reasonable
in such circumstances.
On the other hand, survival may be zero on sites exposed
to excessive wave action, on sites with inappropriate hy-
drologic or salinity regimes, or (rarely) subjected to distur-
bance by grazing animals (e.g., goats, sheep, cattle, horses).
A method of encasing propagules in PVC pipe has been
used in Florida and the Caribbean to protect seedlings in
places with a high likelihood of disturbance.
DISADVANTAGES
In general, red mangroves pose few signifcant disadvan-
tages when planted within their native range. Tey are not
especially susceptible to pests or pathogens and have not
been reported to host major pests or pathogens of impor-
tant crop species.
Potential for invasiveness
Te very successful introduction and rapid spread of red
mangroves in the Hawaiian Islands clearly demonstrates
the potential for invasiveness of mangroves in areas where
suitable habitat is available. Te plants are readily opportu-
nistic due to their relatively wide tolerance for salinity and
Mature hypocotyl of  Rhizophora  racemosa  showing distinc-
tive, slender, smooth propagule, Brazil. photo: u. mehlig
12  Atlantic–East Pacifc Rhizophora species 
soil conditions. Also, their foating propagules are spread
widely by ocean currents over great distances.
Te species R. mangle has unfortunately now taken on the
public status of invasive weed and pest species in Hawai‘i.
To alleviate public concern, eradication eforts have been
carried out in several locations on O‘ahu and Hawai‘i. It
is not clear about the success or efectiveness of this cam-
paign, as it appears to have been based on subjective infor-
mation and no monitoring.
Pests and diseases
Susceptibility to pests and diseases is believed to be low,
with the exception of insect borers and crabs that prey on
propagules. For introduced stands in the Hawaiian islands,
damage to propagules and leaves is notably lower than
within the species’ native range, and productivity (as ex-
pressed in litter fall) is higher.
Host to crop pests/pathogens
No reports are found of red mangroves serving as hosts for
known major crop pests or pathogens. In Hawai‘i, however,
R. mangle stands have served as ideal sites for the non-na-
tive cattle egret (Bubulcus ibis) and also sometimes harbor
signifcant populations of rats.
AGROFORESTRY/ENVIRONMENTAL
PRACTICES
Mulch/organic matter
Mulch in Rhizophora forests is hidden from view. If it were
not for the small mangrove crabs, fallen leaves would be
washed away with each tide. Te crabs actively take leaves
into underground burrows and chambers. Te resulting
mulch is rapidly colonized by bacteria and consumed by
other burrowing fauna to release nutrients that appear to
further enhance the forest.
Soil stabilization
Red mangrove forests stabilize soils with their network
of sturdy overlapping prop roots, which dampen water
movement and promote sedimentation in areas that might
otherwise be eroded. A major reason that red mangrove
was introduced into Hawai‘i was to stabilize mud fats that
were expanding as a result of sugarcane production and
resultant erosion. Rhizophora mangle has proven quite ef-
fective for this purpose and has been shown to improve
water clarity in near-shore environments, presumably due
to its role in sediment trapping and stabilization.
Ience posts
Red mangrove stems make good posts, as the wood is gen-
erally hard and resistant to insect borers.
Windbreaks
Red mangrove forests provide a windbreak along coastal
margins generally, and places to seek sanctuary during ty-
phoons and hurricanes.
Woodlot
Mangroves adjacent to peoples’ homes throughout the Pa-
cifc frequently serve as informal woodlots, particularly on
islands with clear tenure systems that include mangrove
areas. Red mangrove timber is very useful for small con-
struction and for fuel for cooking. Converting the wood
to charcoal can further enhance the timbers’ calorifc value.
Tis is done commercially in SE Asia and Central America
with various Rhizophora species.
Native animal/bird food
Red mangrove is a known source of native animal foods.
Several observations demonstrate the diversity and quan-
tity, and it is thought to be extremely important in man-
grove ecosystems. Numerous insects, crabs, and mollusks
graze on green leaves in the forest canopy and on fallen
leaves on the forest foor. Where present, sesarmid crabs
consume a large quantity of fallen leaves and propagules.
Organic matter processed by these herbivores is believed to
broadly support aquatic food chains in coastal regions. Few
mammals (probably none in introduced Hawaiian stands)
appear to use red mangrove as a major food source.
Wildlife habitat
In addition to aquatic marine organisms (see Fish/marine
food chain), red mangroves serve as habitat for a wide
range of terrestrial and arboreal wildlife. In various loca-
tions throughout the region, these mangrove forests pro-
vide shelter and food for a number of associated fauna, in-
cluding birds, small mammals, shellfsh, and other marine
life.
Bee forage
Rhizophora species have no nectar, but they do produce
copious pollen that is usually distributed by wind. Inter-
estingly, one reason stated for the introduction of red man-
grove to Hawai‘i was as a “pasture plant for bees” (Cooke
1917). Clearly, it wasn’t the best choice for this purpose!
Iish/marine food chain
Mangroves in general are believed to play a vitally impor-
tant role in protecting and supporting marine food chains.
Many fsh species use red mangroves during part of their
 Species Prohles for Pacihc Island Agroforestry (www.traditionaltree.org)  13
life cycles, as do species of shrimp and crabs. As mentioned
above, senescent leaves that have fallen from Rhizophora
trees are taken by grapsid (small mangrove) crabs into their
burrows. In Hawaiian populations, there appears to be ex-
cessive leaf accumulation in some locations, suggesting
that normal associated fauna and other decompositional
biota are lacking.
Coastal protection 
Red mangrove forests and mangroves in general play an
important role in protection of coastlines, fshponds, and
other coastal infrastructure. Red mangroves are planted for
coastal or fshpond protection in some areas (e.g., in Ter-
raba-Sierpe Forest Reserve, Costa Rica), and there are laws
in many locations aimed at protecting mangroves in large
part because of this important function.
USES AND PRODUCTS
Red mangroves are probably of greatest value for their en-
vironmental benefts, because they (and mangroves gener-
ally) are believed to play a vital role in supporting marine
food chains, protecting coastal areas, and improving water
quality.
In terms of direct benefts to people, the most widespread
use of red mangroves is for wood used for a range of pur-
poses from cooking fuel to construction of homes and ca-
noe parts. Other uses of the tree include dyes, medicines,
and tannin for tanning leather. It seems likely that red
mangroves may have several other uses (e.g., as cattle feed)
that to date have not been fully explored.
Staple food 
Leaves and hypocotyls are edible but not widely used for
food.
Medicinal 
Red mangrove bark has reportedly been used to treat an-
gina, boils, and fungal infections. Te leaves and bark have
been used as an antiseptic and to treat diarrhea, dysentery,
fever, malaria, and leprosy, although it is not clear how ef-
fective the treatments have been in these cases.
Animal fodder
One report (Morton 1965) concluded that red mangrove
leaves might serve as a valuable source of cattle feed, but
this potential has yet to be realized.
Timber
Te wood of red mangroves is widely used for structural
components of traditional homes (e.g., poles, beams, foor-
ing, wall-cladding, rafters) and other components includ-
ing underground mine supports, fencing, cabinet works,
tool handles, and boat anchors. Te wood is also used for
other purposes, ranging from traditional uses such as fsh-
ing stakes, spears, and copra huskers to use as a source of
chips for pulp production.
Te heartwood of R. mangle is light red to dark red or
reddish brown or purplish, with uniform or more or less
striped grain. Te sapwood is yellowish, grayish, or pink-
Benefts include, in no particular order (adapted from
Tomlinson 1986):
• visual amenity and shoreline beautifcation
• nutrient uptake, fxation, trapping, and turnover
• habitat use by fauna
• mesoclimate, where forests moderate evapotranspira-
tion to create a specialized niche climate
• nursery habitat for young fauna, where mangroves
provide a source of food and physical protection from
predation
• sanctuary niche for mature fauna, including migratory
birds and fsh, where mangroves provide protection
and a food resource
• primary production based on photosynthesis, giving
rise to forest growth and forest products, notably tim-
ber
• secondary production, including microbial and faunal
production, as well as grazers, and via decomposition
• fshery products, including both estuarine and coastal
• shoreline protection, based on general mangrove tree
and root structure, as well as edge trees, which reduce
erosion and provide stand protection from waves and
water movement
• carbon sequestration and a sink where carbon is bound
within living plant biomass
• sediment trapping, based on mangroves being a dep-
ositional site for both water and airborne sediments,
which in turn reduces turbidity of coastal waters.
BENEFITS OF MANGROVE TIDAL WETLAND
14  Atlantic–East Pacifc Rhizophora species 
ish. Te wood texture is fne to medium; grain
straight to irregular; low-luster; without dis-
tinctive odor or taste. Te specifc gravity (oven-
dry weight/green volume) is 0.89.
Iuelwood
Red mangrove wood is used for fuelwood in
many places along the American Pacifc coast
(e.g., Panama, Costa Rica, Nicaragua). Te
wood is also made into charcoal in many Cen-
tral American countries, including Panama and
Costa Rica.
Canoe/boat/raft making
Te wood has been used to make canoe parts.
Tannin/dye
Te bark and hypocotyls are used to tan leather
and to produce dyes ranging from red-brown to
black (the latter with repeated dying). Tannin
content of the bark is high for most Rhizophora
species.
COMMERCIAL CULTIVATION
Red mangrove timber is harvested commer-
cially for charcoal production through much of
its range, including Central and Latin America.
Te calorifc value of the timber is signifcantly
enhanced by converting it to charcoal. Tis is
done with all local Rhizophora species, as well
as Pelliciera and Laguncularia.
Rehabilitation and replanting
Projects to replant and rehabilitate mangrove
forests have been conducted where they have
been damaged in a signifcant way. For example,
in Bahia las Minas, Atlantic Panama, a large
oil spill killed around 50 ha (124 ac) of man-
groves in 1986 (Duke et al. 1997), and Refneria
Panama lead a project to replant the damaged
areas. Because the soils were oiled it was decided to plant
R. mangle seedlings using clean terrestrial sediments. To-
tal plantings eventually amounted to in excess of 100,000
seedlings. A subsequent investigation (Duke 1996) found
natural recruitment was 40 times greater than planted
numbers, and natural seedlings grew equally as well as, or
better than, planted seedlings. It was of great importance
that naturally recruited sites recovered more quickly than
planted sites, possibly because of site damage during plant-
ing. Furthermore, damage to exposed and damaged man-
grove areas increased after 5–6 years when dead standing
timber dramatically deteriorated and was moved by wave
action with each tide and storm.
Spacing
Mangrove plantations in general are typically planted at
spacings of about 1.0–1.5 m (3.3–5 ft). Spacing wider than
about 2.5 m (8 ft) tends to result in a high proportion of
multiple stemmed and/or shorter trees. Wider spacing and
therefore spreading trees may be desired for coastal protec-
tion projects but not for timber production. In the absence
Top: Charcoal treatment plant established with the DANIDA Mangrove
Project in Pacifc Costa Rica, supported by IUCN and CATIE. Bottom:
Typical earthen kiln used to convert stacked red mangrove poles into char-
coal for local heating needs, Pacifc Panama. photos: N.C. Duke
 Species Prohles for Pacihc Island Agroforestry (www.traditionaltree.org)  15
of signifcant natural mortality, timber plantations should
be thinned to spacing of 2.5 to 3.5 m (8–11 ft) between trees
as the stand develops and becomes crowded.
Management objectives and design consider-
ations
Some published guidelines for mangrove silviculture exist
and are referenced below, but specifc guidelines on thin-
ning, fertilizing, etc., are lacking.
Growing in polycultures
Red mangroves naturally occur in mixed-species stands,
and each species has its own ecological and economic val-
ues. It is also important to plant associated bufer areas,
particularly along the shoreline where mangroves grow
better adjacent to banks stabilized by shoreline upland
plants. Together they will complement and enhance the
richness and stability of the planted environment.
Estimated yield
For the southern (wetter) Pacifc coast of Costa Rica,
Chong (1988), in an FAO-sponsored project, provided in-
formation on potential yields from the 240 ha (590 ac) Pla-
ya Garza Pilot Area in an investigation of the feasibility of
managing around 5200 ha (12,800 ac) of mangrove forests
in the surrounding Terraba-Sierpe Forest Reserve. It was
reported that yields resulting from uncontrolled felling of
100 ha/yr (250 ac/yr) could be achieved by felling just 10
ha/yr (25 ac/yr) if conducted more systematically.
Chong estimated that each hectare of man-
grove in this reserve could produce US$619
($250/ac) annually. A detailed management
plan was proposed in the report. Te average
stand volume for red mangroves was 163 m
3
/ha
(2330 ft
3
/ac), with a total mangrove forest stand
volume around 280 m
3
/ha (4000 ft
3
/ac). Esti-
mated yields of timber volume under bark were
100–150 m
3
/ha/yr (1400–2100 ft
3
/ac/yr) for red
mangroves, and for the co-dominant mangrove
Pelliciera, yields are estimated at 60 m
3
/ha/yr
(860 ft
3
/ac/yr).
Markets
Markets for red mangrove products are local
in nature, with little available except for fre-
wood and charcoal in most places, other than
indirect products such as mangrove crabs and
fruit bats. In Central America, large quanti-
ties of red mangrove wood chips and charcoal
may be moved greater distances and in greater
volumes than other wood products. In Panama there is a
strong trade in red mangrove poles around 3 m (10 ft) in
length (around 10–15 years old) for construction of bohio,
the locally popular outdoor recreation shelters for barbe-
cues and parties.
INTERPLANTING
Some interplanting systems include:
Example 1—Bahia las Minas oil spill (Duke 1996,
Duke et al. 1997)
Iocation
Bahia las Minas, Panama Atlantic coast
Ðescription
Planting was undertaken in 1988 and 1989. Around 40 ha
(100 ac) were planted with Rhizophora mangle, sponsored
and implemented by Refneria Panama.
Crop/tree interactions
Not applicable.
Spacing/density of species
Spacing was about 1.5 m (5 ft).
Planting of red mangroves in an attempt to replace around 50 ha of tree
loss following a major oil spill in Bahia las Minas, Atlantic Panama. photo:
N.C. Duke
16  Atlantic–East Pacifc Rhizophora species 
Example 2— Terraba-Sierpe Forest Reserve
(Chong 1988)
Iocation
Costa Rica, Terraba-Sierpe.
Ðescription
Planned project. Tis project reached the planning stages
only and was sponsored by FAO with IUCN and CATIE.
Te species to be planted included R. racemosa, R. mangle,
and Pelliciera rhizophorae.
Crop/tree interactions
Not applicable, as there were no crops.
Spacing/density of species
Spacing was planned to be 1.5 m (5 ft) for all species.
PUBLIC ASSISTANCE
Centre for Marine Studies, Marine Botany Group
Te University of Queensland
St. Lucia QLD 4072
Australia
Web: <http://www.marine.uq.edu.au/marbot/index.htm>
BIBLIOGRAPHY
(☛  indicates recommended reading)
Allen, J.A. 1998. Mangroves as alien species: Te case of
Hawai‘i. Global Ecology and Biogeography Letters 7:
61–71.
Allen, J.A. 2002. Rhizophora mangle. In: Vozzo, J.A. (ed.).
Tropical Tree Seed Manual. USDA Forest Service, Wash-
ington, D.C.
Bandaranayake, W.M. 1998. Traditional and medicinal uses
of mangroves. Mangroves and Salt Marshes 2: 133–148.
Bandaranayake, W.M. 1999. Economic, traditional and me-
dicinal uses of mangroves. Australian Institute of Marine
Science, AIMS Report #28, Townsville, Australia.
Bosire, J.O., J. Kazungu, N. Koedam, and F. Dahdouh-Gue-
bas. 2005. Predation on propagules regulates regeneration
in a high-density reforested mangrove plantation. Marine
Ecology Progress Series 299: 149–155.
Brook, B.M. 2001. Te efect of Coccotrypes fallax (Coleop-
tera; Scolytidae) on the recruitment of Rhizophora stylosa
(Family Rhizophoraceae) in North Queensland man-
groves. PhD Tesis. School of Biological Sciences, James
Cook University, Townsville, Queensland.
Chong, P.W. 1988. Forest management plan for Playa Gar-
za pilot area: Terraba-Sierpe Mangrove Reserve, Costa
Rica. Food and Agriculture Organization of the United
Nations (FAO), Paris.
Cooke, G.P. 1917. Letter appended to paper by V. Mac-
Caughey. Hawaiian Forester and Agriculturalist 14: 365–
366.
Cox, E.F., and J.A. Allen. 1999. Stand structure and pro-
ductivity of the introduced Rhizophora mangle in Hawaii.
Estuaries 22(2A): 276–284.
Cuong, H.V.V. 1965. Rhizophoraceae. Nat. Mus. Nat. Hist.
4: 131–187.
Dahdouh-Guebas, F., S. Hettiarachchi, D. Lo Seen, O.
Batelaan, S. Sooriyarachchi, L.P. Jayatissa, and N. Koedam.
2005. Transitions in ancient inland freshwater resource
management in Sri Lanka afect biota and human popu-
lations in and around coastal lagoons. Current Biology
15(6): 579–586.
Dahdouh-Guebas, F., L.P. Jayatissa, D.D. Nitto, J.O. Bo-
sire, D.L. Seen, and N. Koedam. 2005. How efective were
mangroves as a defence against the recent tsunami? Cur-
rent Science 15: 443–447.
Dahdouh-Guebas, F., C. Mathenge, J.G. Kairo, and N.
Koedam. 2000. Utilization of mangrove wood products
around Mida Creek (Kenya) amongst subsistence and
commercial users. Economic Botany 54(4): 513–527.
Dahdouh-Guebas, F., M. Verneirt, J.F. Tack, and N. Koedam.
1997. Food preferences of Neosarmatium meinerti de Man
(Decapoda : Sesarminae) and its possible efect on the re-
generation of mangroves. Hydrobiologia 347: 83–89.
Dahdouh-Guebas, F., M. Verneirt, J.F. Tack, D. Van Spey-
broeck, and N. Koedam. 1998. Propagule predators in Ke-
nyan mangroves and their possible efect on regeneration.
Marine and Freshwater Research 49(4): 345–350.
De Lacerda, L.D. 2002. Mangrove Ecosystems: Function
and Management. Springer-Verlag, Berlin, Germany.
Ding, H. 1960. A review of the genus Rhizophora—with
special reference to the Pacifc species. Blumea 10(2):
625–634.
Duke, J.A. n.d. Rhizophora mucronata Lam. In: Handbook
of Energy Crops. <http://www.hort.purdue.edu/newcrop/
duke_energy/Rhizophora_mangle.html>.
Duke, N.C. 1992. Mangrove foristics and biogeography. In:
A.I. Robertson and D.M. Alongi (eds.). Tropical Man-
grove Ecosystems. American Geophysical Union, Wash-
ington, D.C.
Duke, N.C. 1995. Genetic diversity, distributional barriers
and rafting continents—more thoughts on the evolution
of mangroves. Hydrobiologia 295: 167–181.
Duke, N.C. 1996. Mangrove reforestation in Panama: an
evaluation of planting in areas deforested by a large oil
spill. In: Field, C. (ed.). Restoration of Mangrove Eco-
systems. International Society for Mangrove Ecosystems
ISME and International Tropical Timber Organization
(ITTO), Okinawa, Japan.
 Species Prohles for Pacihc Island Agroforestry (www.traditionaltree.org)  17
Duke, N.C. 2001. Gap creation and regenerative processes
driving diversity and structure of mangrove ecosystems.
Wetlands Ecology and Management 9: 257–269.
Duke, N.C., and Z.S. Pinzón. 1992. Aging Rhizophora
seedlings from leaf scar nodes: a technique for studying
recruitment and growth in mangrove forests. Biotropica
24(2a): 173–186.
Duke, N.C., M.C. Ball, and J.C. Ellison. 1998. Factors
infuencing biodiversity and distributional gradients in
mangroves. Global Ecology and Biogeography Letters 7:
27–47.
Duke, N.C., E.Y.Y. Lo, and M. Sun. 2002. Global distribu-
tion and genetic discontinuities of mangroves—emerging
patterns in the evolution of Rhizophora. Trees Structure
and Function 16: 65–79.
Duke, N.C., Z.S. Pinzón, and M.C. Prada. 1997. Large scale
damage to mangrove forests following two large oil spills
in Panama. Biotropica 29: 2–14.
Ellison, J.C. 1999. Present status of Pacifc Island mangroves.
Marine/Coastal Biodiversity Vol. II: 3–19. In: L.G. El-
dredge, J.E. Maragos, and P.L. Holthus (eds.). Te Tropi-
cal Island Pacifc Region: Population, Development and
Conservation Priorities. Pacifc Science Association/East
West Center, Honolulu.
Field, C.D. (ed.). 1996. Restoration of mangrove ecosystems.
International Society for Mangrove Ecosystems (ISME)
and the International Tropical Timber Organisation
(ITTO), Okinawa, Japan.
☛ Hogarth, P.J. 1999. Te Biology of Mangroves. Oxford
University Press, New York.
☛ Jimenez, J.A. 1999. Rhizophora mangle L. Report SO-
ITF-SM-2. Institute of Tropical Forestry, Rio Piedras,
Puerto Rico.
Kairo, J.G., F. Dahdouh-Guebas, J. Bosire, and N. Koedam.
2001. Restoration and management of mangrove sys-
tems—A lesson for and from the East African region.
South African Journal of Botany 67: 383–389.
La-Rue, C.D., and T.J. Muzik 1951. Does the mangrove re-
ally plant its seedlings? Science 114: 661–662.
Lo, E., N.C. Duke, and M. Sun. 2002. Phylogenetic evalu-
ation of Rhizophora taxa for conservation management.
Society for Conservation Biology, United Kingdom.
Marco, H.F. 1935. Systematic anatomy of the woods of the
Rhizophoraceae. Tropical Woods 44: 1–20.
Morton, J.F. 1965. Can the red mangrove provide food, feed
and fertilizer? Economic Botany 19: 113–123.
Riley, R.W., and C.P. Salgado Kent. 1999. Riley encased
methodology: principles and processes of mangrove habi-
tat creation and restoration. Mangroves and Salt Marshes
3: 207–213.
☛ Robertson, A.I., and D.M. Alongi (eds.), Tropical Man-
grove Ecosystems. American Geophysical Union, Wash-
ington, D.C.
☛ Saenger, P. 2002. Mangrove Ecology, Silviculture and
Conservation. Kluwer Academic Publishers, Boston.
Salvoza, F. M. 1936. Rhizophora. Univ. Philip. Nat. Appl. Sc.
Bull. 5(3): 179–237.
Smith, A.C. 1981. Family 96. Rhizophoracaeae. In: A.C.
Smith (ed.). Flora Vitiensis Nova: A New Flora of Fiji,
Vol. 2. Pacifc Tropical Botanical Garden, Lāwa‘i, Kaua‘i,
Hawai‘i.
☛ Snedaker, S.C., and M.S. Brown. 1982. Primary produc-
tivity of mangroves. In: Mitsui, A. and C.C. Black (eds.).
CRC Handbook of Biosolar Resources. CRC Press Inc.,
Boca Raton, Florida.
Sousa, W.P., S.P. Quek, and B.J. Mitchell. 2003. Regenera-
tion of Rhizophora mangle in a Caribbean mangrove for-
est: interacting efects of canopy disturbance and a stem-
boring beetle. Oecologia 137: 436–445.
Steenis, C.C.V. 1958. Introduction to Rhizophoraceae. Flo-
ra Mal. (1)54: 429–444.
Taman, R.R. 1993. One hundred Pacifc island agroforestry
species. In: Clarke, W.C. and R.R. Taman. Agroforestry
in the Pacifc Islands: Systems for Sustainability. United
Nations University Press, Tokyo. <http://www.unu.edu/
unupress/unupbooks/80824e/80824E0p.htm>.
☛ Tomlinson, P.B. 1986. Te Botany of Mangroves. Cam-
bridge, Cambridge University Press.
Tsuda, S., and M. Ajima. 1999. A preliminary study of re-
sprouting ability of some mangrove species after cutting.
Tropics 8(3): 221–224.
USDA Forest Service. n.d. Wood Technical Fact Sheet.
Rhizophora mangle. Adapted from: Chudnof, M. 1984.
Tropical Timbers of the World. Agriculture Handbook
607. USDA Forest Service, Forest Products Laboratory,
Madison, Wisconsin. <http://www2.fpl.fs.fed.us/Tech-
Sheets/Chudnof/TropAmerican/htmlDocs_tropameri-
can/Rhizophoramangle.html>.
Walsh, G.E. 1977. Exploitation of mangal. In: V.J. Chapman
(ed.). Ecosystems of the World: Wet Coastal Ecosystems.
Elsevier Science Publishing Company, Amsterdam.
Walters, B.B. 2005. Patterns of local wood use and cutting
of Philippine mangrove forests. Economic Botany 59(1):
66–76.
Weinstock, J.A. 1994. Rhizophora mangrove agroforestry.
Economic Botany 48(2): 210–213.
Wester, L. 1981. Introduction and spread of mangroves in
the Hawaiian Islands. Association of Pacifc Coast Geog-
raphers Yearbook 43: 125–137.
Wilkinson, K.M., and C.R. Elevitch. 2003. Propagation
protocol for production of container Rhizophora mangle
L. var. samoensis Hochr. plants. In: Native Plant Network.
University of Idaho, College of Natural Resources, Forest
Research Nursery, Moscow, Idaho. <http://www.native-
plantnetwork.org>.
18  Atlantic–East Pacifc Rhizophora species 
Species Profles for Pacifc Island Agroforestry (www.traditionaltree.org)
Rhizophora mangle, R. samoensis, R. racemosa, R. × harrisonii
(Atlantic–East Pacifc red mangroves)
Authors: Norman C. Duke
1
and James A. Allen
2
1. Marine Botany Group, Centre for Marine Studies, Te University of Queensland, St. Lucia QLD 4072, Australia;
<http://www.marine.uq.edu.au/marbot/index.htm>.
2. Paul Smith’s College, PO Box 265, Paul Smiths, New York 12970, USA; Web: http://www.paulsmiths.edu
Acknowledgments: Te authors and publisher thank Tom Cole, Farid Dahdouh-Guebas, Dale Evans, Kathy Ewel, Ariel Lugo, and
Diane Ragone for their input. A photo contribution by Ulf Mehlig is greatly appreciated. Norm Duke acknowledges a host of
people who have over the years supported his journey into Rhizophora and who share the passion for this group of plants. Collabora-
tors and friends include: Jim Allen, Uta Berger, John Bunt, Molly Crawford, Farid Dahdouh-Guebas, Otto Dalhaus, Kathy Ewel,
Candy Feller, XueJun Ge, Gina Holguin, L.P. Jayatissa, Kandasamy Kathiresan, Ken Krauss, Eugenia Lo, Ulf Mehlig, Jaime Polania,
Jurgenne Primavera, Ute Steinicke, Mei Sun, and Nick Wilson.
Recommended citation: Duke, N.C., and J.A. Allen. 2006. Rhizophora mangle, R. samoensis, R. racemosa, R. × harrisonii (Atlantic–
East Pacifc red mangroves), ver. 2.1. In: Elevitch, C.R. (ed.). Species Profles for Pacifc Island Agroforestry. Permanent Agriculture
Resources (PAR), Hōlualoa, Hawai‘i. <http://www.traditionaltree.org>.
Sponsors: Publication was made possible by generous support of the United States Department of Agriculture Western Region Sus-
tainable Agriculture Research and Education (USDA-WSARE) Program; SPC/GTZ Pacifc-German Regional Forestry Project;
USDA Natural Resources Conservation Service (USDA NRCS); State of Hawai‘i Department of Land & Natural Resources Divi-
sion of Forestry & Wildlife; and the USDA Forest Service Forest Lands Enhancement Program. Tis material is based upon work
supported by the Cooperative State Research, Education, and Extension Service, U.S. Department of Agriculture, and Agricultural
Experiment Station, Utah State University, under Cooperative Agreement 2002-47001-01327.
Series editor: Craig R. Elevitch
Publisher: Permanent Agriculture Resources (PAR), PO Box 428, Hōlualoa, Hawai‘i 96725, USA; Tel: 808-324-4427; Fax: 808-324-
4129; E-mail: par@agroforestry.net; Web: <http://www.agroforestry.net>. Tis institution is an equal opportunity provider.
Reproduction: Copies of this publication can be downloaded from <http://www.traditionaltree.org>. Tis publication may be repro-
duced for noncommercial educational purposes only, with credit given to the source. © 2006 Permanent Agriculture Resources. All
rights reserved.

