Rhizophora apiculata, R. mucronata, R. stylosa, R ... - Agroforestry Net

Rhizophora apiculata, R. mucronata, R. stylosa, R. × annamalai, R. × lamarckii 

(Indo–West Pacific stilt mangrove) 

Rhizophoraceae (mangrove family) 

mangle hembra (Guam: Chamorro); tebechel, bngaol, agpat, apgal (Palau); raway, roway, ravey (Yap Islands, FSM); chia, 

cia (Chuuk, FSM); aak, akelel, akapah (Pohnpei, FSM); kahrak, suhkasrik, sakasrik (Kosrae, FSM); petu rogha, ngochangochara 

(Marovo, Solomon Islands); tiri tambua (Fiji); koriki (Daru and Kiwai, PNG); totoa (Motu, PNG); spotted leaved 

red mangrove (Western Australia); red mangrove, tall-stilted mangrove, prop root mangroves (Queensland) 

IN BRIEF 

Species Profiles for Pacific Island Agroforestry 

www.traditionaltree.org 

Distribution Native to tropical and subtropical 

coastal areas from the African east coast, 

throughout Asia to Australia and to most islands 

of the eastern Pacific Ocean. Closely allied 

with Atlantic–East Pacific red mangroves 

whose ranges naturally overlap only in a small 

number of southern Pacific islands. 

Size Can reach 30–40 m (100–130 ft) in height, 

although commonly reaches 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 Mangrove communities. 

Soils Adapted to a wide range but thrives best 

in fine mud sediments of downstream river estuaries. 

Growth rate Grows

INTRODUCTION 

Indo–West Pacific stilt mangroves (IWP Rhizophora species) 

are widespread throughout most tropical coastal areas 

of the western Pacific region to east Africa. This group is 

one of two that make up the genus Rhizophora and consists 

of three species (two being closely allied), R. mucronata, R. 

stylosa, and R. apiculata, and two hybrids, R. × lamarckii 

and R. × annamalai. 

Stilt mangroves thrive under a range of intertidal conditions, 

including a range of salinity levels from near freshwater 

to full strength seawater. They tolerate a range of 

flooding regimes, soil types, and other physical site factors. 

Typically, these mangroves are common in the mid-intertidal 

zone, and particularly along the seaward margin of 

tropical mangrove stands. 

Rhizophora are considered the most important of all mangrove 

genera across the Pacific tropical region. However, 

the benefits provided by stilt mangroves are difficult to 

separate from the larger role of mangroves and mangrove 

ecosystems in general. Mangroves are known to play a vital 

role in shoreline protection, enhancement of water quality 

in nearshore environments (including coral reefs), and in 

supporting estuarine and marine food chains. In most parts 

of the Pacific, trees are harvested for firewood, so the trunk 

is the main part of the tree considered for direct use. 

DISTRIBUTION 

Native range 

Indo–West Pacific stilt mangroves occur widely throughout 

the western Pacific. Specifically, they occur in tropical 

and subtropical, intertidal wetlands from the east coast of 

Africa, through Asia to Micronesia extending east in the 

northwestern Pacific through the Federated States of Micronesia 

to the Marshall Islands, and south to northern 

Australia extending east in the southern Pacific as far as 

Samoa. Distributions appear continuous, but separate species 

have different preferred locations where they dominate: 

• 

• 

• 

• 

R. mucronata occurs mostly in areas subject to regular 

freshwater flows (at least in the eastern part of its 

range) 

R. stylosa occurs in marine situations often preferring 

more exposed offshore sites 

R. apiculata is often found mid–lower estuary in larger 

riverine estuaries and embayments 

the hybrid R. × lamarckii is found downstream in middle 

to higher intertidal locations. 

Indo–West Pacific Rhizophora species 

Rhizophora mucronata and R. stylosa are sibling species (i.e., 

possibly R. stylosa =R. mucronata var. stylosa (Griff.) Salvoza), 

and together they characterize most stands of IWP 

stilt mangroves. The range of R. mucronata is the widest of 

all IWP species, extending from east Africa, where it occurs 

as the sole Rhizophora, to the western Pacific, where it 

overlaps with all other IWP species. In contrast, R. stylosa 

extends exclusively east, notably into the southwestern Pacific. 

The range of R. apiculata also extends further east in 

the south Pacific than R. mucronata, but otherwise its range 

mostly fits within the ranges of other IWP Rhizophora. The 

status of observed differences in similar forms, R. mucronata 

and R. stylosa, will only be resolved in genetic studies 

and selected breeding programs. 

The two hybrid taxa presumably have distributions in keeping 

with their hybrid status. As such, they are located wherever 

the ranges of their putative parents overlap. Confirmed 

records of hybrid distributions show R. × lamarckii to be 

widespread in the western Pacific, while R. × annamalai 

was found in India and Sri Lanka along the western limits 

of R. apiculata. However, considerable confusion is expected 

in the field distribution records, because these hybrids 

WHAT IS A MANGROVE? 

Mangroves form a unique and dominant ecosystem 

comprised of intertidal marine plants, mostly trees, 

predominantly bordering margins of tropical coastlines 

around the world. These halophytic (salt tolerant) 

plants thrive in saline conditions and daily inundation 

between mean sea level and highest astronomical 

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 flooding tides allow 

uptake of river-borne nutrients and frequent dispersal 

by their buoyant propagules. 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, Duke et al. 1998), 

compared with adjacent rainforests that may have 

hundreds of tree species per hectare. Specialized morphological 

and physiological characteristics largely 

define and characterize mangrove plants, such as buttress 

trunks and roots providing support in soft sediments, 

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, representing 

independent instances of co-evolution over 

millions of years to form today’s mangrove habitats.

World distributions of stilt mangroves, the IWP Rhizophora species. Given the hybrid status of Rhizophora × lamarckii (= R. 

stylosa × R. apiculata) and R. × annamalai (= R. mucronata × R. apiculata), their distribution is likely where the distributions of 

their respective parent trees overlap. Coastlines marked in red indicate the distribution of all mangroves. image: N.C. Duke 

are difficult to distinguish based on morphological characters 

alone. 