since which time their presence has become quite noticeable. no mangroves were present until introductions were made in the 1920s. mostly trees.). In the Hawaiian Islands. where introductions of R. R. As such. 1998). Specialized morphological and physiological characteristics largely define and characterize mangrove plants. These halophytic (salt tolerant) plants thrive in saline conditions and daily inundation between mean sea level and highest astronomical tides. samoensis but occurs as a distinct co-inhabitant of man-   Atlantic–East Pacific Rhizophora species  . The status of observed differences in like forms R. The closely related and almost identical “sibling species. not withstanding the disjoint but natural occurrence of R. mangroves have reportedly overgrown channels. reduced tidal flows. In Hawai‘i. This species group is one of two that make up the genus Rhizophora and consists of three species (two being closely allied) and one hybrid: R. mangle and R. soil types. including the IWP stilt mangroves. while flooding tides allow uptake of river-borne nutrients and frequent dispersal by their buoyant propagules. New Hebrides. Mangrove plants exchange gases from exposed roots using special lenticels. Duke et al.. mangle. respectively. will only be resolved in genetic studies and selected breeding programs. including high salinity levels from greater than full strength seawater to freshwater (Cintron et al. enhancement of water quality in near-shore environments (plus coral reef areas). and Samoa. mangle. such as buttress trunks and roots providing support in soft sediments. mangle have been made. the American Pacific coast. samoensis in New Caledonia. The taxonomic status of these sibling taxa is arguable based on morphological data alone.g. for example. Rhizophora racemosa is less common than R. and support of estuarine and marine food chains. as well as in isolated occurrences in the southwestern Pacific islands. Fiji. including reduction in the habitat quality for endangered water birds such as the Hawaiian stilt (Himantopus mexicanus knudensi).INTRODUCTION Atlantic–East Pacific red mangroves (AEP Rhizophora species) are the most important and dominant mangrove species of tropical coastal areas of the Atlantic Ocean. Many of the values of red mangroves are difficult to separate from the larger role played by mangrove plants. 1978). Native range Red mangroves are generally considered non-native to the IWP. New Hebrides. colonization of habitats to the detriment of native species (e. and causing localized drainage problems by reducing the flow through tidal creeks or drainage channels. Tonga. The ecological limits defined by the diurnal tidal range explain the setting and why just 70 species around the world are considered to be mangroves (Tomlinson 1986. Typically. DISTRIBUTION Atlantic–East Pacific red mangroves are native to tropical regions along the American east and west coasts to the African west coast. aboveground roots allowing vital gas exchange in anaerobic sediments. and other physical site factors. in anchialine pools). overgrowing native Hawaiian archaeological sites. notably New Caledonia. compared with adjacent rainforests that may have hundreds of tree species per hectare. and they provide vital structure as habitat and food for similarly adapted resident and transient fauna. samoensis. Fewer than 22 plant families have developed such essential attributes. R. Red mangroves. is native to the American west coast and islands in the southwestern Pacific. representing independent instances of co-evolution over millions of years to form today’s mangrove habitats. and they tolerate a range of flooding. and Samoa. × harrisonii. racemosa. predominantly bordering margins of tropical coastlines around the world. have also been introduced into new sites in the Indo–West Pacific (IWP) region during the past century. several important negative effects have been documented. Fiji. However. notably R. and R. the uses of such plants need to be weighed carefully against their effects as potentially invasive species. samoensis (=R. Rhizophora mangle is the species that most characterizes AEP Rhizophora. they are most common in the middle to low intertidal zone above mean sea level. =R.” R. mangle sensu lato Tomlinson 1986. It occurs naturally and dominates tropical tidal areas along both sides of the Atlantic. Red mangroves thrive under a range of intertidal wetland conditions. and overgrown archeological sites. samoensis. the roles of particular species of Rhizophora are often not distinguished from other members of the genus. Because Rhizophora species dominate most tropical mangroves worldwide they are generally believed to play a vital role for mangrove ecosystems including shoreline protection. and several islands in the southwestern Pacific Ocean. samoensis Hochr. WHAT IS A MANGROVE? Mangroves form a unique and dominant ecosystem comprised of intertidal marine plants. mangle and R. and physiological adaptations for excluding or expelling salt. extending often along the seaward margin of mangrove stands. mangle var. Tonga. however. In the Hawaiian and Society Islands.

In the absence of human intervention. image: N. mangle and R. Duke grove stands in wetter areas and larger catchment estuaries of the Atlantic. R. This hybrid taxon is special because R. the AEP Rhizophora species. there are tantalizing similarities in cross-ocean linkages for several shallow-water reef fish. stylosa (=R. Plants introduced in the early 1900s to Moloka‘i and O‘ahu now extend to most islands of the group. At least one other possible species. and accepting that long-distance dispersal was not possible. However. with distances between the putative ancient islands large enough to restrict all mangrove species except the dispersal-specialist red mangroves. The species favors riverine estuaries. mangle var. mangle/R. Also occurring in this region is another hybrid taxon.traditionaltree. In each case. stylosa is a dominant member of the IWP stilt mangroves.World distribution of red mangroves. racemosa and the putative hybrid R. racemosa and R. Of great interest also is the natural presence of red mangrove in the southwestern Pacific islands. one explanation is island-hopping by natural dispersion across an ancient archipelago of seamounts that crossed the southeastern Pacific during early formation of the Pacific Plate in the Late Cretaceous period (Duke 1992. samoensis in the southwestern Pacific. Rhizophora mangle has been introduced to the Hawaiian and Society Islands from Atlantic populations in Florida. mucronata var. stylosa).C. × harrisonii. its distribution is likely where the distributions R. racemosa. In support of this theory. occurs across the same range. R. × harrisonii appears restricted mostly to equatorial estuaries of larger river systems with more continuous freshwater flows. founder populations have increased and expanded dramatically. The distribution of R. and the expecta- producers. Any migration would have been westward. there is no evidence for this happening. it is feasible that their dispersal westward in the Pacific may have been assisted by aboriginal travelers in the past. samoensis overlap. Given this and its intermediate characters. the taxon is considered the putative hybrid of R. × harrisonii. The occurrence of this hybrid means that there appears to be very little genetic separation between these defining and most divergent of Rhizophora species. beneficial to shoreline stabilization and fisheries. 1995). samoensis) and R. samoensis (=R.org)   . Coastlines marked in red indicate the distribution of all mangroves. selala (Salvoza) Tomlinson. Given the apparent hybrid status of R. whose putative parents include R. Because red mangroves are recognized as valuable timber Current distribution Red mangroves extend notably beyond their native range. so currently there is no agreed explanation for the presence of R.  Species Profiles for Pacific Island Agroforestry (www. especially in the Hawaiian Islands. and it is restricted in the AEP to a few stands along the Pacific coast of the Americas and does not occur in the IWP.