Of great interest also in the southwestern Pacific islands 

is the only occurrence of another hybrid taxon, R. × selala 

(Salvoza) Tomlinson, derived from putative parents R. 

samoensis (=R. mangle?) and R. stylosa (=R. mucronata var. 

stylosa?). This hybrid taxon is special because R. samoensis 

is a key member of the AEP red mangrove Rhizophora 

species. The occurrence of this hybrid means there appears 

to be very little genetic separation between these defining 

and most divergent of Rhizophora species. 

Current distribution 

There are no reports of any stilt mangroves occurring outside 

their natural range. However, these species are recognized 

as valuable timber producers, so it is possible that 

their dispersal east in the Pacific and elsewhere may have 

been assisted by indigenous peoples in the past. 

BOTANICAL DESCRIPTION 

Preferred scientific names 

Rhizophora mucronata Lamk. 

Rhizophora stylosa Griff. (= R. mucronata var. stylosa (Griff.) 

Salvoza) 

Rhizophora apiculata Bl. 

Rhizophora × lamarckii Montr. (= R. stylosa × R. apiculata) 

Rhizophora × annamalai K. Kathiresan. (= R. mucronata × R. 

apiculata) 

Family 

Rhizophoraceae (mangrove family) 

Common names 

aak, akelel, akapah (Pohnpei, FSM) 

abat (Ceram, Indonesia) 

ailave, kailau (Aru, Indonesia) 

bakau (Malay Peninsula, Singapore, Borneo, Sumatra) 

bakauan, bakau, bakhaw (Philippines) 

bako ( Java, Indonesia) 

bangka (Acheh, Indonesia) 

chia, cia (Chuuk, FSM) 

Indo–West Pacific stilt mangroves (English) 

kahrak, suhkasrik, sakasrik (Kosrae, FSM) 

koriki (Daru and Kiwai, PNG) 

lolaro, belukap (Sulawesi, Indonesia) 

mangle hembra (Guam: Chamorro) 

ngochangochara, petu rogha (Marovo, Solomon Islands) 

prop root mangroves (northeastern Australia) 

Species Profiles for Pacific Island Agroforestry (www.traditionaltree.org)

away, roway, ravey (Yap Islands, FSM) 

red, tall-stilted mangrove (Queensland) 

spotted leaved red mangrove (Western Australia) 

tebechel, bngaol, agpat, apgal (Palau) 

tiri tambua (Fiji) 

totoa (Motu, PNG) 

Size 

Indo–West Pacific stilt mangroves are medium to tall trees 

that may reach 30–40 m (100–130 ft) in height, although 

they are commonly much shorter, around 5–8 m (16–26 ft). 

Stem diameters are often about 15–35 cm (6–14 in) taken 

just above the highest prop root. This measure differs fundamentally 

from the standard diameter at breast height 

(dbh) used for most forest surveys, as diameter height 

above the substrate varies from 0.5 to 7 m (1.6–23 ft) (consider 

the tree in the photo on the first page). 

Form 

Stilt mangroves are rambling to columnar trees with distinct 

aboveground prop roots. Trees tend to be of shorter stature 

and more spreading in shape on the seaward edge of stands 

or in areas of higher salinity. Taller, single-stemmed trees 

are found most often just behind the water’s edge of stands 

midstream in major riverine estuaries. Multi-stemmed 

trees are common in more arid or marginal habitats. 

Flowering 

Flowers are perfect. Inflorescences have few to many 

joints with 1,2-chotomous branching and one to many 

buds possible per inflorescence. Open flowers are located 

within or below leaf axils at leaf nodes below the apical 

shoot, depending on species. For R. mucronata, R. stylosa, 

R. × annamalai. R. × lamarckii, and R. apiculata mature 

buds and flowers are located at 1–3, 1–5, 3–5, 3–6, and 6–11 

nodes down from apical shoot, respectively. Calyces are 

typically pale yellow at maturity with 4 lobes, rarely 3. Buds 

are obovate, green when immature to pale yellowish green 

as they mature, dimensions 1–2 cm (0.4–0.8 in) long and ~1 

cm (0.4 in) wide. Petals, usually 4, are lanceolate to linear, 

creamy white, woolly to hairless, ~10 mm (0.4 in) long and 

~2 mm (0.08 in) wide. Stamens number 7–12, pale yellow. 

Style is pale green, terete, 0.5–6 mm (0.02–0.24 in) above 

base, dichotomous tip. Bracts and bracteoles are variable 

depending on species. Mature bud bracts are slender green 

(R. mucronata, R. stylosa), swollen green (R. × annamalai, 

R. × lamarckii) and swollen corky green (R. apiculata). Peduncles 

are 1–7 cm (0.4–2.8 in) long, and ~3 mm (0.12 in) 

wide. Flowering period is chiefly during August–December 

in the southern hemisphere, and during February–June 

in the northern hemisphere. 


Leaves 

Leaves are opposite, simple, light or dark green, obovate, 

leathery, margins revolute, bluntly acute apex with a distinct 

mucronate tip, 1–7 mm (0.04–0.3 in) long. Upper leaf 

surface is smooth, shiny. Cork wart spots occur on undersurface, 

scattered evenly, not raised, present in most species 

in most locations, but absent only from R. apiculata and 

R. × lamarckii in southern New Guinea and Australia. Mature 

leaf dimensions are 6–19 cm (2.4–7.5 in) long, and 3–10 

cm (1.2–3.9 in) wide. Petiole 1–4 mm (0.04–0.16 in) long. 

Leaf emergence is mostly around November–February in 

the southern hemisphere and May–August in the northern 

hemisphere. Leaf fall occurs mostly during the wet summer 

period from October to February in the southern hemisphere 

and April–August in the northern hemisphere. 

Fruit 

Fruits, when mature, are pear-shaped, elongate, waist 

constriction, smooth brown surface, calyx lobes elongate 

spreading (when hypocotyl ready to emerge). For R. mucronata, 

R. stylosa, R. × annamalai, R. × lamarckii, and R. 

apiculata mature fruit located in leaf axils 3–5, 4–7, 5 (rare), 

7 (rare), and 8 nodes down from apical shoot, respectively. 