Petals. mature buds and flowers are located at 1–2. racemosa. Open flowers are located within or below leaf axils at leaf nodes below the apical shoot. mangle var. × harrisonii. American mangrove (English) red mangrove (USA) mangle rojo (Central and Latin America. R. R. single-stemmed trees are often found just behind the seaward edge of stands downstream in major river estuaries. Leaf emergence occurs chiefly during November–February in the southern hemisphere and May– August in the north. bright green. Peduncle is 3–4 cm long (1.2 in) wide.06–0. Style is pale green. to golden brown at maturity.3. racemosa mature fruit located in leaf Flowering BOTANICAL DESCRIPTION Rhizophora mangle L. on the seaward edge of stands or in sites of higher salinity.6 in). containing both male and female parts. creamy white. Cork wart spots occur on most species’ leaf undersurfaces. Rhizophora samoensis (Hochr. margins revolute.) Rhizophora racemosa G. it was destroyed.F.5 in) long and ~4 mm (0. are lanceolate to linear.5–7 m (1. respectively. dichotomous tip.Similar introductions of R.2. October–February in the southern hemisphere and April–August in the northern hemisphere. Inflorescences have few to many joints with 1. Buds elongate to ovate. mangle. mangle.02–0.3 cm (0. For R. obovate. racemosa in northern Brazil.01 in) wide.2–1. Prop roots are sturdy even when relatively thin. × harrisonii.W. Taller. Note that nonspotted leaves are found on an unusual form of R. depending on species. Multi-stemmed trees occur in frontal areas but are more common in upper intertidal regions. with woolly to sparsely hairy margins.12 in) wide. Flowers are perfect. area. They may reach 30–50 m (100–160 ft) in height. R. and R. reddish brown (may be distinguished from infections and wounds by their uniform cover). obtuse blunt apex with a minute lip folded under. not raised. In the Townsville. Leaf fall occurs chiefly over the wet summer period. as diameter height above the substrate varies from 0. calyx lobes elongate spreading (when the hypocotyl is ready to emerge). ~0. mangle × R. Meyer Rhizophora × harrisonii Leechman (=R. are pear-like. although they are commonly much shorter. simple. elongate. waist constriction. filiform. dimensions 1–2 cm (0.5–3 mm (0. Bracts and bracteoles are distinct. usually four. racemosa) Rhizophoraceae (mangrove family) Atlantic–East Pacific red mangroves. 3–5. The calyx is typically waxy yellow to creamy white and green at maturity. Red mangroves are often multi-stemmed rambling to columnar trees with distinct aboveground prop roots. Australia.4–0.16 in) above ovary base. Accordingly. and R.8 in) long.4–chotomous branching and one to many buds possible per inflorescence. ~0. scattered evenly. tion is that they will spread further. with four lobes. leathery. mangle is treated as an invasive species in these islands. terete. local authorities were not prepared to take the risk. so when a small introduced stand of R. Fruits. pale yellow. Pacific and Caribbean coasts) tiri wai (Fiji) togo (Samoa) Atlantic–East Pacific red mangroves are medium to tall trees. around 5–8 m (16–26 ft). when mature. mangle elsewhere are apparently less invasive. ~12 mm (0. smooth brown surface. mangle was discovered in the upper part of Ross River. green when immature to lighter colors as they mature. 0. and 7–9 nodes down from the apical shoot. For R. Leaves are opposite. smooth. 1. samoensis Hochr.5–4 mm (0. pale yellow. Stamens number eight.16 in) wide.5 cm (0. however.) Salvoza (=R. possibly because other mangroves that already occupy the habitat reduce establishment opportunities for any new introductions. Flowering period is chiefly (but not exclusively) August–December in the southern hemisphere and February–June in the northern hemisphere. This method of measurement differs fundamentally from the standard diameter at breast height (dbh) used for most forest surveys. generally curved surface. Stem diameters are about 15–35 cm (6–14 in) taken just above the highest prop root. They tend to be of shorter stature and more spreading in shape Preferred scientific names Family Common names Leaves Size Form Fruit   Atlantic–East Pacific Rhizophora species  .5–23 ft) (consider the tree in the photo on the first page).

Top right: Leafy rosette of Rhizophora mangle (similar to R. Rhizophora mangle and R.org)   . mature hypocotyls are located in leaf axils 3–8. Note the persistent style at the distil tip of the fruiting body. a viviparous seedling. green.  Species Profiles for Pacific Island Agroforestry (www. For R. R. Duke axils 3. photos: N. reflexed calyx lobes. respectively. red mangroves are viviparous.C. prior to emergence of the hypocotyl. samoensis) showing open flowers with elongate. occurs chiefly (but not exclusively) in November–January in the southern hemisphere and May–July in the northern hemisphere. to gray-fissured with smooth.traditionaltree. distal half wider. There is total coverage of gray-fissured bark in some localities. Bark Hypocotyls are narrowly cylindrical.” when mature hypocotyls fall. samoensis both have distinct brown distal portions of other- Bark is smooth and red-brown in seaward and exposed locations (rocky and sandy substrates). 7 (rare) and 10–11 nodes down from apical shoot. distal tip pointed. smooth with irregular small brown lenticels. red-brown prop roots in sheltered locations (mud substrates). The dispersal unit. elongate. wise green. as in this R. Like all Rhizophora species. and R. racemosa. meaning that they produce seeds hidden in the mature fruit. ~8 (rarely observed). × harrisonii. and these germinate while the fruit is still attached to the parent plant. Atlantic Panama. is called a hypocotyl. and 13–15 nodes down from the apical shoot. non-reflexed calyx lobes. slightly hairy petals and stiff-erect. relatively short hypocotyls (see photo). respectively. mangle. Bottom left: Open flower of Rhizophora racemosa showing flat. One hypocotyl is usually produced from each fruit.Top left: The blunt apex of leaves is characteristic of all AEP red mangroves. Bottom right: Mature fruit of Rhizophora mangle (similar to R. Seeds/hypocotyls “Fruiting. samoensis from Fiji. samoensis) showing its elongate pear-shape. although on rare occasions twins may be observed.

samoensis has bracts twice as wide as the pedicle. down from the apical shoot. samoensis is perhaps the most contentious. aboveground prop roots surrounding the stem base that anchor only shallowly in the sediments to 1–2 m (3. as well as their distinct geographic ranges. Hawai‘i. are distinguished by minute/absent bracts on pedicles at the calyx base in R. which may.6 m) depth. which have 3–8 inflorescence joints. racemosa and R. samoensis.C. mangle and R. as they are distinguished by only a few morphological characters. Duke GENETICS Variability of species Atlantic–East Pacific red mangroves are those Rhizophora species that occur naturally along the east and west coasts of the Americas. be shown to be the same species. The sibling Expended fruits of Rhizophora samoensis (left) and R. Rhizophora mangle and R. They appear very closely related. samoensis. samoensis. Discriminating between them reliably is not possible in many instances without detailed examination of key morphological and genetic characteristics. mangle and R. sturdy. R.C. The possible hybrid character is shown in R. Red mangroves are distinguished from IWP stilt mangroves principally by the leaf apex: red mangroves have a blunt leaf apex and lack the spiked. where it has characters intermediate between R. as well as along the west African coast and those island populations in the southwestern Pacific. × harrisonii. mucronate leaf tip present in stilt mangroves. racemosa and R.Rooting habit Mature trees have distinctive. mangle. although at times their morphological and taxonomic differences are questionable. samoensis appear very closely related and are likely to be the same species. × harrisonii. × harrisonii. are distinguished from R. mangle. Duke   Atlantic–East Pacific Rhizophora species  . on closer examination. The uncertainty is chiefly based on: 1) the presence of one intermediate individual that is recognized as a distinct hybrid. Rhizophora racemosa is distinguished from R.3–6. As noted. Rhizophora mangle and R. mangle and R. This conforms to the anoxic conditions commonly observed in mangrove sediments. Similar species Mature hypocotyls of Rhizophora mangle (similar to R. showing distinctive brown distal end. the sibling species. photo: N. stylosa (right). Fiji. R. while R. samoensis). namely R. In this group there appear to be four relatively distinct taxa. photo: N. and 2) the occurrence of two sibling species. × harrisonii by the node position of mature buds and flowers in leaf axils. being 7–9 nodes and 3–5 respectively. The relationship of R. with mostly 0–3 inflorescence joints.

racemosa based on its intermediate and shared morphological characteristics. All Rhizophora species are closely similar in tree form. The same recent investigation also discovered two intermediate forms. are considered close in form and value also. One fits the type description in every respect including the presence of cork wart spots on leaf undersurfaces. Rhizophora harrisonii is the apparent hybrid of R. while R. pers. stylosa (a stilt mangrove). The question raised by this natural discontinuity remains unresolved. mangle and R.C. samoensis where they occur elsewhere in the Indo–West Pacific. samoensis in the southwestern Pacific islands. and the species now dominates most intertidal wetland areas of each location. mangle in the Atlantic and R. mangle in the Hawaiian and Society Islands. and cultural groups in the Pacific region may not always distinguish between them. racemosa were found to co-exist in northern Brazil.org)   . racemosa forms and hybrid grow in proximity to a regular freshwater source. and how do these compare with populations in western Africa? There are clearly more questions than answers concerning genetic variation in red mangroves. mangle. This species appears more common in estuaries influenced by larger and more continuous freshwater flows in equatorial regions of three distinct regions: the east Pacific (Costa Rica to Ecuador). This observation emphasizes the profound limitations of long-distance dispersal for this mangrove group of otherwise long-distance dispersal specialists. The Australian introduction was eradicated a few years ago. with R. By contrast. In Hawai‘i changes in shoreline ecology have been dramatic. samoensis in the Pacific. preferably within intertidal wetland conditions above mean sea level elevations in tropical regions. mangle and R. Further investigations are needed to adequately describe Rhizophora taxa and their distributions throughout the AEP region. samoensis. Aberrant individuals are reported to produce chlorophylldeficient propagules. Curiously. Current evidence indicates the situation may be more complex. This is the only example where AEP and IWP Rhizophora species naturally co-exist.traditionaltree. mangle and R. samoensis. while the other form lacked these spots. mangle. Key questions arise from these new observations including: what is the distribution of the two forms of R. but in both other locations the spread of R. and these together are often used in similar ways by indigenous peoples. R. R. but no detailed reports are available. while R. Known varieties and hybrids Recognizing the morphological differences between R. This was particularly notable since all red mangroves until now have been reported to possess such spots on their leaf undersurfaces. mangle demonstrate that island habitats without mangroves existed between Samoa and South America. In this case. racemosa is readily distinguished from R. Their relative numbers on an individual tree are believed to quantify the amount of outcrossing that occurs among neighboring normal trees. two forms of R. Yellow or red propagules can be observed hanging in affected trees alongside normal green propagules. especially because no mangroves grew in this isolated location previously. mangle stands throughout the Caribbean area to northern Brazil. with another possible hybrid where R. the human introductions of R. mangle (mostly from around Atlantic Florida) was introduced to the Pacific in several isolated instances. called albinos. obs. A similar situation is expected in the Society Islands. frequent freshwater runoff. Duke. In Fiji. Other trees have also been observed with variegated foliage. notably to Hawai‘i. however. mangle and R. and that these are suitable for mangrove colonization. samoensis is considered useful in distinguishing between introduced populations of R. Furthermore. apiculata occurring in Australasia and the northwestern Pacific. R. samoensis occurs more fre-  Species Profiles for Pacific Island Agroforestry (www. There are no reports of differences within this taxon between these areas. racemosa were observed in Brazil. racemosa are commonly found in lower to middle tidal reaches of rivers and streams with regular. racemosa apparently occurs on the eastern Pacific coast of the Americas with R. The situation became more complex during the 1900s when R. such as Bruguiera. and east Atlantic (western Africa). samoensis grow in marine saline conditions. two forms of R. samoensis occurs together with R.species separate naturally across the American land barrier. racemosa forms and R. recognized as potential hybrids between the two R. the Society Islands. as compared with natural populations of R. samoensis occur in coastal embayments and coral islands. racemosa. R. and northeastern Australia. mangle and R. mangle has proceeded unchecked. This character is present in IWP stilt mangroves and in a similar way defines the two forms of R. This is particularly notable and common in R. These introductions highlight the unexplained discontinuity of R. the allied partner to R. Other mangrove genera.). west Atlantic (Venezuela to Brazil). Culturally important related species ASSOCIATED PLANT SPECIES AEP red mangroves can be found growing with other mangrove species. as noted above. but recently (N. R.

and clays). Associated species commonly found Soils ENVIRONMENTAL PREFERENCES AND TOLERANCES Atlantic–East Pacific red mangroves thrive in tropical and subtropical environments characterized by moderately high and well distributed rainfall. They are also anaerobic with high concentrations of sulfide. characterized by fine clay. The optimal salinity range is reported to be from 8 to 26 ppt (parts per thousand). In Hawai‘i. presumably due to decreases in freshwater availability. red mangroves occur on estuarine margins just above mean sea level. especially during drier months. Dry season duration (consecutive months with <0  mm [1.   Atlantic–East Pacific Rhizophora species  . and coral ramparts. medium. in] rainfall)  Because of access to groundwater.5 Special soil tolerances  Plants grow best in saline soils but survive well in fresh water. samoensis (=R. Soil drainage The trees grow in soils with free and unimpeded drainage. Soil acidity  pH 6–8. stylosa and Bruguiera gymnorrhiza predominantly. Mean annual rainfall  These mangroves grow in all rainfall conditions. including R. as well as waterlogged soils. The types of associated species are notably variable where this species has been introduced to the IWP region. Across a wide variety of climatic regions. red mangroves grow in regions with up to 8 months of drought. Best development has been observed in lower tidal reaches and on soft. sandy loams. Rainfall pattern  They grow in climates with summer or uniform rainfall patterns. R. Mean annual temperature  20–30°C (68–86°F) (estimate) Mean maximum temperature of hottest month  32–38°C (90–100°F) (estimate) Mean minimum temperature of coldest month  0–5°C (0–41°F) Minimum temperature tolerated  0°C (32°F) (estimate) Trees develop greatest stature and columnar growth form in estuaries of larger tropical rivers. Their extent. Climate Soil texture  Plants grow in light. red mangroves occur commonly with Avicennia germinans and Laguncularia racemosa. sandy clays. In Atlantic and east Pacific populations. in reference to mean sea level. such as the Baja Peninsula on the northern Pacific coast of Mexico. while the species frequently occurs in nearly mono-specific stands. while R. Trees also grow well in sites with aerobic sediments consisting of fine sands to coarse stones and rocks. preferring downstream and exposed coastal locations. and heavy texture soils (sands. the stunted but dense thickets of R. This has been evident in correlations between El Niño events and reduced growth as possible causes of some damage to mangroves. clay loams. it does sometimes occur alongside a mangrove introduced from the IWP (Bruguiera sexangula) and species that typically occur in lower-salinity environments. loams. mangrove cover expands and contracts through time. In the southwestern Pacific. such as Hibiscus tiliaceus. In other locations. compared with approximately 34–36 ppt for seawater. Mangrove plants appear to depend on groundwater to sustain optimal growth. Elevation range  0–6 m (0–20 ft). mangle?) attest to the great adaptability of red mangroves to a wide range of climatic types. form. muddy sediment substrates. and sandy clay loams. black mud sediments with relatively high loads of organic carbon.Red mangroves often extend up the tidal profile to the terrestrial fringe in areas of higher rainfall. In drier locations. fine. samoensis is associated with IWP mangroves. red mangroves are associated with other mangrove species. quently in upstream locations of the freshwater dominated streams. stylosa remains the marine specialist. In lower-rainfall areas. and biomass reflect the different rainfall conditions.

Shade They appear to have low tolerance of shade. Since mangroves have narrow optimal temperature ranges. Fire Red mangroves have no tolerance of fire in close proximity. such changes have occurred throughout history. as expected with changes in longer-term rainfall patterns as this affects moisture stress in saline environments. Waterlogging Red mangroves are tolerant of daily tidal flooding up to depths of up to 1. rainfall. Full sun They grow best in full sun. rising temperatures will cause their distributions to shift north or south to areas where temperature conditions are most suitable.5 m (5 ft). Sousa et al. Changes along contours can be quantified from longterm spatial assessments over decade. 2003). Generally. red mangroves are believed to provide significant buffering of coastal areas during storm and tsunami surge events (Dahdouh-Guebas 2005). and the availability of suitable new space. so the distribution of mangroves today represents the survivors of all past changes. but they Coppice The trees have notably poor coppice ability. depending on the rates of change. These might be observed as shifts in vegetation. and they will die off in areas where they are not suited. as sea level rises their communities must move upland to survive. Abilities Self-prune Red mangroves self-prune well in dense stands. Of course. They generally grow less than  Species Profiles for Pacific Island Agroforestry (www. GROWTH AND DEVELOPMENT Growth rates vary with age. and sea level. like other mangrove species. However. While tolerating permanently saturated soils. it will die. shaded seedlings (Brook 2001. Key indicators of change can be identified and mapped as incremental shifts and responses of mangrove communities. MANGROVES AND CLIMATE CHANGE Atlantic–East Pacific red mangroves.and centurylong time periods. The unique physiological characteristics of each species define its capacity for survival in the face of change. the response zones will follow elevation contours. Frost Tolerance of sub-freezing temperatures is low to none.Tolerances Drought Red mangroves can tolerate drought periods well. commonly maintain lower branches in more open-growing locations. Wind Red mangroves are typically found in seaward areas subject to wind and salt spray but largely protected from waves. In areas where wave action is infrequent. In both cases.org)   . are affected by climate change. recent evidence shows this more likely due to weevils that infest and kill cooler. Knowledge of these changes and their causes allows better prediction of future change. Drought conditions presumably cause soil salinity to increase in excess of tolerable limits for these plants. because mangroves are characteristically restricted to elevations between mean sea level and highest tides.traditionaltree. For instance. although trees have apparently been killed by drought in some sites on Moloka‘i. but only where required and within the tree’s native range. Clearly. if 50% or more of the leaves are removed from a tree. Mangroves are expected to respond rapidly and decisively to shifts in key factors. their success in making these shifts depends on their successful dispersal and re-establishment. and 2) in the salt marsh–mangrove ecotone. Planting in such highly wind-prone locations is recommended. Hawai‘i where groundwater sources appear to be limited. Salt spray The trees are highly tolerant of salt spray. they are intolerant of drying soils. Waves In general. like temperature. wave-prone coastlines are inhospitable to mangroves. as expected with changes in sea level. for example: 1) in the total tidal wetland habitat zone. as each species has defined ranges of tolerance for each factor. exposed.