Seeds/hypocotyls 

Like all Rhizophora species, stilt mangroves are viviparous, 

meaning that the trees produce seeds hidden in the mature 

fruit, and these germinate on the parent tree. The 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 is 

slightly wider, distal tip is pointed in most taxa, but rounded 

to blunt for R. apiculata. For R. mucronata, R. stylosa, 

R. × annamalai, R. × lamarckii, and R. apiculata mature 

hypocotyls are located in leaf axils 4–10, 4–9, none, 8–9 

(rarely observed), and 9–13 nodes down from the apical 

shoot, respectively. Hypocotyl dimensions are variable and 

not consistently species-specific, 14–80 cm (6–31 in) long, 

1–2 cm (0.4-0.8 in) at the widest point, and 0.5–1.5 cm 

(0.2–0.6 in) wide at the “collar,” the fruiting structure that 

envelops the plumule (embryonic leaves). 

“Fruiting,” when mature hypocotyls fall, occurs chiefly (but 

not exclusively) from November to January in the southern 

hemisphere and May to July in the northern hemisphere. 

Bark 

The bark is gray to dark gray and heavily fissured, occa-

Top left: The distinct mucronate (abrupt, pointed) tip at the apex of leaves is characteristic of all IWP stilt Rhizophora. The 

coloration of the mucronate tip shown in this image of R. stylosa is variably red or green. Northeastern Australia. Top right: 

Inflorescence of Rhizophora mucronata (similar to R. stylosa) showing open flowers with distinctive woolly petals and slender 

smooth bract beneath the calyx. Bottom left: Inflorescence of Rhizophora apiculata showing open flowers with distinctive hairless 

petals, non-reflexed calyx lobes, and corky bract beneath the calyx. Bottom right: Leafy rosette of Rhizophora mucronata 

(similar to R. stylosa) showing mature buds and mature fruit. photos: N.C. Duke 

sionally red-brown and smooth. Prop roots are sturdy even 

when relatively thin. 

Rooting habit 

Mature trees have distinctive, sturdy, aboveground prop 

roots surrounding the stem base that anchor only shallowly 

in the sediments to 1–2 m (3.3–6.6 ft) depth. This 

conforms to the oxygen deficient (anoxic) conditions commonly 

measured in mangrove sediments. 

Similar species 

Stilt mangroves are distinguished from Atlantic–East Pacific 

red mangrove species principally by the spiked, mu- 

cronate tip at the leaf apex of stilt mangroves that is absent 

in red mangroves. 

Rhizophora mucronata and R. stylosa have slender (i.e., length 

much greater than the width) bracts at the base of mature 

buds as distinguished from R. apiculata, R. × lamarckii, and 

R. × annamalai that have bracts almost as wide, or wider 

than the length. 

Rhizophora apiculata is distinguished from R. × lamarckii 

and R. × annamalai plus other IWP species by swollen 

(wider than long), corky brown bracts, one inflorescence 

joint, and node position of mature buds and flowers in leaf 

axils at 6–11 nodes down from apical shoot, well below 

leaves in the leafy shoot. 

Hybrid characteristics are shown in R. × lamarckii where 


Mature hypocotyls of Rhizophora stylosa (similar to R. mucronata). 

Great Sandy Straits, NE Australia. photo: N.C. Duke 

Left to right: Mature flower buds of Rhizophora apiculata, 

R. × lamarckii, R. stylosa, and R. mucronata. These samples 

were collected in NE Australia, where the four species grow 

side by side in several estuaries. photo: N.C. Duke 


it has key characters intermediate between R. apiculata 

and R. stylosa. Similarly, hybrid characteristics are shown 

in R. × annamalai, which has key characters intermediate 

between R. apiculata and R. mucronata. Hybrids can often 

be larger and taller than neighboring parent trees. 

Rhizophora × lamarckii is distinguished from R. × annamalai 

by style length

elated to differences in habitat. It is clear that 

R. stylosa is the taxon best adapted to marine 

exposed locations wherever it occurs. 

It is of great interest to fully evaluate the relative 

and precise distributions of R. mucronata 

and R. stylosa, because these are likely to explain 

the way species have naturally evolved and dispersed. 

In some island groups to the north of 

Australia, for instance, there is a curious pattern 

in separate sympatric occurrences where these 

species are easily distinguished in some populations 

(Philippines, Sulawesi) while in others 

(Federated States of Micronesia, Palau) their 

style lengths overlap, making it impossible to 

always tell them apart using this character. This 

may be explained by a progressive lengthening 

of the style in R. stylosa, corresponding possibly 

with its more recent introduction in locations 

with longer styles. If this is correct, R. stylosa 

has spread from Australia and Federated States 

of Micronesia both westward and eastward; 

for example, in Sulawesi and Fiji where style 

lengths are the longest recorded. This idea concurs 

with the specialized marine habit of this 

species. 

By contrast, R. apiculata is readily distinguished 

from R. mucronata and R. stylosa. This species 

appears more prevalent in estuaries influenced 

by larger and more continuous freshwater flows, 

and it is found in a continuous distributional 

range, from India to the western Pacific and northern 

Australia. Across this range there are also two forms of R. 

apiculata, and their occurrences do not overlap. One form, 

found north and west from the northern New Guinea 

coast, fits the type description in every respect including 

the presence of cork wart spots on leaf undersurfaces. The 

other form lacking these spots occurs south and east from 

the southern New Guinea coast. This character was thought 

to be unique to R. apiculata until recently, when trees of R. 

racemosa (an AEP red mangrove) without spots were discovered 

in Brazil. This character helps define two forms of 

R. apiculata in Australasia and the western Pacific. 

Known varieties and hybrids 

Rhizophora mucronata and R. stylosa are both known to 

hybridize with R. apiculata, giving rise to morphologically 

distinct and geographically widespread hybrid forms, 

R. × annamalai and R. × lamarckii, respectively. Their hybrid 

status was initially based on intermediate morphological 

characters but has since been confirmed in genetic 

investigations. 