spaced at about 1–1. Propagules can be sown in nursery beds. slower plants die and decompose quickly. Propagule maturation occurs later in relatively higher latitude locations north or south beyond the tropic zone (e. vigor. Trees have notable and relatively long periods of reproductive development. Low maintenance is generally sufficient for maximizing seedling establishment success in sheltered areas. leaving faster competitors the benefit of not only the space they occupied but also their nutrients. Although propagules with only minor borer damage may survive and grow. Jimenez (in Chong 1988) reported annual diameter growth increments for R. and neither nursery preparation nor stakes are needed. However. and 0. This affords them maximal protection from physical damage by drift logs and erosive waves. but should not be stored for long. these species are relatively easy to propagate.06–0. Propagules that already have some root or leaf development can be used in most cases. racemosa trees of 0. Propagule processing Processing of mature propagules is not required for red mangroves. PROPAGATION Although natural regeneration is generally relied upon 10  Atlantic–East Pacific Rhizophora species  . Only healthy looking propagules should be selected. The following pretreatment section gives possible additional steps. It is likely that propagules can be kept Rapid early growth of red mangrove seedlings in full sunlight ensures their success and dominance in preferred estuarine and intertidal conditions. Reaction to competition Propagule collection Propagules may be available throughout the year. selection of propagules with no signs of borer or crab damage are strongly preferred. Since competition is high between neighboring seedlings. Growth rate slows when trees approach a site-specific maximum canopy height. and age. Flowering and fruiting Flowering and fruiting periods of red mangroves are distinctly seasonal. The estimated mean annual increment (MAI) was 6 m3/ha/yr (86 ft3/ac/yr) with some stands as high as 11–14 m3/ha/yr (157–200 ft3/ac/yr). Peak pheno-events are expected to shift into later months with cooler temperatures and higher latitudes. Pretreatment is considered essential if considering such storage (see below).. racemosa with an expected period of around 30 months for each reproductive cycle. July–October in Hawai‘i). Mature propagules may be collected from the ground after they have fallen or picked directly from trees. Growth rates vary with species. spatial position in the stand. taking 18–30 months from first emergence of flower bud primordia until maturation and drop of mature hypocotyls. or installation of temporary structures to dampen wave action and reduce debris drift across restoration sites. When near maturity and maximum height. Yields/growth rates Chong (1988) provides information on potential yields from Playa Garza Pilot Area in an investigation of the feasibility of managing mangrove forests the Terraba-Sierpe Forest Reserve on the southern (wetter) Pacific coast of Costa Rica.03–0.1 m/yr (3. The duration depends on species. or preferably planted directly in the field soon after collection.g.17 cm (0.08–0. Propagation is simple and relies on the special feature of the genus in having large viviparous propagules.3–5 ft) intervals. but peak production occurs around July–August in northern parts of the range and around January–February in the southern hemisphere.19 cm (0. Newly established seedlings grow best in close proximity with their same species cohort. competition. Height growth is rapid shortly after establishment while food reserves are taken up from the hypocotyls of established seedlings. Such protection methods may include encasement of individual seedlings in PVC piping (Riley and Kent 1999). Propagule storage Propagules can be kept viable for at least 6–7 days by storing them in brackish water or by wrapping them in wet burlap bags and keeping them out of direct sunlight.5 m (3.3 ft/yr) but can exceed this in favorable circumstances.14–0. from Pacific to Atlantic regions.07 in) for diameter size classes greater than 10 cm. Planting simply entails gently pushing the distal end of the 20–60 cm (8–24 in) long hypocotyl one third of its length into the sediment. For northern Pacific sites in Costa Rica. Propagules that are shrunken or desiccated in appearance or that exhibit significant physical damage should be rejected. mangle and R. substantial protection is required in more exposed coastal locations during the first decade of establishment. with the longest being R. as observed in Vietnam and China (Field 1996).07 in) for diameter size classes less than 10 cm (4 in). trees broaden their canopies and increase stem diameter rather than grow taller. although damaged and insect-infested individuals should be removed. No holes need be dug.

survival may be zero on sites exposed to excessive wave action. The plants are readily opportunistic due to their relatively wide tolerance for salinity and Potential for invasiveness  Species Profiles for Pacific Island Agroforestry (www.org)  11 . sheep. Brazil. a study of scolytid beetle larvae (Coccotrypes fallax) infections on Rhizophora propagules in Australia found at least 18% were heavily infested (Brook 2001). Heat treatment might be easily achieved by leaving the collected propagules in the sun for a short period (a few hours) before planting. The study went further to find that pretreatment in a 50°C (122°F) water bath for 5 minutes killed the beetles and removed the risk of establishment losses due to borer damage. The very successful introduction and rapid spread of red mangroves in the Hawaiian Islands clearly demonstrates the potential for invasiveness of mangroves in areas where suitable habitat is available. On the other hand. Direct planting of large numbers of propagules is restricted by their peak seasonal availability. which can take up to 6 months to achieve.5 m (1. cattle. A method of encasing propagules in PVC pipe has been used in Florida and the Caribbean to protect seedlings in places with a high likelihood of disturbance. Propagule pretreatment Pretreatment of propagules is generally considered unnecessary. red mangroves pose few significant disadvantages when planted within their native range. photo: u. if appropriate. and long-term storage is not feasible.traditionaltree. Propagules or nursery-grown seedlings usually have excellent survival in sites correctly selected and. However. mehlig longer. Time to outplanting Seedlings are ready for outplanting at the six-leaf (threenode) stage if grown up in nursery conditions. horses). but storage beyond 2 weeks is not recommended.g. Media Although a wide variety of soil media are acceptable. DISADVANTAGES In general. as propagules do not keep for extended periods.5 ft) tall have also been successfully transplanted. but this is not recommended. smooth propagule. Seedling establishment Leaves and roots may begin to develop within a week or two of sowing. on sites with inappropriate hydrologic or salinity regimes. goats. Mature hypocotyl of Rhizophora racemosa showing distinctive. protected from disturbance. Older seedlings up to 0. a mix of sand and peat in equal parts has been recommended for mangrove nurseries.Growing area Red mangroves should be grown in full sunlight. They are not especially susceptible to pests or pathogens and have not been reported to host major pests or pathogens of important crop species. or (rarely) subjected to disturbance by grazing animals (e. Survival rates of 90% or greater are reasonable in such circumstances.. slender. Guidelines for outplanting Outplanting anytime of the year is suitable. Infested propagules were established under canopy-shaded areas.

Fence posts Red mangrove stems make good posts. Clearly. Bee forage Rhizophora species have no nectar. which dampen water movement and promote sedimentation in areas that might otherwise be eroded. Many fish species use red mangroves during part of their Pests and diseases Susceptibility to pests and diseases is believed to be low. it wasn’t the best choice for this purpose! Fish/marine food chain Mangroves in general are believed to play a vitally important role in protecting and supporting marine food chains. Soil stabilization Red mangrove forests stabilize soils with their network of sturdy overlapping prop roots. particularly on islands with clear tenure systems that include mangrove areas. however. 1  Atlantic–East Pacific Rhizophora species  . Rhizophora mangle has proven quite effective for this purpose and has been shown to improve water clarity in near-shore environments. as the wood is generally hard and resistant to insect borers. Windbreaks Red mangrove forests provide a windbreak along coastal margins generally. shellfish. Where present. It is not clear about the success or effectiveness of this campaign. R. Several observations demonstrate the diversity and quantity. fallen leaves would be washed away with each tide. If it were not for the small mangrove crabs. In various locations throughout the region. including birds. For introduced stands in the Hawaiian islands. The resulting mulch is rapidly colonized by bacteria and consumed by other burrowing fauna to release nutrients that appear to further enhance the forest. their floating propagules are spread widely by ocean currents over great distances. Also. soil conditions. red mangroves serve as habitat for a wide range of terrestrial and arboreal wildlife. Interestingly. Host to crop pests/pathogens AGROFORESTRY/ENVIRONMENTAL PRACTICES Mulch/organic matter Mulch in Rhizophora forests is hidden from view. sesarmid crabs consume a large quantity of fallen leaves and propagules. Converting the wood to charcoal can further enhance the timbers’ calorific value. Numerous insects. Native animal/bird food Red mangrove is a known source of native animal foods. these mangrove forests provide shelter and food for a number of associated fauna. one reason stated for the introduction of red mangrove to Hawai‘i was as a “pasture plant for bees” (Cooke 1917). Few mammals (probably none in introduced Hawaiian stands) appear to use red mangrove as a major food source. mangle stands have served as ideal sites for the non-native cattle egret (Bubulcus ibis) and also sometimes harbor significant populations of rats. small mammals. A major reason that red mangrove was introduced into Hawai‘i was to stabilize mud flats that were expanding as a result of sugarcane production and resultant erosion. mangle has unfortunately now taken on the public status of invasive weed and pest species in Hawai‘i.The species R. and mollusks graze on green leaves in the forest canopy and on fallen leaves on the forest floor. and other marine life. To alleviate public concern. and productivity (as expressed in litter fall) is higher. The crabs actively take leaves into underground burrows and chambers. This is done commercially in SE Asia and Central America with various Rhizophora species. Organic matter processed by these herbivores is believed to broadly support aquatic food chains in coastal regions. presumably due to its role in sediment trapping and stabilization. Red mangrove timber is very useful for small construction and for fuel for cooking. as it appears to have been based on subjective information and no monitoring. crabs. Wildlife habitat In addition to aquatic marine organisms (see Fish/marine food chain). with the exception of insect borers and crabs that prey on propagules. No reports are found of red mangroves serving as hosts for known major crop pests or pathogens. but they do produce copious pollen that is usually distributed by wind. and places to seek sanctuary during typhoons and hurricanes. and it is thought to be extremely important in mangrove ecosystems. In Hawai‘i. Woodlot Mangroves adjacent to peoples’ homes throughout the Pacific frequently serve as informal woodlots. damage to propagules and leaves is notably lower than within the species’ native range. eradication efforts have been carried out in several locations on O‘ahu and Hawai‘i.