Variation in style length of Rhizophora stylosa and R. mucronata in the IWP 

region. Note that for each species, but especially for R. stylosa, larger styles 

occur in eastern and western populations. images: N. C. Duke 

As discussed above, R. mucronata and R. stylosa are sibling 

species with few diagnostic characters to separate them 

and a corresponding lack of genetic separation. This clearly 

makes it difficult to separate respective hybrid forms with 

R. apiculata, although this may be done by also using the 

style length character. The situation is more complex, however, 

when it must be considered further that there are the 

two forms of R. apiculata. These each have hybrids with 

R. stylosa based on occurrences north and south of New 

Guinea. 

IWP stilt mangroves are also characterized by a number of 

notable genetic, morphological, and growth variants. Aberrant 

traits are observed in a small number of individuals 

throughout the range. Perhaps the most consistent aberrant 

trait, however, is “albino” hypocotyls (i.e., those lacking 

any green pigment, leaving them red or yellow) observed in 

Australia, Federated States of Micronesia, Palau, and other 

places. Yellow or red propagules can be observed hanging 

in affected trees alongside normal green propagules of all 

IWP species. Their relative number on an individual tree is 

thought to quantify the amount of outcrossing that occurs 

with neighboring normal trees. On rare occasions, other 


trees have also been observed with variegated 

foliage. 

Culturally important related species 

All Rhizophora species are closely similar in tree 

form, and different cultural groups in the Pacific 

region may not always distinguish between 

them. Other mangrove genera, like Bruguiera, 

are considered close in form and value also, and 

these are often used in similar ways by indigenous 

peoples. 

ASSOCIATED PLANT SPECIES 

IWP stilt mangroves are naturally associated 

with a number of other mangrove and nonmangrove 

species across their wide range of 

tidal and estuarine locations. Planting with 

other species is highly recommended, particularly 

terrestrial and beach strand associates 

growing naturally above mean sea level elevations. 

The idea is to establish and achieve wholeof-bank 

stabilization as soon as possible. Since 

stilt mangroves also occupy a range of substrate 

types, including coral, rocks, gravel, sand, and mud, it is 

important to choose associated species best suited to the 

particular local conditions. 

Associated species commonly found 

The associated species vary with geographic location, latitude, 

soil type, estuarine upstream location, and tidal position. 

In frontal stands, IWP stilt mangroves may be associated 

with Sonneratia alba and Avicennia marina downstream. 

Where downstream stands of stilt mangroves are backed 

by sand ridges and beach margins, the associated species 

include Calophyllum spp., Thespesia spp., Casuarina spp., 

Barringtonia asiatica, and Cocos nucifera. 

Upstream in riverine estuaries of tropical coastlines of 

larger islands and continental margins, associated species 

include Sonneratia caseolaris, S. lanceolata, and Aegiceras corniculatum. 

In such upstream locations, stilt mangroves are 

commonly backed by Bruguiera and other higher intertidal 

mangrove species, as well as associated upland species including 

Barringtonia racemosa and Hibiscus tiliaceus. 


Foliage of Rhizophora mucronata showing a tree with a rare yellow “albino” 

form of hypocotyls among the normal green hypocotyls. Johnstone River, 

NE Australia. photo: N. C. Duke 

ENVIRONMENTAL PREFERENCES 

AND TOLERANCES 

Climate 

IWP stilt mangroves thrive in tropical and subtropical 

environments characterized by moderately high and well 

distributed rainfall. However, in drier locations, like in 

northern areas of Western Australia, the stunted but dense 

thickets of R. stylosa attest to the great adaptability of stilt 

mangroves to a wide range of climatic types. 

Elevation range 

0–6 m (0–20 ft), in reference to mean sea level. 

Mean annual rainfall 

These mangroves grow in all rainfall conditions. Their extent, 

form, and biomass reflect the different rainfall conditions. 

Rainfall pattern 

Grows in climates with summer or uniform rainfall patterns. 

Dry season duration (consecutive months with < 0 

mm [1. in] rainfall) 

Mangrove plants appear to depend on groundwater to sustain 

optimal growth, especially during drier months. Across 

a wide variety of climatic regions, mangrove cover expands

and contracts through time. This 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 

23–38°C (73–100°F) (estimate) 

Mean minimum temperature of coldest month 

13–18°C (55–64°F) 

Minimum temperature tolerated 

10°C (50°F) (estimate) 

Soils 

Trees develop greatest stature and columnar growth form 

in estuaries of larger tropical rivers, characterized by fine 

clay, black mud sediments with relatively high loads of organic 

carbon, and anaerobic soils with high concentrations 

of sulfide. Trees also grow well in sites with aerobic sediments 

consisting of fine sands to coarse stones and rocks, 

as well as coral ramparts. 

Soil texture 

Plants grow best in light, medium, and heavy texture soils 

(sands, sandy loams, loams, sandy clay loams, sandy clays, 

clay loams, and clays). 

Soil drainage 

Plants grow best in soils with free and un-impeded drainage, 

as well as waterlogged soils. 

Soil acidity 

pH 6–8.5 

Special soil tolerances 

Plants grow best in saline soils but can survive well in fresh 

water. The optimal salinity range is reported to be 8–26 ppt 

(parts per thousand), compared to approximately 34–36 ppt 

for seawater. 

Tolerances 

Drought 

Stilt mangroves usually grow best in wetter conditions. 

However, R. stylosa and western range R. mucronata can 

tolerate drought periods well. 

Full sun 

They usually grow best in full sun. 

Shade 

MANGROVES AND CLIMATE 

CHANGE 

Indo–West Pacific stilt mangroves, like other mangrove 

species, are affected by climate change. The unique 

physiological characteristics of each species define its 

capacity for survival in the face of change. Mangroves 

are expected to respond rapidly and decisively to shifts 

in key factors, like temperature, rainfall, and sea level, 

as each species has defined ranges of tolerance 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 mangroves have 

narrow optimal temperature ranges, rising temperatures 

will cause their distributions to shift north or 

south to areas where temperature conditions are most 

suitable, and they will die off 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 represents 

the survivors of all past changes. 