Staple food  Leaves and hypocotyls are edible but not widely used for food. and tannin for tanning leather. wall-cladding. as do species of shrimp and crabs. flooring.BENEFITS OF MANGROVE TIDAL WETLAND Benefits include. as well as edge trees. with uniform or more or less striped grain. based on mangroves being a depositional site for both water and airborne sediments. the most widespread use of red mangroves is for wood used for a range of purposes from cooking fuel to construction of homes and canoe parts. tool handles. protecting coastal areas. dysentery. In Hawaiian populations. The wood is also used for other purposes. and there are laws in many locations aimed at protecting mangroves in large part because of this important function. notably timber • secondary production. fixation. • shoreline protection. trapping. malaria. where forests moderate evapotranspiration to create a specialized niche climate rise to forest growth and forest products.. and via decomposition • fishery products.g. and improving water quality..traditionaltree. fencing. mangle is light red to dark red or reddish brown or purplish. giving life cycles. Animal fodder One report (Morton 1965) concluded that red mangrove leaves might serve as a valuable source of cattle feed. including microbial and faunal production. although it is not clear how effective the treatments have been in these cases. or pink- Coastal protection  Red mangrove forests and mangroves in general play an important role in protection of coastlines. Medicinal  Red mangrove bark has reportedly been used to treat angina. mangroves may have several other uses (e. poles. medicines. and copra huskers to use as a source of chips for pulp production. In terms of direct benefits to people. The leaves and bark have been used as an antiseptic and to treat diarrhea. because they (and mangroves generally) are believed to play a vital role in supporting marine food chains.g. fishponds. and leprosy. including migratory birds and fish. The sapwood is yellowish. beams. fever. Costa Rica). senescent leaves that have fallen from Rhizophora trees are taken by grapsid (small mangrove) crabs into their burrows. in Terraba-Sierpe Forest Reserve. As mentioned above. Other uses of the tree include dyes. including both estuarine and coastal • nursery habitat for young fauna. Timber The wood of red mangroves is widely used for structural components of traditional homes (e. USES AND PRODUCTS Red mangroves are probably of greatest value for their environmental benefits. as well as grazers. there appears to be excessive leaf accumulation in some locations. rafters) and other components including underground mine supports. The heartwood of R. where mangroves provide protection and a food resource • primary production based on photosynthesis. spears. cabinet works. and boat anchors.g. and turnover • mesoclimate. which in turn reduces turbidity of coastal waters.org)  1 . It seems likely that red  Species Profiles for Pacific Island Agroforestry (www. based on general mangrove tree and root structure. grayish. as cattle feed) that to date have not been fully explored. which reduce erosion and provide stand protection from waves and water movement • carbon sequestration and a sink where carbon is bound within living plant biomass • sediment trapping. boils. suggesting that normal associated fauna and other decompositional biota are lacking. and fungal infections. ranging from traditional uses such as fishing stakes. where mangroves provide a source of food and physical protection from predation • sanctuary niche for mature fauna. and other coastal infrastructure.. but this potential has yet to be realized. in no particular order (adapted from Tomlinson 1986): • visual amenity and shoreline beautification • habitat use by fauna • nutrient uptake. Red mangroves are planted for coastal or fishpond protection in some areas (e.

This is done with all local Rhizophora species.C. Canoe/boat/raft making The wood has been used to make canoe parts.3–5 ft). It was of great importance about 2. Wider spacing and planted sites. Costa Rica. A subsequent investigation (Duke 1996) found Spacing natural recruitment was 40 times greater than planted Mangrove plantations in general are typically planted at numbers.000 seedlings. Duke groves in 1986 (Duke et al. tal plantings eventually amounted to in excess of 100.0–1. Atlantic Panama. supported by IUCN and CATIE.5 m (8 ft) tends to result in a high proportion of that naturally recruited sites recovered more quickly than multiple stemmed and/or shorter trees. possibly because of site damage during planttherefore spreading trees may be desired for coastal protecing. Projects to replant and rehabilitate mangrove forests have been conducted where they have Top: Charcoal treatment plant established with the DANIDA Mangrove been damaged in a significant way. Bottom: in Bahia las Minas. a large Typical earthen kiln used to convert stacked red mangrove poles into charoil spill killed around 50 ha (124 ac) of man. mangle seedlings using clean terrestrial sediments. grain straight to irregular. The calorific value of the timber is significantly enhanced by converting it to charcoal.89. without distinctive odor or taste. Fuelwood Red mangrove wood is used for fuelwood in many places along the American Pacific coast (e. COMMERCIAL CULTIVATION Red mangrove timber is harvested commercially for charcoal production through much of its range. Toaction with each tide and storm. and natural seedlings grew equally as well as. Spacing wider than better than. The specific gravity (ovendry weight/green volume) is 0. For example. or spacings of about 1. damage to exposed and damaged mantion projects but not for timber production. Because the soils were oiled it was decided to plant timber dramatically deteriorated and was moved by wave R. Tannin content of the bark is high for most Rhizophora species. low-luster. In the absence Rehabilitation and replanting 1  Atlantic–East Pacific Rhizophora species  .coal for local heating needs.g. as well as Pelliciera and Laguncularia. The wood is also made into charcoal in many Central American countries. Panama.ish. Pacific Panama. Tannin/dye The bark and hypocotyls are used to tan leather and to produce dyes ranging from red-brown to black (the latter with repeated dying). 1997).5 m (3. including Central and Latin America. photos: N. Project in Pacific Costa Rica. Furthermore. Nicaragua). including Panama and Costa Rica. planted seedlings. and Refineria Panama lead a project to replant the damaged grove areas increased after 5–6 years when dead standing areas. The wood texture is fine to medium..

Some published guidelines for mangrove silviculture exist and are referenced below. Duke et al. with a total mangrove forest stand volume around 280 m3/ha (4000 ft3/ac). The average stand volume for red mangroves was 163 m3/ha (2330 ft3/ac).5 to 3. and each species has its own ecological and economic values. It is also important to plant associated buffer areas. Atlantic Panama.C. with little available except for firewood and charcoal in most places. other than indirect products such as mangrove crabs and fruit bats. Duke  Species Profiles for Pacific Island Agroforestry (www.org)  1 . INTERPLANTING Some interplanting systems include: Red mangroves naturally occur in mixed-species stands. are lacking. particularly along the shoreline where mangroves grow better adjacent to banks stabilized by shoreline upland plants. For the southern (wetter) Pacific coast of Costa Rica. but specific guidelines on thinning. Markets Markets for red mangrove products are local in nature. yields are estimated at 60 m3/ha/yr (860 ft3/ac/yr). 1997) Location Bahia las Minas. Chong (1988). Around 40 ha (100 ac) were planted with Rhizophora mangle. fertilizing.5 m (5 ft). Together they will complement and enhance the richness and stability of the planted environment. A detailed management plan was proposed in the report. and for the co-dominant mangrove Pelliciera. In Panama there is a strong trade in red mangrove poles around 3 m (10 ft) in length (around 10–15 years old) for construction of bohio. Estimated yield Chong estimated that each hectare of mangrove in this reserve could produce US$619 ($250/ac) annually. In Central America. Panama Atlantic coast Description Planting was undertaken in 1988 and 1989. timber plantations should be thinned to spacing of 2. It was reported that yields resulting from uncontrolled felling of 100 ha/yr (250 ac/yr) could be achieved by felling just 10 ha/yr (25 ac/yr) if conducted more systematically. Estimated yields of timber volume under bark were 100–150 m3/ha/yr (1400–2100 ft3/ac/yr) for red mangroves. sponsored and implemented by Refineria Panama. large quantities of red mangrove wood chips and charcoal Planting of red mangroves in an attempt to replace around 50 ha of tree may be moved greater distances and in greater loss following a major oil spill in Bahia las Minas. the locally popular outdoor recreation shelters for barbecues and parties. etc. Spacing/density of species Spacing was about 1. in an FAO-sponsored project.5 m (8–11 ft) between trees as the stand develops and becomes crowded. Management objectives and design considerations volumes than other wood products. Growing in polycultures Example 1—Bahia las Minas oil spill (Duke 1996.. Crop/tree interactions Not applicable.traditionaltree.800 ac) of mangrove forests in the surrounding Terraba-Sierpe Forest Reserve.of significant natural mortality. photo: N. provided information on potential yields from the 240 ha (590 ac) Playa Garza Pilot Area in an investigation of the feasibility of managing around 5200 ha (12.

Predation on propagules regulates regeneration in a high-density reforested mangrove plantation. and N. Koedam. 2005. Stand structure and productivity of the introduced Rhizophora mangle in Hawaii.G. 1999.M. Dahdouh-Guebas. Forest management plan for Playa Garza pilot area: Terraba-Sierpe Mangrove Reserve. H. Hydrobiologia 295: 167–181.M. 4: 131–187. L. In: Field. PhD Thesis. De Lacerda. Hydrobiologia 347: 83–89. Paris.marine. Sooriyarachchi. Bosire. Mangrove reforestation in Panama: an evaluation of planting in areas deforested by a large oil spill. n. R. Letter appended to paper by V.). D. Rhizophora mangle. 1997. Townsville. 1917.D. J.P.d. 1  Atlantic–East Pacific Rhizophora species  . Washington. and N. Okinawa. Traditional and medicinal uses of mangroves. Genetic diversity.V. Alongi (eds. F. Koedam. Koedam. 2002.edu. Economic Botany 54(4): 513–527. Duke.. Verneirt. The species to be planted included R. Mangrove Ecosystems: Function and Management. Jayatissa. In: A.C. Dahdouh-Guebas. G.purdue. mangle. <http://www. P. D. Verneirt. L. W. and F. S.au/marbot/index.L. D. Mangroves and Salt Marshes 2: 133–148.uq. C. Global Ecology and Biogeography Letters 7: 61–71.O. Duke. S.. 1992.). 2005. Koedam. 1995. C. J. Cooke. Ding. J. James Cook University.F. Mangroves as alien species: The case of Hawai‘i. AIMS Report #28.A. H. In: Vozzo. 1998. Estuaries 22(2A): 276–284. J. Marine Ecology Progress Series 299: 149–155. M. Restoration of Mangrove Ecosystems. Nitto. Duke.C. W. Terraba-Sierpe. This project reached the planning stages only and was sponsored by FAO with IUCN and CATIE. J. Nat. N.P.A.. Hist. Mus. Dahdouh-Guebas.W. Crop/tree interactions Not applicable. Costa Rica. Kairo. Tropical Mangrove Ecosystems. Tack.edu/newcrop/ duke_energy/Rhizophora_mangle. Australia. 2005. Dahdouh-Guebas.C.hort. racemosa. Tack. The effect of Coccotrypes fallax (Coleoptera. USDA Forest Service. B. Mangrove floristics and biogeography. Blumea 10(2): 625–634. Rhizophora mucronata Lam.A. In: Handbook of Energy Crops. 1960. Food and Agriculture Organization of the United Nations (FAO).html>.I. Koedam.A.. Propagule predators in Kenyan mangroves and their possible effect on regeneration. Mathenge. Van Speybroeck.C. Utilization of mangrove wood products around Mida Creek (Kenya) amongst subsistence and commercial users. N. 1998. School of Biological Sciences. 2001. Berlin. Allen. J. Japan. 1998. Townsville.M. 1965. Lo Seen. Lucia QLD 4072 Australia Web: <http://www. and N. J. O.A. Kazungu. M. Hawaiian Forester and Agriculturalist 14: 365– 366. Economic. Washington. Bosire. Spacing/density of species Spacing was planned to be 1. traditional and medicinal uses of mangroves.. Marine Botany Group The University of Queensland St. Duke. F. 2002. Current Biology 15(6): 579–586.C. F. 2000. Dahdouh-Guebas. Description Planned project. and J. J. International Society for Mangrove Ecosystems ISME and International Tropical Timber Organization (ITTO). and N. J. MacCaughey.O. D. Koedam.).M.htm> BIBLIOGRAPHY (☛  indicates recommended reading) Allen. Chong. (ed. D. Bandaranayake. Australian Institute of Marine Science. Scolytidae) on the recruitment of Rhizophora stylosa (Family Rhizophoraceae) in North Queensland mangroves.V. N. 1996. N. Robertson and D. Tropical Tree Seed Manual. D. Queensland. Nat. (ed. Bandaranayake. American Geophysical Union. Seen. Cuong. F.Example 2— Terraba-Sierpe Forest Reserve (Chong 1988) Location Costa Rica. 1988. F. Rhizophoraceae. E. Brook. L.5 m (5 ft) for all species. How effective were mangroves as a defence against the recent tsunami? Current Science 15: 443–447. and Pelliciera rhizophorae. Dahdouh-Guebas. Cox. Germany. Transitions in ancient inland freshwater resource management in Sri Lanka affect biota and human populations in and around coastal lagoons. Springer-Verlag. Batelaan. Marine and Freshwater Research 49(4): 345–350..D.P. Hettiarachchi. distributional barriers and rafting continents—more thoughts on the evolution of mangroves.F. Jayatissa. as there were no crops. 1999. J. PUBLIC ASSISTANCE Centre for Marine Studies. and N.F.. Allen. A review of the genus Rhizophora—with special reference to the Pacific species. Food preferences of Neosarmatium meinerti de Man (Decapoda : Sesarminae) and its possible effect on the regeneration of mangroves.