Key indicators of change can be identified and mapped 

as incremental shifts and responses of mangrove communities. 

These might be observed as shifts in vegetation, 

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 affects 

moisture stress in saline environments. In both cases, 

the response zones will follow elevation contours. 

Changes along contours can be quantified from longterm 

spatial assessments over decade- and centurylong 

time periods, depending on the rates of change. 

Knowledge of these changes and their causes allows 

better prediction of future change. 

Stilt mangroves are usually considered to have a very low 

tolerance of shade. However, in eastern Australia at its 

southern latitudinal limit, R. stylosa trees commonly grow 

under a closed canopy of Avicennia marina. This may be 


due to the absence of scolytid beetle larvae that 

usually infect shaded propagule-seedlings in 

the tropics (Brook 2001). If this is the case, it 

implies Rhizophora seedlings might be more 

shade tolerant than originally thought (Sousa 

et al. 2003). This is supported further by the 

observation that Bruguiera species, the apparent 

shade specialists of tropical areas, had few 

scolytid infections. 

Fire 

The trees have no tolerance of fire in close proximity. 

Frost 

They have low tolerance of sub-freezing temperatures. 

Waterlogging 

Stilt mangrove trees are tolerant of daily tidal 

flooding up to depths of 2.5 m (8 ft). While 

tolerating permanently saturated soils, they are 

intolerant of drying soils. 

Salt spray 

They are highly tolerant of salt spray. 

Wind 

Some species of stilt mangrove trees are typically 

found on the exposed water’s edge of large 

riverine estuaries. 

Abilities 

Self-prune 

Stilt mangroves self-prune well in dense stands, but they 

commonly maintain lower branches in more open locations. 

Coppice 

The trees have notably poor coppice ability. Generally, if 

greater than 50% or more of the leaves are removed from 

a tree, they will die. There appears to be a variant of R. 

apiculata in the Philippines and in some islands of the 

Federated States of Micronesia (Pohnpei and Kosrae) that 

is used by local people for sustainable firewood collection. 

These trees regrow from regular cutting, which keeps them 

stunted. 

GROWTH AND DEVELOPMENT 

Growth rates vary with age. The tree generally grows less 

10 Indo–West Pacific Rhizophora species 

Trunk of Rhizophora mucronata showing distinctive stilt roots. Kenya, E 

Africa. photo: N. C. Duke 

than 1 m/yr in height but can exceed this in favorable circumstances. 

Height growth is rapid shortly after establishment 

while food reserves are taken up from the hypocotyls 

of established propagules. The growth rates appear to slow 

when trees approach a site maximal canopy height, dependant 

on specific site/location conditions. When near maturity, 

trees tend to spread and broaden their canopy and 

stem diameter rather than grow taller. 

Flowering and fruiting 

Flowering and fruiting periods of stilt mangroves are distinctly 

seasonal. Furthermore, peak pheno-events shift 

later with cooler temperatures and higher latitudes. Trees 

have notable and relatively long periods of reproductive 

development, taking 18–30 months from first emergence 

of flower bud primordia until maturation and drop of mature 

hypocotyls. The duration depends on species, with the

longest being Rhizophora apiculata, with around 30 months 

for each reproductive cycle. 

Yields/growth rates 

Growth rates vary with species, spatial position in the 

stand, competition, vigor, and age. In the Matang Mangrove 

Forest Reserve in Malaysia, Putz and Chan (1986) 

reported that diameter growth rates of R. apiculata trees 

were 0.24–0.29 cm (0.09–0.11 in) for diameter size classes 

from 10 to 60 cm (4–24 in). 

Watson (1928) estimated that under Malaysian conditions 

mean annual increment (MAI) of stilt mangroves culminated 

at around 10.6 m 3 /ha/yr (152 ft 3 /ac/yr) at 39–40 years. 

Reaction to competition 

Rapid early growth of seedlings of stilt mangroves in full 

sunlight ensures their success and dominance in preferred 

estuarine and intertidal conditions. Newly established 

seedlings grow best in close proximity with their same 

species cohort. This affords them maximal protection from 

physical damage by drift logs and erosive waves. Since 

competition is high between neighboring seedlings, slower 

plants die and decompose quickly, leaving faster competitors 

the benefit of not only the space they occupied but 

also their nutrients. 

PROPAGATION 

IWP stilt mangroves are readily propagated by direct 

planting of their propagules. Although natural 

regeneration is generally relied upon around the 

Pacific region, these species are relatively 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 distal end of the 20–80 cm 

(8–31 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 

nursery preparation nor stakes are needed. Low 

maintenance is generally required for maximizing 

seedling establishment success in sheltered 

areas. However, substantial protection is required 

in more exposed coastal locations during 

the first decade of establishment. Such protection 

methods may include encasement of indi- 

vidual 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. Mature propagules may be collected after they 

have fallen or been picked directly off trees. Only healthy 

looking propagules should be selected. Propagules that 

are shrunken or desiccated in appearance or that exhibit 

significant 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 is 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 stilt 

mangroves, although damaged and insect infested individuals 

should be removed. Also see the following pretreatment 

section for possible additional steps. Propagules can 

be sown in nursery beds, or preferably planted directly in 

the field soon after collection. 

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 

Left: Mature hypocotyl and fruits of Rhizophora mucronata showing distinctive 

collar on attached propagule and expended fruit. photo: N. C. Duke 

Right: A sectioned wilting seedling of Rhizophora stylosa shows the extent 

of scolytid infection marked by brown frass. photos: B. m. Brook 

Species Profiles for Pacific Island Agroforestry (www.traditionaltree.org) 11

urlap bags and keeping them out of direct 

sunlight. Pretreatment is considered essential if 

considering such storage (see below). It is likely 

that propagules can be kept longer, but storage 

beyond 2 weeks is not recommended, and longterm 

storage is not feasible. 

Propagule pretreatment 

Pretreatment of propagules is generally considered 

unnecessary. However, a study of scolytid 

beetle larvae (Coccotrypes fallax) infections 

on Rhizophora propagules in Australia found 

at least 18% that were heavily infested (Brook 

2001). Infested propagules were established under 

canopy-shaded areas. The study went further, 

finding that pretreatment in a 50°C (122°F) 

water bath for 5 minutes killed the beetles and 

removed the risk of establishment loses due to 

borer damage. Heat treatment might be easily 

achieved by leaving the collected propagules in 

the sun for a short period (a few hours) before 

planting. 

Growing area 

Stilt 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 

Anytime during the year is suitable. Seedlings are ready 

for outplanting at the 6-leaf (=3 node) stage if grown up 

in nursery conditions. Older seedlings up to 0.5 m (20 in) 

tall have also been successfully transplanted, but this is not 

considered beneficial. 

Direct planting of large numbers of propagules is restricted 

by their peak seasonal availability, as propagules do not 

keep for extended periods unless planted out in nursery 

conditions. 

Guidelines for outplanting 

Propagules or nursery-grown seedlings usually have excellent 

survival in sites correctly selected and, if appropriate, 


New planting on seaward edge of R. apiculata at Ao Khung Krabaen Mangrove 

Forestry Center, Som Lek, Thailand. photo: C. elevitCh 

protected from disturbance. Survival rates of 90% or greater 

are not unreasonable in such circumstances. 

On the other hand, survival may be zero on sites exposed 

to excessive wave action, on sites with inappropriate hydrologic 

or salinity regimes, or (rarely) disturbance by grazing 

animals (e.g., goats, sheep, cattle, and 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, the planting of stilt mangroves poses few significant 

disadvantages when planted within their native ranges. 

They are not especially susceptible to pests or pathogens 

and they have not been reported to host major pests or 

pathogens of important crop species. 

Potential for invasiveness 

Although the invasiveness of stilt mangroves has not been 

demonstrated, they are likely to be opportunistic due to 

their relatively wide tolerance for salinity and soil conditions. 

The spread of the closely allied AEP red mangrove 

species R. mangle when introduced into Hawai‘i has clearly 

demonstrated the potential for invasiveness of these mangroves. 

Although they were not native, suitable growing 

conditions existed, demonstrating that the global distribution 

of Rhizophora spp. is currently limited by their natural

Benefits include, in no particular order (adapted from 

Tomlinson 1986): 

• visual amenity and shoreline beautification 

• nutrient uptake, fixation, trapping, and turnover 

• habitat use by fauna 

dispersal range. This also means geographically isolated 

populations are vulnerable to introductions of genotypes 

from elsewhere. 

Susceptibility to pests/pathogens 

Susceptibility to pests and diseases is believed to be low, 

with the exception of insect borers and crabs that feed on 

propagules. 

Host to crop pests/pathogens 

No reports were found of stilt mangroves serving 

as hosts for known major crop pests or 

pathogens. The scolytid beetles are specific to 

the genus Rhizophora. 

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. 

The crabs actively take leaves into underground 

burrows and chambers. The resulting mulch is 

rapidly colonized by bacteria and consumed by 

BENEFITS OF MANGROVE TIDAL WETLAND 

• mesoclimate, where forests moderate evapotranspiration 

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 fish, where mangroves provide protection 

and a food resource 

• primary production based on photosynthesis, giving 

rise to forest growth and forest products, notably timber 

• secondary production, including microbial and faunal 

production, as well as grazers, and via decomposition 

• fishery 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 depositional 

site for both water and airborne sediments, 

which in turn reduces turbidity of coastal waters. 

other burrowing fauna to release nutrients that appear to 

further enhance the forest. 

Soil stabilization 

Stilt mangrove forests stabilize soils with their network 

of sturdy overlapping prop roots dampening water movement 

and promoting sedimentation in areas that might 

otherwise be eroded. 

Rhizophora mucronata tree growing on the edge of an estuarine channel. 

While the stem leans out over the water to get the most light, the stilts 

both support the tree and stabilize the bank. photo: N. C. Duke 


Fence posts 

Stilt mangrove stems make good posts since they are generally 

hard wood and resistant to insect borers. 

Windbreaks 

Rhizophora forests provide a windbreak along coastal margins, 

generally, and as places to seek sanctuary during typhoons 

and cyclones. Planting in highly wind-prone locations 

is not recommended unless the location has some 

history, or reasonable expectation, that mangrove survival 

is likely. 

Woodlot 

Mangroves adjacent to peoples’ homes throughout the Pacific 

frequently serve as informal woodlots, particularly on 

islands with clear tenure systems that include mangrove 

areas. Stilt mangrove wood is very useful for small construction, 

and for cooking fuel. Converting it to charcoal 

can further enhance the timbers’ calorific value. This is 

done commercially in SE Asia and Central America with 

various Rhizophora species. 

Native animal/bird food 

IWP stilt mangroves are largely an unknown source of 

native animal foods. However, several observations demonstrate 

that the diversity and quantity of the source is 

thought to be extremely important in mangrove ecosystems. 

Numerous insects, crabs, and mollusks graze on 

green leaves in the forest canopy. 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 

appear to use stilt mangroves as a major food source, 

although native rats often chew into the wood in search of 

boring insect larvae. 

Wildlife habitat 

In addition to aquatic marine organisms (see Fish/marine 

food chain), stilt mangroves serve as habitat for a wide 

range of terrestrial and arboreal wildlife. In various locations 

throughout the region, these forests provide shelter 

and food for a number of associated fauna, including birds, 

fruit bats, small mammals, shellfish, and other marine life. 

Bee forage 

Rhizophora species have no nectar, but they do produce copious 

pollen that is usually distributed by wind. 

Fish/marine food chain 

Mangroves in general are believed to play a vitally important 

role in protecting and supporting marine food chains. 