2003. Quek. La-Rue.. Z. B. 1935. Dahdouh-Guebas.C.L. and M. and C. Lo.J. Hawai‘i. Weinstock. Gap creation and regenerative processes driving diversity and structure of mangrove ecosystems.J.G. Smith.htm>. Development and Conservation Priorities. Kairo. C.. E. Lāwa‘i. A. CRC Press Inc. and D. Restoration and management of mangrove systems—A lesson for and from the East African region. <http://www.S. Muzik 1951. ☛ Tomlinson.P. <http://www2.P. Japan.D. Appl. In: L. Duke. W. Report SOITF-SM-2. S. and J.G. A preliminary study of resprouting ability of some mangrove species after cutting.. Flora Vitiensis Nova: A New Flora of Fiji.html>.C. ☛ Hogarth. Association of Pacific Coast Geographers Yearbook 43: 125–137..B. Cambridge. A. Wilkinson. 2003. The Biology of Mangroves. Exploitation of mangal. United Nations University Press.). 5(3): 179–237.C. Factors influencing biodiversity and distributional gradients in mangroves. N.unu. International Society for Mangrove Ecosystems (ISME) and the International Tropical Timber Organisation (ITTO). M. Flora Mal.. Systematic anatomy of the woods of the Rhizophoraceae. Idaho. 1997. and C. Lo. W.I. Adapted from: Chudnoff. 2002. (ed.edu/ unupress/unupbooks/80824e/80824E0p.E. Economic Botany 48(2): 210–213.R. and C. Ellison. Nat. C. Primary productivity of mangroves. 1982..).).traditionaltree. Amsterdam.. S. J. J. In: V. Boston.us/TechSheets/Chudnoff/TropAmerican/htmlDocs_tropamerican/Rhizophoramangle. E.. 2001.. Alongi (eds. Rio Piedras.M. Marine/Coastal Biodiversity Vol. Rhizophora. Global distribution and genetic discontinuities of mangroves—emerging patterns in the evolution of Rhizophora. South African Journal of Botany 67: 383–389. J. II: 3–19.C. In: A. Pinzón. Black (eds.Duke. 2. n. N. F. H. Ecosystems of the World: Wet Coastal Ecosystems. Aging Rhizophora seedlings from leaf scar nodes: a technique for studying recruitment and growth in mangrove forests. Propagation protocol for production of container Rhizophora mangle L. 1996. Biotropica 29: 2–14. Philip. <http://www. Sc. G. Duke.R.D. and P. and M. and M. Eldredge. J. Society for Conservation Biology. F. and R.. Maragos. Riley encased methodology: principles and processes of mangrove habitat creation and restoration. Chapman (ed.d. L. M. Mangroves and Salt Marshes 3: 207–213.Y. Silviculture and Conservation. 1986.C. N. Does the mangrove really plant its seedlings? Science 114: 661–662. Duke. 1977. Brown. 2001. Elevitch. Marco. Puerto Rico. J. Salgado Kent. One hundred Pacific island agroforestry species. (1)54: 429–444. Agriculture Handbook 607.C. Madison. Wood Technical Fact Sheet. K. 2002. Moscow. Vol. and M. USDA Forest Service. Ellison.C. Wisconsin. Tropical Timbers of the World. J. ☛ Saenger. 1998. 2002. Okinawa. feed and fertilizer? Economic Botany 19: 113–123. Bull. Morton. 1965. Sun. Thaman.C. Thaman.B. 1994. N. 1981. Cambridge University Press. CRC Handbook of Biosolar Resources. 1999. Can the red mangrove provide food. USDA Forest Service.org)  1 . Pacific Tropical Botanical Garden. 1936. College of Natural Resources. Bosire.A. Kaua‘i. R. samoensis Hochr.V. Walsh. Duke. and Z. Tropical Woods 44: 1–20.F. N. 1999. Tsuda. Tokyo. D.fpl.). New York.C. Univ. 1958. Restoration of mangrove ecosystems. Field. N.Y. Forest Products Laboratory. Wetlands Ecology and Management 9: 257–269. C. Biotropica 24(2a): 173–186. Koedam.C.org>. ☛ Robertson.W..). Holthus (eds.C. Rhizophoracaeae. Introduction to Rhizophoraceae. Agroforestry in the Pacific Islands: Systems for Sustainability. Regeneration of Rhizophora mangle in a Caribbean mangrove forest: interacting effects of canopy disturbance and a stemboring beetle. American Geophysical Union. Ajima. In: Native Plant Network. Pacific Science Association/East West Center.M. Sousa. P. Tropics 8(3): 221–224. Washington.C.  Species Profiles for Pacific Island Agroforestry (www.. In: Mitsui.J. P. Elsevier Science Publishing Company. Mangrove Ecology. University of Idaho.P. Salvoza. Oecologia 137: 436–445. The Botany of Mangroves. Sun. The Tropical Island Pacific Region: Population.nativeplantnetwork.C. Large scale damage to mangrove forests following two large oil spills in Panama.J.C. Global Ecology and Biogeography Letters 7: 27–47. Florida. plants. Pinzón. In: Clarke. Trees Structure and Function 16: 65–79.S. 1981. Kluwer Academic Publishers. Family 96. Present status of Pacific Island mangroves. and N. 1999. Oxford University Press. and B. S. Prada. Duke. J. ☛ Snedaker. P. Tropical Mangrove Ecosystems. Boca Raton.C. Rhizophora mangle L. Honolulu. A. Patterns of local wood use and cutting of Philippine mangrove forests. 1984. 2005.). Introduction and spread of mangroves in the Hawaiian Islands. R. Rhizophora mangle. and M..fed. 1993. Walters. United Kingdom. and T. M. ☛ Jimenez. Steenis. Riley. 1999.C.E.S. Smith (ed. Economic Botany 59(1): 66–76. var.R. 1999. Forest Research Nursery. Rhizophora mangrove agroforestry..fs. Institute of Tropical Forestry. Wester.F. Ball. 1992. Mitchell.A. Phylogenetic evaluation of Rhizophora taxa for conservation management.C.

R. John Bunt. × harrisonii (Atlantic– East Pacific red mangroves). Dale Evans. 2006. Allen2 1. Candy Feller. R.org>. Paul Smith’s College. Ken Krauss. The University of Queensland. Gina Holguin. Hōlualoa. Fax: 808-3244129. Web: http://www. USA. Hōlualoa.org) Rhizophora mangle. Tel: 808-324-4427. USDA Natural Resources Conservation Service (USDA NRCS). C. Ulf Mehlig.marine.traditionaltree. R. R. Hawai‘i 96725. In: Elevitch.org>. L. with credit given to the source. 2. and Nick Wilson.traditionaltree. Duke1 and James A. This institution is an equal opportunity provider. Eugenia Lo. samoensis. Kathy Ewel. racemosa. Reproduction: Copies of this publication can be downloaded from <http://www.S. Norm Duke acknowledges a host of people who have over the years supported his journey into Rhizophora and who share the passion for this group of plants. ver. and Extension Service. Rhizophora mangle. and Diane Ragone for their input. Kandasamy Kathiresan. Farid Dahdouh-Guebas. Collaborators and friends include: Jim Allen.traditionaltree.Species Profiles for Pacific Island Agroforestry (www. 2. (ed. Jayatissa. Farid Dahdouh-Guebas. © 2006 Permanent Agriculture Resources. Molly Crawford.net. Utah State University. Department of Agriculture. Mei Sun. U. racemosa. XueJun Ge. E-mail: par@agroforestry.P. Ariel Lugo. Australia. under Cooperative Agreement 2002-47001-01327. Recommended citation: Duke. This material is based upon work supported by the Cooperative State Research. 1  Atlantic–East Pacific Rhizophora species  . Ute Steinicke. R. USA.net>. R.1. <http://www. SPC/GTZ Pacific-German Regional Forestry Project.. Education. and the USDA Forest Service Forest Lands Enhancement Program. Lucia QLD 4072. Paul Smiths.au/marbot/index. PO Box 265. Marine Botany Group. State of Hawai‘i Department of Land & Natural Resources Division of Forestry & Wildlife. Series editor: Craig R.agroforestry. and J.). A photo contribution by Ulf Mehlig is greatly appreciated. All rights reserved. Elevitch Publisher: Permanent Agriculture Resources (PAR). Uta Berger. <http://www. Allen. Jurgenne Primavera.A. Web: <http://www.uq. Hawai‘i. Kathy Ewel. Sponsors: Publication was made possible by generous support of the United States Department of Agriculture Western Region Sustainable Agriculture Research and Education (USDA-WSARE) Program. PO Box 428.htm>. Species Profiles for Pacific Island Agroforestry.edu Acknowledgments: The authors and publisher thank Tom Cole. × harrisonii (Atlantic–East Pacific red mangroves) Authors: Norman C. Jaime Polania.C. Otto Dalhaus. St. New York 12970.paulsmiths. Centre for Marine Studies. Permanent Agriculture Resources (PAR). and Agricultural Experiment Station.R.edu. samoensis. N. This publication may be reproduced for noncommercial educational purposes only.