Many fish species use stilt mangroves during part of their 


Rhizophora stylosa sprouted propagules for sale in Iriomote, 

Japan. photo: N. C. Duke 

life cycles, as do species of shrimp and crabs. Species such 

as the mangrove mud crab (Scylla serrata) are common in 

IWP mangroves, and these are important sources of food 

and income on many islands in the region. Populations 

of some smaller species may exceed 10 crabs/m 2 (1 crab/ 

ft 2 ) in parts of the Pacific. Senescent leaves having fallen 

from Rhizophora trees are taken by grapsid (small mangrove) 

crabs into their burrows. Buried leaves decompose 

and contribute to nutrient recycling in mangrove forests. 

Nutrients also feed directly and indirectly to associated estuarine 

and marine food chains. 

Coastal protection 

Stilt mangrove forests, and mangroves in general, play an 

important role in protection of coastlines, fishponds, and 

other coastal infrastructure. Stilt mangroves are planted 

for coastal or fishpond protection in some areas (e.g., in 

Kalibo Bakhawan Eco Park, the Philippines) and there are 

laws in many locations aimed at protecting mangroves in 

large part because of this important function. 

Ornamental 

No significant ornamental trade is known. However, as an

apparently isolated example, a small number of plants are 

sold as souvenirs to departing tourists from Iriomote and 

other southern islands of Japan. Sprouted hypocotyl seedlings 

are used to make bonsai “forest gardens.” 

USES AND PRODUCTS 

IWP stilt mangroves are probably of greatest value for their 

environmental benefits, because they (and mangroves generally) 

are believed to play a vital role in supporting 

marine food chains, protecting coastal 

areas, and improving water quality. 

In terms of direct benefits to people, the most 

widespread use of stilt mangroves is for wood 

for a range of purposes from cooking fuel to 

construction of homes and canoe parts. Other 

uses of the stilt mangroves include tannin and 

dyes. A less lauded benefit has been derived 

from some ecotourism ventures. The dual benefit 

in this practice comes from both a moderate 

economic return combined with a substantial 

longer-term educational value in raising environmental 

awareness in the community. 

Staple food 

Leaves and hypocotyls are edible but not widely 

used for food. 

Medicinal 

Stilt mangrove bark has reportedly been used 

to treat angina, boils, and fungal infections. The 

leaves and bark have been used as an antiseptic 

and to treat diarrhea, dysentery, fever, malaria, 

and leprosy, although it is not clear how effective 

the treatments have been in each of these 

cases. 

Timber 

The wood of stilt mangroves is widely used 

for structural components (e.g., poles, beams, 

flooring, wall-cladding, and rafters) of traditional 

homes and other structures like underground 

mine supports, fencing, cabinet works, 

tool handles, and boat anchors. The wood is also 

used for other purposes, ranging from traditional 

uses such as fishing stakes, spears, and copra-huskers 

to use as a source of chips for pulp 

production. However, pulp from Rhizophora 

species does not have good strength properties, 

so these species are not often sought after for 

this purpose (Percival and Womersley 1975). 

Fuelwood 

Stilt mangrove wood is used for fuelwood on many Pacific 

islands (e.g., Kosrae and Chuuk). The wood is also made 

into charcoal in countries such as the Philippines, Indonesia, 

and Malaysia (Ong et al. 1980, 1995). 

Canoe/boat/raft making 

The wood has been used to make canoe parts. 

Top: Mangrove tourism in Hainan, China, complete with Rhizophora 

motifs on the jetty handrail panels. photo: N. C. Duke Bottom: Boardwalk 

and signage for visitors (R. mucronata on left and R. apiculata on right), Ao 

Khung Krabaen Mangrove Forestry Center, Som Lek, Thailand. photo: C. 

elevitCh 


Tannin/dye 

The bark and hypocotyls are used to produce dyes ranging 

from red-brown to black (the latter with repeated dying). 

Rhizophora species are the best producers of tannins of all 

the mangroves with 20–25% of bark weight (Percival and 

Womersley 1975). Chopped bark is treated by continuous 

countercurrent extraction with first cold then warm water 

producing an extract called “cutch.” The cutch can be 

exported for processing and final tannin extraction. Tannins 

were used extensively for preserving cotton ropes and 

netting. Mangrove tannins can be used to prepare tannin 

formaldehyde adhesives. These adhesives are recognized 

for their high moisture resistance and waterproof grades in 

plywood and particleboard production. Rhizophora tannins 

are also known to impart a red color to finished leather 

tanning, but this is considered undesirable since blending 

with other tannin extracts is needed. 

Ecotourism 

The mangrove environment is attractive and interesting 

to both local people and to visiting tourists. In China, for 

example, facilities have been provided to assist and regulate 

this form of ecotourism, including a dedicated hotel 

and restaurants. Simple boardwalks in other places (e.g., 

Australia, Thailand) are relatively low key but the intention 

is the same, to provide public access for those wanting 

to see and learn about this fascinating environment. The 

boardwalks are there to make the mangrove experience as 

effortless as possible while protecting the mangroves from 

direct damage. Walks are often also accompanied with informative 

and educational signage. 

COMMERCIAL CULTIVATION 

Stilt mangrove timber is harvested commercially for charcoal 

production in SE Asia. The calorific value of the timber 

is significantly enhanced by converting it to charcoal. 

This is done with various Rhizophora species. Charcoal 

production has been conducted in a sustainable way on 

the Malay Peninsula for over 100 years using silvicultural 

practices developed for Rhizophora species, especially R. 

apiculata (Ong et al. 1980, 1995). In addition, forests in Malaysia, 

New Guinea, and the Solomon Islands have been 

clear-felled for wood chips using unsustainable harvesting 

practices. 

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 


Eleven-year-old plantation of Rhizophora apiculata in the 

63 ha (156 ac) Bakhawan Eco Park at Kalibo, northern Panay, 

Philippines. photo: N. C. Duke 

and the resulting spreading trees may be desired for coastal 

protection projects, but not for timber production. In the 

absence of significant 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 

In areas where the mangrove fern (Acrostichum speciosum) 

is common, it may need to be controlled to promote early 

growth of stilt mangroves. Some published guidelines for 

mangrove silviculture exist and are referenced below, but 

specific guidelines on thinning, fertilizing, etc., are currently 

unavailable. 

Growing in polycultures 

Stilt mangroves naturally occur in mixed-species stands, 

and each species has its own ecological and economic values. 

Mixed-species plantings are recommended together 

with large-leaf mangrove, Bruguiera gymnorrhiza. It is also

important to plant associated buffer areas, especially along 

the shoreline where mangroves grow better adjacent to 

banks stabilized by shoreline upland plants. Suitable upland 

tree genera include Calophyllum, Casuarina, Hibiscus, 

Thespesia, and Barringtonia. Within the mangrove forest, 

natural recruitment will often bring additional mangroves 

such as Bruguiera species, but these might be planted as 

well. Together they will complement and enhance the 

richness and stability of the planted environment. 

Estimated yield 

Snedaker and Brown (1982) state that mangrove forest products 

in Bangladesh have an annual value of US$36,000,000. 

Ong (1982) reports that the Matang Reserve in Malaysia 

with 35,000 ha (86,000 ac) of productive, almost monospecific, 

forests of R. apiculata, the value of mangrove wood 

alone is close to US$9,000,000 per year. 

The Matang Mangrove Forest Reserve in Malaysia has 

been managed for timber production of chiefly Rhizophora 

apiculata since the beginning of the century and is reputedly 

the best managed mangrove forest in the world (Khoon 

and Ong 1995, Ong et al. 1995). The average stand volume 

for these stilt mangroves was 153 m 3 /ha (2190 ft 3 /ac). Stand 

volumes up to 226 m 3 /ha (3230 ft 3 /ac) have been reported 

from Thailand. 

The present management plan for the Matang Reserve 

is a 30-year rotation period with two thinnings, at 15 and 

20 years. The 30-year rotation harvested yields have been 

around 136–299 mt/ha (61–133 t/ac) of green wood. However, 

there has been a decline in yield from 299 mt/ha (133 

t/ac) from virgin stands to the second-generation yields of 

158 mt/ha (70 t/ac) in 1967–69 to an even lower 136 mt/ha 

(61 t/ac) in 1970–77. Because the standing biomass of the 

trees did not increase from 23 years (155 mt/ha [69 t/ac]) 

to 28 years (153 mt/ha [68 t/ac]), it was suggested that a 

rotation of 25 years be used instead of the previous recommendation 

of 30 years. 

Markets 

Markets on most Pacific islands are local in nature, with 

little in the way of stilt mangrove products (other than indirect 

products like mangrove crabs and fruit bats) being 

exported from one island to another. The exception is probably 

firewood and charcoal, which is available from local 

markets in most places. In Southeast Asia, large quantities 

of stilt mangrove wood chips and charcoal may be moved 

greater distances and in greater volumes than wood products 

on smaller islands. 

INTERPLANTING/FARM 

APPLICATIONS 

Some interplanting systems include: 

Example 1—Bakhawan Eco Park (Primavera et al. 

2004) 

Location 

Aklan, New Buswang, and Kalibo, Philippines 

Description 

Planting was undertaken in 1989 and 1993. Species planted 

included R. apiculata and R. mucronata on a total area 

planted of 63 ha (156 ac). Sponsors and implementers were 

DENR, PACAP AusAID, USWAG, and KASAMA. 

Crop/tree interactions 

Not known. 

Spacing/density of species 

Spacing was about 1.5 m (5 ft) for both species. 

Example 2 (Primavera et al. 2004) 

Location 

Guimaras and Nueva Valencia, Philippines 

Description 

The planting was undertaken in 1994. Species planted included 

R. apiculata and R. mucronata on a total area of 

149 ha (368 ac). Funding source and implementers were 

DENR-CEP. 


Not known. 


Spacing was around 1.5 m (5 ft) for both species. 

Example 3 (Primavera et al. 2004) 

Location 

Iloilo and Carles, Philippines 

Description 

The planting was done in 2001. Species planted included 

R. apiculata and R. mucronata on a total area of 530 ha 

(1300 ac). Funding source and implementers were DENR 

Forestry Sector Program: MACABATA-ARM Federated 

People’s Organisation. 



Not known. 


Spacing was around 1.5 m (5 ft) for both species. 

PUBLIC ASSISTANCE 

Centre for Marine Studies, Marine Botany Group 

The University of Queensland 

St. Lucia QLD 4072 

Australia 

Web: . 

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0 Indo–West Pacific Rhizophora species

Species Profiles for Pacific Island Agroforestry (www.traditionaltree.org) 

Rhizophora apiculata, R. mucronata, R. stylosa, R. × annamalai, R. × lamarckii 

(Indo–West Pacific stilt mangroves) 

Authors: Norman C. Duke, Marine Botany Group, Centre for Marine Studies, The University of Queensland, St. Lucia QLD 4072, 

Australia; . 

Acknowledgments: The author and publisher thank Tom Cole, Farid Dahdouh-Guebas, Dale Evans, Ariel Lugo, and Diane Ragone 

for their input. Photo contributions by Beth Brook and Craig Elevitch are greatly appreciated. The author 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. Collaborators 

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. 2006. Rhizophora apiculata, R. mucronata, R. stylosa, R. × annamalai, R. × lamarckii (Indo–West 

Pacific stilt mangroves), ver. 2.1. In: Elevitch, C.R. (ed.). Species Profiles for Pacific Island Agroforestry. Permanent Agriculture 

Resources (PAR), Hōlualoa, Hawai‘i. . 

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; SPC/GTZ Pacific-German Regional Forestry Project; 

USDA Natural Resources Conservation Service (USDA NRCS); State of Hawai‘i Department of Land & Natural Resources Division 

of Forestry & Wildlife; and the USDA Forest Service Forest Lands Enhancement Program. This 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: . This institution is an equal opportunity provider. 

Reproduction: Copies of this publication can be downloaded from . This publication may be reproduced 

for noncommercial educational purposes only, with credit given to the source. © 2006 Permanent Agriculture Resources. All 

rights reserved.

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